Professional Documents
Culture Documents
CHRISTENSEN
EVANS OF
THE DOG
ANATOMY
OF
THE DOG
by
MALCOLM E. MILLER, D.V.M., M.S., Ph.D.
Late Professor and H ead o f the Department o f Anatomy,
New York State Veterinary College, Cornell University
12 Dyott Street
London WC1A 1DB
The majority of the illustrations used in this volume were prepared at the Department of
Anatomy at New York State Veterinary College at Cornell University and are used by permission
of and remain the property of Cornell University. © Assigned to Cornell University 1964.
© 1964 by W. B. Saunders Company. Copyright under the International Copyright Union. All
rights reserved. Made in the United States of America. Press of W. B. Saunders Company.
Library of Congress catalog card number 63—7038.
Print No.: 15 14 13 12
Foreword
T h i s t e x t is a monument to the memory of the author, Malcolm E. Miller, who labored faith
fully and long to produce it. During many of the years while it was being written the author
worked under the handicap of ill health. The struggle for health was a losing one that culmi
nated in his death in 1960 while he was in his fiftieth year.
From the beginning it was the author’s intent to produce a wholly original work. He was
not content to accept and use descriptions of others. His original goal, to which I think he ad
hered to the end, was to base each of his anatomical descriptions and illustrations on not less
than five original dissections. This work was meticulously done. It was slow, and it was inter
rupted by several hospitalizations and other periods when he was incapable of working. Alto
gether more than 15 years elapsed betw een the time the work was begun and when the task
had to be relinquished. By the time some of the later sections had been finished it was neces
sary to revise parts that had been finished years earlier.
Unfortunately time ran out on him before he was able to complete the manuscript and the
work had to be finished by two of his friends, former students and colleagues, who have done
it as a labor of love.
Since I am not an anatomist, I cannot adequately judge of the excellence of the work.
Since I saw the manuscript in the making, know of the devotion of the author to his specialty,
and know of the large amount of conscientious labor that went into it, I am led to believe that
the volume will be a fitting memory to “ M ac” Miller, an excellent and well-loved teacher, a
devoted veterinary anatomist, and a long-time colleague and friend.
The generous spirit of George C. Christensen and Howard E. Evans, the two friends who
assumed the task of completing the manuscript and preparing it for publication, deserves men
tion here. Without their efforts the volume could not have been published.
W ILLIAM A. HAGAN
Late Professor Emeritus and form er Dean,
New York State Veterinary College,
Cornell University, Ithaca, New York
Late Director, National Animal Disease Laboratory,
U. S. Department o f Agriculture, Ames, Iowa
Page v
M\l c o l m E M iller
B .S ., D .V .M ., M .S ., PH.D.
1909 - 1960
Page vii
D e d ic a t io n and P r efa ce
in the preparation, in 1947, of a “Guide to the Dissection of the Dog,” which is now in its
third edition.
The major portions of the manuscript and plates for this volume were near completion
at the time of Dr. Miller’s death. W e have endeavored to accept and edit the chapters that
had been previously solicited, and to make the necessary alterations and additions to
the manuscript while preserving the original style.
The terminology employed is based on the Nomina Anatomica (second edition, 1961
Excerpta Medica Foundation), with modifications suggested by the Nomenclatorial Com
missions of the World Association of Veterinary Anatomists and the American Association
of Veterinary Anatomists. Structures are generally designated by the anglicized forms in
common use. Each term, when introduced for the first time in the main discussion of the part,
is followed by its Latin equivalent. Eponyms and synonyms have been used occasionally for
reference to antecedent systems of nomenclature.
W e wish to thank Mrs. Mary Wells Miller for entrusting to us the completion and editing
of the unfinished manuscript. W e welcome this opportunity to show our gratitude to our for
mer colleague and good friend.
G eo rge C. C h r is t e n s e n
H ow ard E. E vans
Page viii
Acknowledgments
T h e c o m p l e t i o n of this volume at the present time would not have been possible without
the cooperation of the following contributing authors.
Most of the illustrations have been prepared from repeated dissections by Dr. Miller and
his students or colleagues at Cornell University and have not appeared previously. The great
est number of illustrations have been prepared by Miss Marion Newson, Medical Illustrator in
the Department of Anatomy at the New York State Veterinary College since 1951. Her skill as
an illustrator and her knowledge of anatomy have proved invaluable. Several illustrations for
the Skeletal and Muscular systems were drawn by Miss Pat Barrow at Cornell University; those
for the Introduction to the Nervous System and the Autonomic Nervous System were pre
pared by Algernon R. Allen and Neil Harris of Purdue University; and illustrations for the
Sense Organs and Skin were drawn by Robert R. Billiar, Dan J. Hillmann, and Santiago B.
Plurad of Iowa State University. Permission to use the illustrations which appeared in “Das
Lymphgefasssystem des Hundes” by Baum (1918) was kindly granted by Springer-Verlag.
The editors of the American Journal of Veterinary Research and American Journal of Anatomy
have also granted permission to use illustrations.
Page ix
A ckn o w led g m en ts
The Smith Kline & French Foundation of Philadelphia made a generous grant which
enabled us to double the number of colored illustrations originally intended to appear in this
volume.
Aid in securing pertinent literature and photocopied materials was cheerfully given by
Miss Mia Reinap and the staff of the New York State Veterinary College Library. Some trans
lations from the foreign literature were provided by Drs. Lisabeth Kraft, Karl Reinhard, and
Hermann Meyer. By generous permission of author and publisher parts of the text on the Mus
cular System has been freely translated and adapted from the Baum-Zietzschmann Anatomie
des Hundes (Paul Parey Verlag, Berlin).
Our relations with the publishers and their efficient staff have been most cordial. We
wish to acknowledge the helpful corrections and suggestions made by Miss Jean Husted while
working on the manuscript. Mr. John Dusseau, Vice President and Editor of the W. B.
Saunders Company, has continuously given his personal attention to all major and minor
problems that arose. His gracious advice and expeditious assistance are deeply appreciated.
G eorge C. C h r is t e n s e n
H ow ard E. E vans
Page x
Contents
Chapter 1
THE SKELETAL SYSTEM .......................................................................................................... 1
Chapter 2
ARTHROLOGY ................................................................................................................................ 95
Chapter 3
MYOLOGY ........................................................................................................................................ 131
Chapter 4
THE HEART AND A R T E R IE S ................................................................................................. 267
Chapter 5
THE VENOUS SYSTEM .............................................................................................................. 389
Chapter 6
THE LYMPHATIC SYSTEM ...................................................................................................... 430
Chapter 7
INTRODUCTION TO THE NERVOUS SYSTEM ............................................................... 464
By Ralph L. Kitchell
Chapter 8
THE BRAIN ...................................................................................................................................... 480
By Hermann Meyer
Chapter 9
THE SPINAL CORD AND M EN IN G ES.................................................................................... 533
By Robert C. McClure
Chapter 10
THE CRANIAL N E R V E S ............................................................................................................... 544
By Robert C. McClure
C o n ten ts
Chapter 11
THE SPINAL N E R V E S .................................................................................................................. 572
Chapter 12
THE AUTONOMIC NERVOUS SY STEM ................................................................................. 626
By M. W. Stromberg
Chapter 13
THE DIGESTIVE SYSTEM AND ABDOMEN......................................................................... 645
Chapter 14
THE RESPIRATORY SYSTEM ................................................................................................... 713
Chapter 15
THE UROGENITAL SYSTEM AND MAMMARY G LA N D S............................................... 741
By George C. Christensen
Chapter 16
THE ENDOCRINE SYSTEM ..................................................................................................... 807
By J. F. Smithcors
Chapter 17
THE SENSE ORGANS AND IN TEGUM EN T......................................................................... 837
The Eye, Orbit, and Adnexa.............................................................................................. 837
By Robert Getty
The Ear ................................................................................................................................. 847
By Robert Getty
The Nasal C avity.................................................................................................................. 863
By Robert Getty and Robert Hadek
The Organ of T a s te .............................................................................................................. 868
By John G. Bowne and Robert Getty
The Integument.................................................................................................................... 875
By James E. Lovell and Robert Getty
INDEX .........................................................................................................................................................889
x ii
CHAPTER 1
Long bones (ossa longa) occur only in the ex in the ligamentous tissue over which tendons
tremities, or limbs. The bones of the thigh and pass. They usually possess only one articular sur
arm, that is, the femur and humerus, are good face, which glides on a flat or convex surface of
examples. Typically a long bone, during its one or more of the long bones of the extremities.
growth, possesses a long middle part, the shaft Their chief function seems to be to alter the
or diaphysis, and two ends, the epiphyses. Dur course of tendons and to protect tendons at the
ing development each end is separated from the places where greatest friction is developed.
shaft by a plate of growing cartilage, the epi Flat bones (ossa plana) are found in the proxi
physeal cartilage (cartilago epiphysialis), or mal portions of the limbs, and in the head and
plate. At maturity the epiphyseal cartilage thorax. The most obvious function of these
ceases to grow, and the epiphysis fuses with the bones is protection. The ribs and the bones of
shaft as both share in the bony replacement of the cranium are primarily for this purpose. The
the epiphyseal cartilage. Fractures sometimes bones of the face are also flat, providing maxi
occur at the epiphyseal plate. Usually after ma mum shielding without undue weight, and
turity no distinguishable division exists between streamlining the head. Furthermore, the heads
epiphysis and diaphysis. The ends of most long of all quadrupeds overhang their centers of grav
bones enter into the formation of freely movable ity; a heavy head would be a handicap in loco
joints. Long bones form levers and possess great motion. The flat bones of the cranium consist of
tensile strength. They are capable of resisting outer and inner tables of compact bone and an
many times the stress to which they are nor intermediate uniting spongy bone, called diploe.
mally subjected. The stress on long bones is both In certain bones of the head the diploe is pro
through their long axes, as in standing, and at gressively invaded, during growth, by extensions
angles to these axes, as exemplified by the pull from the nasal cavity which displace the diploe
of muscles which attach to them. Although and cause a greater separation of the tables than
bones appear to be rigid and not easily influ would otherwise occur. The intraosseous air
enced by the soft tissues which surround them, spaces of the skull formed in this way are known
soft tissues actually do contour the bones. In as the paranasal sinuses. Bones which contain air
dentations in the form of grooves are produced cavities are called pneum atic hones (ossa pneu-
by blood vessels, nerves, tendons, and ligaments matica).
that lie adjacent to them, whereas roughened Irregular bones (ossa irregulata) are those of
elevations or depressions are produced by the the vertebral column, but the term also includes
attachments of tendons and ligaments. The ends all bones of the skull not of the flat type, and the
of all long bones are enlarged and smooth. In three parts of the hip bone (os coxae). Jutting
life, these smooth surfaces are covered by a layer processes are the characteristic features of ir
of hyaline cartilage, as they enter into the for regular bones. Most of these processes are for
mation of joints. The enlargement of each ex muscular and ligamentous attachments; some
tremity of a long bone serves a dual purpose. It are for articulation. The vertebrae of quad
diminishes the risk of dislocation and provides a rupeds protect the spinal cord and furnish a
large bearing surface for the articulation. The relatively incompressible bony column through
distal end of the terminal phalanx of each digit which the propelling force generated by the
is an exception to the stated rule. Since it is cov pelvic limbs is transmitted to the trunk. The
ered by horn and is not articular, it is neither en vertebrae also partly support and protect the
larged nor smooth. abdominal and thoracic viscera, and give rigid
Short bones (ossa brevis) are confined to the ity and shape to the body in general. The
carpal (wrist) and tarsal (ankle) regions, which amount of movement between any two verte
contain seven bones each. They vary in shape brae is small, but the combined movement per
from the typical cuboidal shape with six surfaces mitted in all the intervertebral articulations is
to irregularly compressed rods with only one flat, sufficient to allow considerable mobility of the
articular surface. In those bones having many whole body in any direction.
surfaces, at least one surface is nonarticular.
This surface provides an area where ligaments Development of Bone
may attach and blood vessels may enter and
leave the bone. Bone develops in both cartilage and mem
Sesamoid bones (ossa sesamoidea) are pres branous connective tissue. By far the greater
ent near freely moving joints. They are usually number of bones develop in cartilage, or, to
formed in tendons, but they may be developed speak more accurately, replace it. These bones
4 Chapter 1. T h e S k e l e t a l S y s te m
are known as endochondral, replacement, or various planes. Two types of bone structure will
cartilage bones. Some bones, such as those be seen. One is compact, or dense, bone, which
which form the roof of the cranium, develop in forms the outer shell of all skeletal parts. The
connective tissue sheets or membranes. Bones other is spongy, or cancellous, bone, which oc
developed in such a way are called m em brane, cupies the interior of the extremities of all long
or dermal, bones. bones and the entire interior of most other bones,
Bone is about one-third organic material, except certain of the skull bones and the bones of
wli i h is both intracellular and extracellular in the pectoral and pelvic girdles. Spongy bone is
location. Within or around the bone cells, not found in the girdles, where the two compact
known as osteoblasts, the bone matrix is laid plates are fused.
down. The osteoblasts later become the osteo- Compact bone (substantia compacta, or sub
cytes of mature bone. The cells which seem to stantia corticalis) is developed in direct ratio to
direct the deposition of cartilage and bone are the stress to which the bone is subjected. It is
derived from mesenchyme, which forms the thicker in the shafts of long bones than in their
greater part of the middle germ layer, or meso extremities. It attains its greatest uniform thick
derm, of the embryo. Since most bones are pre ness where the circumference of the bone is
formed in cartilage, the cartilage appears early least. The maximum thickness of the compact
in development, followed by perichondral and bone found in the femur and humerus of an
endochondral ossification. The first evidence of adult Great Dane was 3 mm. Local areas of in
ossification in an embryonic long bone is seen as creased thickness are present at places where
a collar around the middle of the shaft. Forma there is increased tension from muscles or liga
tion of the inorganic material is preceded by dis ments.
solution of the cartilage at the site of its deposi Spongy bone (substantia spongiosa) is elabo
tion. One stage follows another so rapidly that all rated in the extremities of long bones, forms the
calcified tissue soon takes the form of true bone internal substance of short and irregular bones,
of the spongy type. Secondary centers of ossifi and is interposed between the two compact lay
cation appear in the epiphyses. From this stage ers of most flat bones. Spongy bone consists of a
bone formation, much like the writing of a book, complicated maze of crossing and connecting
consists of altering or destroying the first-formed osseous leaves and spicules which vary in shape
material and the building of a more perfect and direction. The spongy bone of the skull is
structure. Osteoclasts are thought to be the cells known as diploe.
of bone destruction. Bones grow in length by The shafts of long bones in the adult are
endochondral ossification, but their increase in largely filled with yellow bone marrow (medulla
circumference, and the entire formation of cer ossium flava). This substance is chiefly fat. In the
tain bones of the skull, occurs through a differ fetus and the newborn, red bone marrow (me
ent process, described in the following para dulla ossium rubra) occupies this cavity and
graph. functions in forming red blood cells. No spongy
The bones of the face and dorsum of the cra bone is present in the middle of the shafts of
nium develop in sheets of connective tissue, not long bones, and the marrow-filled spaces thus
in cartilage. This type of bone formation is formed are known as medullary cavities (cava
known as intramembranous ossification. The os medullaria).
teoblasts and the osteoclasts continue to be the Spongy bone is developed where greatest
laborers in this activity. The compact bone stress occurs. The leaves or lamellae and bars are
formed by the periosteum is identical with mem arranged in planes where pressure and tension
brane bone in its elaboration. Bony tissue of are greatest, this structural development for
either type is capable of growing in any direc functional purposes being best seen in the proxi
tion. mal end of the femur. The interstices between
The larger sesamoid bones are preformed in the leaves and bars of spongy bone are occupied
cartilage, whereas the smaller ones may develop by red marrow. The spongy bone of ribs and
in membrane. vertebrae and of many other short and flat bones
is filled with red marrow throughout life. In the
Structure of Bone emaciated or the extremely aged, red marrow
gives way to fatty infiltration.
The gross structure of a dried, macerated The periosteum is an investing layer of con
bone is best revealed if the bone is sectioned in nective tissue which covers the nonarticular sur
G en eral 5
faces of all bones in the fresh state. The connec a specific bone by studying its eminences and
tive tissue covering of cartilage, known as depressions. There is a functional, embryologi-
perichondrium, does not differ histologically cal, or pathological reason for the existence of
from periosteum. Perichondrium covers only the every irregularity.
articular margins of articular cartilages, but in Most eminences serve for muscular and liga
vests cartilages in all other locations. Periosteum mentous attachments. Grooves and fossae in
blends imperceptibly with tendons and liga some instances serve a similar function. Facets
ments at their attachments. Muscles do not ac are small articular surfaces which may be flat,
tually have the fleshy attachment to bone which concave, or convex. Trochleas and condyles are
they are said to have, since a certain amount of usually large articular features of bone. The
connective tissue, periosteum, intervenes be roughened enlarged parts which lie proximal to
tween the two. At places where there are no the condyles on the humerus and femur are
tendinous or ligamentous attachments it is not known as epicondyles.
difficult, when bone is in the fresh state, to
scrape away the periosteum from it. Vessels and Nerves of Bone
The endosteum is similar in structure to peri
osteum, but is thinner. It lines the larger medul Bone, unlike cartilage, has both a nerve and
lary cavities, being the condensed peripheral a blood supply. Long bones and many flat and
layer of the bone marrow. Both periosteum and irregular bones have a conspicuous nutrient
endosteum, under emergency conditions, such (medullary) artery and vein passing through the
as occur in fracture of bone, provide cells (osteo compact substance to serve the marrow within.
blasts) which aid in repair of the injury. Some Such arteries pass through a nutrient foram en
times the broken part is over-repaired with bone (foramen nutricium) and canal (canalis nutric-
of poor quality. Such osseous bulges at the site of ius) of a bone and, upon reaching the marrow
injury are known as exostoses. cavity, divide into proximal and distal branches
Mucoperiosteum is the name given to the which repeatedly subdivide and supply the bone
covering of bones which participate in forming marrow and the adjacent cortical bone. In the
boundaries of the respiratory or digestive sys long and short bones terminal branches reach
tem. It lines all of the paranasal sinuses and con the epiphyseal plate of cartilage where, in
tains mucous cells. young animals, they end in capillaries. In adults
it is likely that many twigs nearest the epiphyses
Physical Properties of Bone anastomose with twigs arising from vessels in
Bone is about one-third organic and two- the periosteum. Nutrient veins pursue the re
thirds inorganic material. The inorganic matrix verse course. Not all of the blood supplied by the
of bone has a microcrystalline structure com nutrient artery is returned by the nutrient vein
posed principally of calcium phosphate. The or veins; much of it, after traversing the capillary
exact constitution of the crystal lattice is still bed, returns through veins which perforate the
under study, but it is generally agreed that bone compact bone adjacent to the articular surfaces
mineral is largely a hydroxyapatite 3 Ca3(P 0 4)2 at the extremities of these bones. The periosteal
•Ca(OH), with adsorbed carbonate. Some con arteries and veins are numerous but small; these
sider that it may exist as tricalcium phosphate arteries supply the extremities of long bones and
hydrate 3 Ca3(P 04)2 •2 H20 with adsorbed cal much of the compact bone also. They enter
cium carbonate (Dixon and Perkins 1956). The minute canals which lead in from the surface,
organic framework of bone can be preserved and ramify proximally and distally in the micro
while the inorganic part is dissolved. A 20 per scopic tubes which tunnel the compact and
cent aqueous solution of hydrochloric acid will spongy bone. The arterioles of the nutrient
decalcify any of the long bones of a dog in about artery anastomose with those of the periosteal
arteries deep within the compact bone. It is
one day. Such bones retain their shape but are
chiefly through enlargement of the periosteal
pliable. A slender bone, such as the fibula, can be
arteries and veins that an increased blood supply
tied into a knot after decalcification. The organic
and increased drainage are obtained at the site
material is essentially connective tissue, which
of a fracture. Veins within bone are devoid of
upon boiling yields gelatin.
valves, the capillaries are large, and the endo
thelium from the arterial to the venous side is
Surface Contour of Bone
continuous. Lymph vessels are present in the
Much can be learned about the role in life of periosteum as perivascular sheaths and probably
6 Chapter 1. T h e S k e l e t a l S y s te m
also as unaccompanied vessels within the bone body. In all lever movements of bones by mus
marrow. The nerves of bone are principally sen cles the force or the fulcrum is always at one
sory. They serve as an inner defense against in end and the weight at the other. The weight is
jury. The sensory nerves of the skin form the never between the force and the fulcrum, which
outer defense. Both carry impulses which result is necessary for a second class lever. Nearly all
in pain. Kuntz and Richins (1945) state that both muscles act at a mechanical disadvantage. The
the afferent and sympathetic fibers probably speed at which the weight travels is in direct
play a role in reflex vasomotor responses in the proportion to the shortness of the force arm, and
bone marrow. this is determined by the distance of the inser
tion of the muscle from the joint, or fulcrum.
Function of Bone 3. Bone serves as a storehouse for calcium
The skeleton of the vertebrate body serves and phosphorus and for many other elements in
four functions. small amounts. These substances are withdrawn
1. Bone forms the supporting and in many in from the bone as complicated compounds. The
stances the protecting framework of the body. greatest drain occurs during pregnancy; con
The supportive function does not need explana versely, the greatest deposition takes place dur
tion. The essential organs of vertebrates receive ing growth. In the large breeds, such as the
protection from the skeleton. These are the Great Dane and St. Bernard, the skeleton is the
brain and spinal cord, heart, and liver. To these system most likely to show the effects of a nutri
may be added certain pelvic organs which, al tional deficiency. Undermineralization of the
though not essential for life, are protected by the skeleton is a common manifestation of under
pelvis. (The urinary bladder is largely an abdom feeding, improper feeding, or inability of the in
inal organ in the dog.) The lungs further protect dividual to assimilate food adequately.
the heart, and are in turn protected by the ribs. 4. Bone serves as a factory for red blood cells
2. Many bones serve as levers by which the and for several kinds of white blood cells. In the
muscles move the body. Of the three types of normal adult it also stores fat. Red marrow,
levers, only two are represented by bones. where the red and many white blood cells de
Many bones may serve as either a first or a velop, occurs most richly in the bones of the
third class lever, owing to the action of differ axial skeleton and in the proximal epiphyses of
ent muscles at different times and to changes the humerus and femur; yellow or fatty marrow
in the positions of force and fulcrum. No lever is most abundant in the long bones of the ex
of the second class is represented in the living tremities.
A XIA L SK ELET O N
BRACHY M E S A T I D O L IC H O
M EASUREM ENT
C E P H A L IC C E P H A L IC C E P H A L IC
(1962) briefly reviews the phylogenetic history of size than does any other part of the skeleton. In
the vertebrate skull. Skulls differ more in size brachycephalic breeds the facial skeleton is
and shape among domestic dogs than in any shortened and broadened. In some brachyceph
other mammalian species. For this reason, cra alic breeds, the English bulldog, for example, the
niometry in dogs takes on added significance. lower jaw protrudes anterior to the upper jaw,
Certain points and landmarks on the skull are producing the undershot condition known as
recognized in making linear measurements and prognathism o f the mandible. Most other breed
have been used by Stockard (1941) and others. types have brachygnathic mandibles, that is, re
The more important of these are: ceding lower jaws. Although brachygnathism of
Inion: Central surface point on the external the mandibles is relative, both the collie and the
occipital protuberance. dachshund frequently exemplify this condition
Bregma: Junction on the median plane of the to a marked extent.
right and left frontoparietal sutures, or the point Table 2 shows average measurements in milli
of crossing of the coronal and sagittal sutures. meters taken from randomly selected adult-
Nasion: Junction on the median plane of the skulls of the three basic types. From these data
right and left nasofrontal sutures. it can be seen that the greatest variation in skull
Prosthion: Anterior end of the intermaxillary shape occurs in the facial part. In making com
suture, located between the roots of the upper parisons of skull measurements it is essential that
central incisor teeth. the over-all size of the individuals measured is
Pogonion: Most anterior part of the mandi taken into consideration. As a rule the dolicho
ble, at the symphysis, located between the roots cephalic breeds are larger than the brachyce
of the lower central incisor teeth. phalic, whereas the working breeds fall in the
Basion: Middle of the ventral margin of the mesaticephalic group, and these as a division
foramen magnum. have the greatest body size. The only measure
The center o f the external acoustic meatus: ment in which the brachycephalic type exceeds
Although unnamed, this spot also serves as a ref the others, in the small sampling shown, is facial
erence point. width. To obviate the size factor among the
Three terms are frequently used to designate breed types, indices are computed. These indi
head shapes: cate relative size and are expressed by a single
Dolichocephalic, meaning long, narrow term representing a two-dimensional relation
headed. Breed examples: collie, Russian wolf ship. The cranial index is computed by multiply
hound. ing the cranial width by 100 and dividing the
Mesaticephalic, meaning a head of medium product by the cranial length. Skull and facial
proportions. Breed examples: German shepherd, indices are computed in the same manner.
beagle, setter. Differences among the breeds in facial skele
Brachycephalic, meaning short, wide-headed. tal development are the most salient features re
Breed examples: Boston terrier, Pekingese. vealed by craniometry. The face is not only
The face of the dog varies more in shape and short in the brachycephalic breeds but it is also
T h e Sk u l l 9
Incisi ve
M axilla- -
Z y g om atic-■
Pp e s p h e n o i d - -
P i e r i j 2 oi d
P a n i e i a I ----------
B a s i s p hen o l d
Temponal'
O ccipital' '
actually wider than it is in the heavier, longer- from a flat surface. Disproportionate growth of
headed breeds. These data do not show that ap the length of the face occurs after the early
preciable asymmetry exists, especially in the weeks of life, so that suckling the dam is not im
round-headed types. Even though the neurocra paired. Dental malocclusion is treated under the
nium varies least in size, it frequently develops description of the teeth in Chapter 12 on the
asymmetrically. The caudal part of the skull is Digestive System.
particularly prone to show uneven development. Cranial capacity varies but little among the
The further a breed digresses from the ancestral different breeds and skull types. The terms mi-
German shepherd type, the more likely are dis crocephalic, m esocephalic, and m egacephalic
tortions to be found. This is particularly true of indicate skulls with small, medium, and large
the round-headed breeds, as these types have cranial capacity, respectively. The following
been developed to please man’s fancy without data were computed by filling the crania with
regard to the health of the strain or even to their mustard seed after the foramina had been closed
expected survival without special attention. with modeling clay, and then determining the
Their is little rationale in developing a breed of volume of seed used. Average Boston terrier
dog like the Boston terrier, with large, round skulls held 82 cc. A sampling of skulls of medium
heads and small pelves. In this instance the size and medium length showed an average ca
transgression against nature is twofold. The pacity of 92 cc.; the average skull capacity of the
large crania of the young frequently exceed the crania of the Russian wolfhound and of the collie
dimensions of the dam’s pelvis, and normal par was 104 cc.
turition is impossible. The breed would soon be The names of the individual bones making up
extinct if cesarean sections were not performed. the 50 which compose the skull are listed in
The ugly appearance of the English bulldog is Table 3.
partly produced by the prognathic condition of
the lower jaw, as well as the brachygnathic con
dition of the upper jaw. This structural dishar B on es o f t h e B r a in c a s e
mony results in poor occlusion of the teeth.
Stockard (1941) found, by crossing purebred Occipital Bone
breeds of extreme jaw sizes, that the lengths of
the upper and of the lower jaw are inherited in The occipital bone (os occipitale) (Figs. 1-5,
dependently. Dogs with prognathic upper jaws 1-6) forms the posterior portion of the skull. It
and brachygnathic lower jaws are unable to eat develops from four centers: a squamous part
Unpaired: 1. Vomer
Unpaired: 1. Basihyoid
T h e Sk u l l 11
- I n t e r p a r i e t a l process
D o r s a l nuchal l i n e - £*/■. o c c i p i t a l p r o t u b e r a n c e
Ventral nuchal l i n e - - -|J! A _ _ Exf. o c c i p i t a l c r e s t
Foramen i m p a r
Int. o c c i p i t a l p r o t u b e r a n c e v
sagittal crt
Transverse sulcus
V e r mi f o r m i mpressi on
Int. o c c i p i t a l c r e s t ------
L o c at i o n of s u p r a m a s t o i d f o r a m e n - - ^ — Nuchal tu b e rc le
A n t e r i o r v p o s t er i o r o p e n i n g s of
~ condyloid ca n a l
Ventral o p e n i n g of c o n d y l o i d c a n a l
S u lc u s of ventral p e tr o s a l sinus
' Int. o p e n i n g of h y p o g l o s s a l c a n a l
dorsally, two lateral condylar parts, and a basilar cerebellar surface of the squamous part of the
part ventrally. occipital bone which houses a part of the vermis
The squamous part (pars squamosa), or supra- of the cerebellum. The vermiform impression is
occipital bone, is the largest division. Dorsoan- bounded laterally by the paired internal occipi
teriorly it is wedged between the parietal bones tal crest (crista occipitalis interna), which is usu
to form the interparietal process (processus in- ally asymmetrical and convex laterally. Lateral
terparietalis). This process represents the un to the internal occipital crest, as well as on the
paired interparietal bone which fuses prenatally ventral surface of the interparietal process, there
with the supraoccipital. From the interparietal are elevations, juga cerebralia et cerebellaria,
process arises the mid-dorsal external sagittal and depressions, impressiones digitatae. Ven
crest (crista sagittalis externa), which, in some trally the squamous part is notched to form the
specimens, is confined to this bone. The anterior dorsal part of the foramen magnum. On either
end of the interparietal process is narrower and side the supraoccipital is fused with the paired
thinner than the caudal part, which turns ven exoccipital. This union represents the former ar
trally to form a part of the posterior surface of ticulation (synchondrosis intraoccipitalis squa-
the skull. The dorsal nuchal line (linea nuchalis molateralis) which extended from the foramen
dorsalis) marks the division between the dorsal magnum to the temporal bone.
and posterior surfaces of the skull. It is an un The lateral parts (partes laterales), or exoccip
paired sharp-edged crest of bone which reaches ital bones, bear the occipital condyles (condyli
its most dorsal point at the external occipital occipitales), which are convex and, with the at
protuberance. On each side it arches ventrally las, form the atlanto-occipital joints. The jugular
before ending on a small eminence located dor- process (processus jugularis) is located, one on
soposterior to the external acoustic meatus. The either side, lateral to the condyle, and ends in a
ventral nuchal line (linea nuchalis ventralis) is a rounded knob ventrally, usually on a level with
line located in a frontal plane, ventral to the the bottom of the anteriorly located tympanic
middle part of the dorsal nuchal line, and forms bulla. Between the jugular process and the oc
the base of an uneven triangular area. It extends cipital condyle is the ventral condyloid fossa
transversely between the dorsolateral parts of (fossa condylaris ventralis). On a ridge of bone
the dorsal nuchal line. It is not distinct. The ex anterior to this fossa is the hypoglossal foram en
ternal occipital protuberance (protuberantia oc (foramen hypoglossi), which is the external
cipitalis externa) is the median, triangular pro opening of the hypoglossal canal (canalis hypo-
jection forming the most dorsoposterior portion glossi), a direct passage through the ventral part
of the skull. The external occipital crest (crista of the occipital bone. The dorsal condyloid fossa
occipitalis externa) is a smooth median ridge ex (fossa condylaris dorsalis) is located dorsal to the
tending from the external occipital protuber occipital condyle. The rather large condyloid
ance to the foramen magnum. It is poorly devel canal (canalis condylaris) runs through the me
oped in some specimens. dial part of the lateral occipital bone. There is an
Within the dorsal part of the occipital bone intra-osseous passage between the condyloid
and opening bilaterally on the cerebral surface canal and the hypoglossal canal. Usually there is
is the transverse can al (canalis transversa), also a small passage between the condyloid canal
which, in life, contains the venous transverse and the petrobasilar fissure.
sinus. The transverse canal is continued later The basilar part (pars basilaris), or basioccipi-
ally, on each side, by the sulcus fo r the trans tal bone, is unpaired, and forms the posterior
verse sinus (sulcus sinus transversi). Mid-dor- third of the cranial base. It is roughly rectangu
sally or to one side, the sagittal sinus enters the lar, although caudally it tapers to a narrow, con
transverse sinus via the foram en impar. Between cave end which forms the central portion of the
the laterally located sulci the skull protrudes an- inter condyloid notch (incisura intercondyloidea).
teroventrally to form the internal occipital pro The adjacent occipital condyles on each side
tuberance (protuberantia occipitalis internus). deepen the incisure as they contribute to its for
Extending anteriorly from the internal occipital mation. The incisure bounds the ventral part of
protuberance is the variably developed, usually the foramen magnum. The foram en magnum is
paramedian and always small internal sagittal a large, transversely oval opening in the postero-
crest (crista sagittalis interna). The vermiform ventral portion of the skull, through which pass
impression (impressio vermis), forming the thin the spinal cord and its associated structures, the
nest part of the caudal wall of the skull, is an ir meninges, vertebral venous sinuses, the spinal
regular excavation of the median portion on the portion of the accessory nerve, and the various
T h e Sk u l l 13
arteries associated with the spinal cord. In between the petrosal and the occipital bones
brachycephalic breeds it is more circular than which forms the petro-occipital suture is the
oval, and it is frequently asymmetrical. The dor synchondrosis petro-occipitalis. Laterally, and
sal boundary of the foramen magnum is featured proceeding dorsally, the occipital bone first ar
by the caudally flared ventral part of the supra ticulates with the squamous temporal bone su
occipital bone. The caudal extension is increased perficially, the occipitosquamous suture (sutura
by the paired nuchal tubercles (tubercula nu- occipitosquamosa), and with the mastoid part
chalia). These projections are sufficiently promi of the petrous temporal bone deeply, the occip
nent to make spinal punctures at this site diffi itom astoid suture (sutura occipitomastoidea);
cult. The dorsal surface of the basioccipital bone further dorsally it articulates with the parietal
is concave to form the sulcus medulla oblongata. bone, the lam bdoid suture (sutura lambdoidea).
The lateral surfaces of the caudal half of the Where the squamous and lateral parts of the oc
basioccipital bone fuse with the exoccipital cipital bone articulate with each other and with
bones along the former ventral intraoccipital the mastoid part of the temporal bone, the su-
synchondrosis (synchondrosis intraoccipitalis pram astoid foram en (foramen supramastoi-
basilateralis). The ventral surface of the basioc deum) is formed.
cipital bone adjacent to the petrotympanic syn Variations in the occipital bone are numer
chondrosis possesses muscular tubercles (tuber ous. The foramen magnum varies in shape and
cula muscularia). These are rough, sagittally is not always bilaterally symmetrical. The con
elongated areas, located medial to the smooth, dyloid canal may be absent on one or both sides.
rounded tympanic bullae. The pharyngeal tu Even when both canals are present, connections
bercle (tuberculum pharyngeum) is a single tri between the hypoglossal and condyloid canals
angular rough area anterior to the intercondy- may fail to develop. The jugular processes may
loid incisure. Laterally the basioccipital bone is extend several millimeters ventral to the tym
grooved to form the ventral petrosal sulcus, panic bullae so that they will support a skull
which concurs with the pyramid of the temporal without the mandibles when it is placed on a
bone to form the petrobasilar canal (canalis horizontal surface; conversely, they may be
petrobasilaris). short, retaining the embryonic condition. The
Ventrally the anterior end of the basioccipital vermiform impression may be deep, causing a
bone articulates with the body of the basisphe- posteromedian rounded, thin protuberance on
noid bone at the cartilaginous spheno-occipital the posterior face of the skull. The foramen im-
joint (synchondrosis spheno-occipitalis). Ventro- par may be double. It is rarely median in posi
laterally the occipital bone articulates with the tion. A sutural bone may be present at the an
tympanic part of the temporal bone to form the terior end of the interparietal process.
cartilaginous tympano-occipital joint (synchon
drosis tympano-occipitalis). Deep to this joint is Parietal Bone
the important petro-occipital suture (sutura
petro-occipitalis), in which the foramen lacerum The parietal bone (os parietalis) (Fig. 1-7) is
caudalis, or jugular foramen, opens. The joint paired and forms most of the dorsolateral part
- 1 n f e r p a r i e t a l s u t ur e
Tentorium o s s e u m - -
T r a n s v e r s e sul cus —
V a s c u l a r gr oo ve f o r m e d . m e n i n g e a l a. -
|»|,NfwsoN
of the cranial wall. It articulates dorsally with parietal or coronal suture (sutura frontoparie-
its fellow and with the interparietal process of talis). The posterior or occipital border (margo
the occipital bone. Each parietal bone lies di occipitalis) meets the occipital bone to form the
rectly anterior to the squamous occipital and occipitoparietal suture (sutura occipitoparie-
dorsal to the squamous temporal. In the new talis). The anterior half of the dorsal or sagittal
born no elevation is present at the sagittal in border (margo sagittalis) articulates with its
terparietal suture or on the interparietal proc fellow on the midline to form the sagittal suture
ess, but soon thereafter in the heavily muscled (sutura sagittalis). The posterior half of the dorsal
breeds, particularly in the male, the mid-dorsal border articulates with the interparietal process
external sagittal crest is developed. This crest, of the occipital bone to form the parietointer-
which increases in size with age, forms the me parietal suture (sutura parietointerparietalis).
dial boundary of the temporal fossa (fossa tem The ventral or squamous border (margo squamo-
poralis), a large area on the external surface sus) is overlaid by the squamous temporal bone in
(facies externa) of the cranium from which the forming the squamous suture (sutura squamosa).
temporal muscle originates. In dolichocephalic A small area of the squamous border at its an
breeds with heavy temporal muscles, the exter terior end articulates with the temporal wing of
nal sagittal crest may reach a height of more the sphenoid bone to form the parietosphenoi-
than 1 cm. and extend from the external occipi dal suture (sutura parietosphenoidalis). Over
tal protuberance to the parietofrontal suture. lapping of the bones at the squamous and coro
Anteriorly, it continues as the diverging frontal nal sutures allows for cranial compression of the
crests. In most brachycephalic skulls the exter fetal skull during its passage through the pelvic
nal sagittal crest is confined to the interparietal canal.
part of the occipital bone and is continued an
teriorly as the diverging tem poral lines (lineae Frontal Bone
temporales). The temporal lines at first are con The frontal bone (os frontale) (Figs. 1-8, 1-9)
vex laterally, then become concave as they cross is irregular in shape, being broad posteriorly and
the parietofrontal, or coronal, suture and are somewhat narrower anteriorly. Laterally, the
continued as the external frontal crests to the anterior part is concave and forms the medial
zygomatic processes. The temporal lines replace wall of the orbit. Posterior to this concavity, it
the external sagittal crest in forming the medial flares laterally to form part of the temporal fossa.
boundaries of the temporal fossae in most The frontal sinus (sinus frontalis) is an air cavity
brachycephalic skulls. located between the inner and outer tables of
The internal surface (facies interna) of the the anterior end of the frontal bone and is di
parietal bone presents digital impressions and vided into two or three compartments. It is dis
intermediate ridges corresponding, respectively, cussed in greater detail under the heading Para
with the cerebral gyri and sulci. A well-defined nasal Sinuses.
vascular groove, the sulcus fo r the middle me For descriptive purposes the frontal bone is
ningeal artery (sulcus arteriae meningeae me divided into orbital, temporal, frontal, and nasal
diae), starts at the ventrocaudal angle of the parts.
bone and arborizes over its internal surface. The The orbital part (pars orbitalis) is a segment of
groove runs toward the opposite angle of the a cone with the apex located at the optic canal
bone, giving off smaller branched grooves along and the base forming the medial border of the
its course. A leaf of bone projects anteromedially orbital margin (margo orbitalis). Lateral to the
from the dorsal part of the posterior border. This most dorsal part of the frontomaxillary suture
leaf concurs with its fellow and with the internal (sutura frontomaxillaris) the orbital margin is
occipital protuberance to form the curved ten slightly flattened for the passage of the vena an-
torium ossium. On the internal surface of the gularis oculi. Ventrally, a long, distinct, dorsally
parietal bone near its caudal border is a portion arched muscular line marks the approximate
of the transverse sulcus, which leads dorsally ventral boundary of the bone. The ethm oidal
into the transverse canal of the occipital bone foram in a (foramina ethmoidalia) are two small
and ventrally into the temporal meatus. openings about 1 cm. anterior to the optic canal.
The borders of the parietal bone are anterior, The smaller opening is in the frontosphenoidal
posterior, dorsal, and ventral in position, since suture; the larger foramen, located dorsopos-
the bone is essentially a curved, square plate. terior to the smaller, parses obliquely through
The anterior or fron tal border (margo frontalis) the orbital part of the frontal bone. Sometimes
overlaps the frontal bone, forming the fronto the two ethmoidal foramina are confluent. At
T h e Sk u l l
Septum o f fr o n ta l sinus
N ^ Maxillary incisure
E th m o id a l incisure
' ■Ma x i l l a r y p r o c e s s
To f r o n t a l s i n u s \Ethmoida! foramina
F ig . 1-8. Left frontal bone, medial aspect.
-P A R S FRONTALIS
G r o o v e f o r a n g u l a r i s o c u l i v.
PAR S N A S A L I S -PARS T E M P O R A L I S
xExt; f r o n t a l c r e s t
Fossa f o r l a c r i m a l g l a n d
Zygomatic p ro c e s s
Frontal foramen
the orbital margin, the frontal and orbital sur teriorly and a small portion of the nasal cavity
faces meet, forming an acute angle. The supra anteriorly. The salient ethmoidal notch sepa
orbital or zygomatic process (processus zygoma- rates the two parts. The posterior part is deeply
ticus) is formed where the orbital margin meets concave and divided into many fossae by the
the external fron tal crest (crista frontalis ex digital impressions and the cerebral juga. Fine,
terna), which curves anterolaterally from the dorsocaudally running vascular grooves indi
temporal line or sagittal crest. On the orbital cate the position occupied in life by the an
surface of the zygomatic process is a small fora terior meningeal vessels. The large aperture for
men which is only large enough to admit a horse the fron tal sinus is located dorsal to the eth
hair. Ventroanterior to this foramen in some moidal notch. The nasal part of the internal
adult skulls the fo ssa fo r the sm all lacrim al surface of the frontal bone is marked by many
gland (fossa glandulae lacrimalis) can be seen. longitudinal lines of attachment for the eth-
The temporal part (pars temporalis) forms moturbinates.
that part of the frontal bone posterior to the or The mid-dorsal articulation of the frontal
bital part. Dorsally the two tables of the frontal bones forms the frontal suture (sutura inter-
bone are separated to form the frontal sinus, frontalis). This suture is a forward continuation
whereas ventrally and posteriorly the two tables of the sagittal suture between the parietal bones.
are fused or united by a small amount of diploe Posteriorly the frontal bone is overlapped by the
to form the braincase. parietal bone, forming the frontoparietal suture
The frontal part (pars frontalis), or frontal (sutura frontoparietalis). Ventrally the rather firm
squama (squama frontalis), is roughly triangular, sphenofrontal suture (sutura sphenofrontalis) is
with its base facing medially, and articulating formed. Anteriorly the frontal bone articulates
with that of the opposite bone. It is gently with the nasal, maxillary, and lacrimal bones to
rounded externally and is largely subcutaneous form the frontonasal suture (sutura fronto-
in life. Its posterior boundary is the external nasalis), the frontomaxillary suture (sutura
frontal crest and the lateral part of its anterior frontomaxillaris), and the frontolacrimal suture
boundary is the orbital margin. (sutura frontolacrimalis). Deep in the orbit, the
The nasal part (pars nasalis) is the anterior ex frontal bone articulates with the palatine bone to
tension of the frontal bone. Its sharp, pointed form the frontopalatine suture (sutura fronto-
nasal process (processus nasalis) lies partly under palatina). Medially, hidden from external view,
and partly between the posterior parts of the the frontal bone articulates with the ethmoid
nasal and maxillary bones. The septum o f the bone in forming the frontoethm oidal suture
fron tal sinus (septum sinuum frontalium) is a (sutura frontoethmoidalis).
vertical median partition which closely articu
lates with its fellow in separating right and left Sphenoid Bone
frontal sinuses. It is widest near its middle,
which is opposite the cribriform plate. Anteri The sphenoid bone (os sphenoidale) (Figs.
orly it is continuous with the septal process of 1-10, 1-11, 1-12) forms the anterior two-thirds
the nasal bone. The ventral part of the septum of the base of the neurocranium, between the
of the frontal sinus is the internal fron tal crest basioccipital posteriorly and the ethmoid an
(crista frontalis interna). The conjoined right teriorly. It consists of two parts, each possess
and left crests articulate with the perpendicular ing a pair of wings and a median body. The
plate of the ethmoid bone ventrally and with the anterior part is the presphenoid (os presphe-
conjoined right and left septal processes of the noidale); the posterior part, with the larger
nasal bones anteriorly. The sagittally located wings, is the basisphenoid (os basisphenoidale),
notch between the pointed anterior end of the or postsphenoid.
septum and the nasal process is the maxillary The dorsal part of the body of the presphe
incisure (incisura maxillaris). The ethm oid in noid is roofed over by the fusion of right and
cisure (incisura ethmoidalis), which lies dorsal left orbital wings (alae orbitales) to form the
and lateral to the cribriform plate of the eth yoke (jugum sphenoidale). The yoke forms the
moid bone, is formed by the smooth concave base of the anterior cranial fossa. A small,
edge of the internal table of the nasal part of median tubercle, the rostrum (rostrum sphenoi
the frontal bone. dale), divided in the newborn, projects from
The internal su rface (facies interna) of the the anterior border of the yoke. Posteriorly, the
frontal bone forms a part of the brain case pos yoke forms a shelf, the orbitosphenoidal crest
T he Sk u l l 17
Sphenoidal s i n u s .
Jugum sphenoidale
O rbital w ing_ __
- OrbitosphenoidaI c r e s t
Optic c o nal — 4 j/p>
A nterior clinoid process- - - ^~ - S u l c u s c h i a s m a t i s
T u b e rc u lu m sellae A n t e r i o r end o f p t e r y g o i d p r o c e s s
^ M - - Notch f o r o r b i t a l fi ss ur e
Po s t e r i or cl i n o i d p r o c e s s
- -Foramen rotundum
Gr oove foi------r - t
, . , UlVr" „j'1--
med- m e n i n g e a l a. ' - Foramen ovale
/
L i n g u l a sphenoida I i s / / ■Hypophyseal f o s s a
Orbital wing
Temporal winy
O rb ita l fis s u re \
„ - O p t ic canal
F o r a m e n f o r z y y o m a t i c n.
A n t e r i o r o p e n i n g of
p te ry g o id ca n a l-
''Sphenoidal sinus
Po s t e r i or a l a r f o r a m e n Body of p re s p h e n o id
A nterior a la r f o r a m e n 1 NP t e r y y o i d p r o c e s s e s
(crista orbitosphenoidalis), under which lie the These large openings are at a lower level and
diverging optic canals (canales opticae). On are located slightly posterolateral to the optic
either side the anterior clinoid process (pro canals.
cessus clinoideus anterior) projects posteriorly The temporal or great wings (alae temporales
from the orbitosphenoidal crest and overhangs s. majores), or alisphenoids, of the basisphe
the orbital fissure. On the dorsum of the body, noid are larger than the orbital wings, and
posterior to the optic canals, is the unpaired curve outward and upward. Anteriorly they
sulcus chiasm atis, in which lies the optic extend to the lateral margin of each frontal
chiasma. The body of the basisphenoid forms bone to form the sphenofrontal suture. The
the base of the middle cranial fossa. The mid posterior two-thirds of the temporal wings are
dle of its dorsal surface is slightly dished to covered laterally by the squamous temporal
form the oval h yp op h y seal fo s s a (fossa hypo- bone in forming the sphenotemporal suture. At
physeos). The fossa is limited anteriorly by the the base of each wing, near its junction with
tuberculum sellae, an upward sloping ridge of the body, are a series of foramina. The oval
bone formed at the junction of the presphenoid foram en (foramen ovale) is a large opening
and basisphenoid. The hypophyseal fossa is which leads directly through the cranial wall.
limited posteriorly by a bony process, the It is located about 0.5 cm. medial to the tem
dorsum sellae, which, in adult skulls, is flat poromandibular joint. A small notch or even a
tened and expanded at its free end. Projecting foramen, foram en spinosum, may be present in
anteriorly on either side of the dorsum sellae is its posterolateral border for the transmission of
a posterior clinoid process (processus clinoideus the middle meningeal artery. The alar canal
posterior). This complex of bony structures, (canalis alaris) runs through the anterior part of
consisting of the tuberculum sellae and anterior the base of the temporal wing. Its smaller pos
clinoid processes, the hypophyseal fossa, and the terior opening is the posterior alar foram en
dorsum sellae with its two posterior clinoid proc (foramen alare posterius), and its larger an
esses, is called the sella turcica, or Turkish sad terior one is the anterior alar foram en (foramen
dle. In life it contains the hypophysis. Occasion alare anterius). Entering the canal from the
ally the small cran iopharyn geal can al (canalis cranium is the round foram en (foramen ro-
craniopharyngeus) persists in the adult. This tundum). It can be seen by viewing the medial
canal is a remnant of the pharyngeal diverticu wall of the alar canal through the anterior alar
lum to the hypophyseal fossa from which the foramen. Dorsoanterior to the alar canal is the
pars glandularis of the hypophysis develops. orbital fissure. A small foram en alare parvum
The orbital or lesser wings (alae orbitales s. may be present as the dorsal opening of a small
minores), or orbitosphenoids, leave each side of canal which leaves the alar canal. It is located
the presphenoid and roof across its body. An on the ridge of bone separating the orbital fis
teriorly, at the junction of the wings and the sure from the anterior alar foramen. When
body, the presphenoid is hollow and divided by present it conducts the zygomatic nerve from
a longitudinal septum to form the sphen oidal the maxillary trunk. Two pairs of grooves are
fo ssa e (fossae sphenoidales) into which extend present on the basisphenoid bone. The ex
the ventrocaudal parts of the ethmoturbinates. tremely small pterygoid groove (sulcus nervi
The orbital wings articulate ventrally with the pterygoidei) leads into the minute pterygoid
palatine and dorsally with the frontal bones. In canal (canalis pterygoideus). It begins anterior
the frontosphenoidal suture is located the eth to the small, pointed, muscular process of the
m oidal foram en (foramen ethmoidale); a sec temporal bone where it is located in the suture
ond larger ethmoidal foramen is usually present between the pterygoid and basisphenoid bones.
in the frontal bone dorsoposterior to the one in It ends in the posterior part of the pterygo
the suture. These foramina may be confluent. palatine fossa. Probing with a horse hair will
The posterior parts of the orbital wings slope reveal that it runs medial to the pterygoid proc
upward and outward and are thicker, but ess of the sphenoid in the suture between this
smaller, than the anterior parts. Their bases process and the pterygoid bone. The second
are perforated by the optic canals. Medially, in groove of the basisphenoid is the sulcus fo r the
young specimens, the two elliptical optic canals middle m eningeal artery (sulcus arteriae men-
are confluent across the midline. The orbital ingeae mediae). This groove runs obliquely dor-
fissures (fissurae orbitales) are located lateral to solaterally from the oval foramen on the cerebral
the body of the sphenoid in the sutures be surface of the temporal wing and continues
tween the orbital wings and the temporal wings. mainly on the temporal and parietal bones. Two
T h e Sk u l l 19
notches indent the posterior border of the tem pyramid houses the cochlea and the semicircular
poral wing. The medial notch (incisura carotica) canals, and is the last to fuse with the other parts
concurs with the temporal bone to form the ex in development. It is located completely within
ternal carotidtforamen (foramen caroticum ex the skull. The pars mastoidea is the only part of
ternum). The lateral notch, with its counterpart the temporal bone to form a part of the posterior
on the temporal bone, forms the short osseous surface of the skull. It is located between the
auditory tube (tuba auditiva ossea). A low ridge ventral end of the dorsal nuchal line laterally
of bone, the lingula (lingula sphenoidalis), end and the jugular process ventromedially. The
ing in a process, separates the two openings. tympanic part, or bulla tympanica, is a sac-
The pterygoid processes (processus ptery- shaped protuberance, roughly as large as the
goidei) are the only ventral projections of the end of one’s finger, which lies ventral to the
basisphenoid. They are thin, sagittal plates about mastoid part and is in opposition to the jugular
1 cm. wide, 1 cm. long, and a little over 1 cm. process posteriorly. The squamous part consists
apart. Attached to their medial surfaces are the of two basic divisions, an expanded plate which
posteriorly hooked, approximately square ptery lies above the bulla, and the anteriorly project
goid bones. The processes and pterygoid bones ing zygomatic process which forms the posterior
separate the posterior parts of the pterygopal half of the zygomatic arch.
atine fossae from the nasal pharynx. The petrosal part (pars petrosa), pyramid, or
The body of the basisphenoid articulates pos petrosum (Fig. 1-14), is fused around its pe
teriorly with the basioccipital, forming the riphery laterally to the medial surfaces of the
spheno-occipital synchondrosis (synchondrosis tympanic and squamous parts; posteriorly, it may
spheno-occipitalis); and anteriorly with the pre on rare occasions be united with the mastoid
sphenoid, forming the intersphenoidal syn part. It is roughly pyramidal in shape, and is
chondrosis (synchondrosis intersphenoidalis). called the pyramid for this reason. The part im
Anteriorly, the presphenoid contacts the vomer, mediately surrounding the membranous laby
forming the vomerosphenoidal suture (sutura rinth ossifies first and is composed of very dense
vomerosphenoidalis). The ethmoid also contacts bone. The cartilage which surrounds the inner
the body of the presphenoid, forming the spheno ear, known as the otic capsule, is a conspicuous
ethmoidal suture (sutura sphenoethmoidalis). feature of early embryos. Its sharp petrosal crest
As the orbital wing of the presphenoid bone ex (crista petrosa) extends downward and forward;
tends dorsoanteriorly, the sphenopalatine suture its axis forms an angle of about 45 degrees poste
(sutura sphenopalatina) is formed ventrally, and riorly with a longitudinal axis through the skull.
the sphenofrontal suture (sutura sphenofron- It nearly meets the tentorium ossium dorsally to
talis), dorsally. Posterodorsally, the temporal form a partial partition between the cerebral
wing is overlapped by the squamous temporal and cerebellar parts of the brain; anteriorly it
bone, forming the sphenosquamous suture ends in a sharp point, the apex pyram idalis. Its
(sutura sphenosquamosa). The dorsal end of the cerebral and cerebellar surface (facies encepha-
temporal wing overlaps the parietal bone, form lica) is divided by the petrosal crest into antero-
ing the sphenoparietal suture (sutura spheno- dorsal and posteromedial parts. The third
parietalis). The medial surface of the pterygoid surface, facing ventrolaterally, is the tympanic
process, with the pterygoid bone, forms the surface (facies tympanica).
pterygosphenoid suture (sutura pterygosphe- The posteromedial surface presents several
noidalis). features. The most dorsal of these is the cerebel
lar fossa (fossa cerebellaris), which attains its
Temporal Bone greatest relative size in puppies and houses the
The temporal bone (os temporale) (Figs. 1-13 paraflocculus of the cerebellum. Ventral to the
to 1-16) forms a large part of the ventrolateral cerebellar fossa is a very important recess, the in
wall of the cranium. Its structure is intricate, ternal acoustic meatus (meatus acusticus inter-
owing to the presence of the cochlea and the nus). The opening into this recess is the porus
semicircular canals, and an extension of the acusticus internus. The meatus is an irregularly
nasal pharynx into the middle ear. In a young elliptical depression which is divided deeply by
skull the temporal bone can be separated into the transverse crest (crista transversa). Dorsal to
petrosal, tympanic, and squamous parts. The the crest is the opening of the facial canal, which
petrosal part can further be divided into the contains the facial nerve as well as the cribri
pyramid and mastoid part (pars mastoidea). The form dorsal vestibular area (area vestibularis
20 Chapter 1. T h e S k e l e t a l S y s te m
utriculo-ampullaris) for the passage of nerve cerebrum. Its lateral border is usually grooved
bundles from the semicircular canals. Ventral to by the small middle meningeal artery.
the crest is the ventral vestibular area (area ves The tym panic or ventral surface of the pyra
tibularis saccularis), through which pass addi mid forms much of the dorsal wall of the tym
tional vestibular nerve bundles that come from a panic cavity (cavum tympani). At its periphery
deep, minute depression, the foram en singulare. it articulates with the squamosum dorsally and
The spiral cribriform tract (tractus spiralis fo- the tympanicum ventrally. It can be seen from
raminosus) is formed by the wall of the hollow the outside through the external acoustic meatus.
modiolus of the cochlea. The perforations are An eminence, two openings (windows), and
formed by the fascicles of the cochlear nerve three fossae are the prominent features of this
which arise from the spiral ganglion on the out surface. The barrel-shaped eminence, or pro
side of the modiolus. The cochlea and semicircu montory (promontorium), has at its larger poster
lar canals can be seen by removing a portion of olateral end the round window (fenestra coch
the pyramid. These parts are described in Chap lea), which, in life, is closed by the secondary
ter 17, on the Special Sense Organs. Ventroan- tympanic membrane. Just anterior and slightly
terior to the internal acoustic meatus is the short dorsolateral to the round window is the oval or
canal through the pyramid for the passage of vestibular window (fenestra vestibuli), which is
the trigeminal nerve (canalis trigemini). The occluded by the foot plate of the stapes. The
posteroventral part of the pyramid articulates fossae lie at the angles of a triangle located an
with the occipital bone. On the cerebral surface, terolateral to the windows. The smallest fossa is
or on the border between the cerebral surface a curved groove with its concavity facing the
and the suture for the occipital bone, is the ex oval window; it is the open part of the canal for
ternal opening o f the cochlear canaliculus (aper- the facial nerve peripheral to the genu and ante
tura externa canaliculi cochleae). This opening rior to the stylomastoid foramen. The largest is
is in the anterior edge of the jugular foramen the fossa fo r the tensor tympani muscle (fossa
and is large enough to be probed with a horse m. tensoris tympani); it is a spherical depression
hair. The jugular foram en (foramen jugulare) is which lies anterior to the oval window. A thin
located between the pyramid and the occipital scale of bone with a point extending caudally
bone. A smaller opening for the vestibular aque forms part of its ventral wall. The epitym panic
duct, the apertura externa aqueductus vestibuli, recess (recessus epitympanicus), the third fossa,
is located posterodorsal to the opening of the lies posterolateral to the fossa m. tensoris tym
cochlear canaliculus in a small but deep cleft in pani and at a higher level. The incus and the
the bone. head of the malleus lie in this recess.
The anterodorsal part of the cerebral sur The petrosum contains the osseous labyrinth
face of the pyramid is gently undulating, its only (capsula ossei labyrinthi), which is divided into
features being the digital impressions and eleva three parts: the cochlea, semicircular canals,
tions corresponding to the gyri and sulci of the and vestibule. The basal turn of the cochlea is
Petrosal cre s t
C a n a l f o r m a j ■ s up e rf . p e t r o s a l n
C a n a l f o r t r i g e m i n a l n. ~~ fo r chorda tympani
P e tro sa l crest
C a n a l f o r a c o u s t i c n.
T r a n s v e r s e sulcus
PARS P ETR O SA o r py ra m id
Cerebe-I l a r fossa
- Zuqom atic process
Openincj f o r v e s t i b u l a r a q u e d u c t
T r a n s v e r s e c r e s t of
P A R S M A S T O I D E A -------- ------- i n f a c o u s t i c m e a t u s
M a n d i b u l a r fossa
--------- M a s t o i d p r o c e s s
R e t r o g l e n o i d p r o c e s s - - ~ _ -S-
A rea of a t t a c h m e n t of tym panohyoid
A p e x o f p y r a m i d -----
s nS t y l o m a s t o i d foram en
Carotid n o tch --'
/ Round w in d o w
M a n u b riu m of m a l l e u s /
Ext. a c o u s t i c meatus
Tympa ni c bulla 1
F ig . 1-15. Left temporal bone, lateral aspect.
R e t r o g l e no i d f o r a m e n
E p i ty m p a n i c recess-
y i p m r ~ PARS m a s t o i d e a
Fossa f o r t e n s o r t y m p a n i m-
f _S \ XN.
\ ^^Openiny f o r c o c h l e a r c a n a l i c u i u s
' \
P r o m o n t o r y of p y r a m i d ' ' ' Oval w i n d o w
F ig . 1-16. Left temporal bone, ventral aspect. (Tympanic bulla removed.)
22 Chapter 1. T h e S k e l e t a l S y s te m
located lateral to the ventral part of the internal rection of the auditory tube. The chorda tym
acoustic meatus, its initial turn producing the pani usually passes through a small canal in the
bulk of the promontory. The semicircular canals anterodorsal wall of the bulla tympanica and
(canales semicirculares ossei) are three in num emerges through the petrotym panic fissure (fis-
ber, each located in a different plane posterior sura petrotympanica) by a small opening medial
to the cochlea. The bony vestibule (vestibulum) to the retroglenoid process. When the canal fails
is the osseous common chamber where the three to develop, the opening is through the antero
semicircular canals and the cochlea join. The lateral wall of the tympanic bulla. Both the
oval and round windows of the vestibule com petrosal canal and the canaliculus chordae tym
municate with the tympanic cavity in well pani leave the facial canal at an acute angle and
cleaned skulls. For details of the labyrinth, see course toward the brain.
Chapter 17, on the Special Sense Organs. Two minute canals run from the labyrinth to
The fa c ia l canal (canalis facialis) carries the the cerebral surface of the pyramid. The coch
seventh cranial, or facial nerve. It enters the lear aqu edu ct (aqueductus cochleae) runs ven-
petrosum in the dorsal part of the internal acous trad from a point on the ventral wall of the scala
tic meatus and, after pursuing a sigmoid course tympani near the round window to the border
through the temporal bone, emerges at the sty of the jugular foramen. It carries the perilym
lomastoid foramen. The initial 3 mm. of the phatic duct to the subarachnoid space. The ves
canal, starting at the internal acoustic meatus, is tibular aqu edu ct (aqueductus vestibuli) from
straight. The canal makes its first turn on arriv the vestibule passes posteroventrally to the pos
ing at the thin medial wall of the fossa m. ten terior part of the cerebral surface of the pyramid
soris tympani. At this turn, or genu o f the facia l about 3 mm. dorsoanterior to the cochlear open
canal (geniculum canalis facialis), there is an in ing. This duct is too small to be probed easily. It
distinct enlargement for the sensory geniculate carries the endolymphatic duct to the epidural
ganglion of the facial nerve. In the concavity of space.
this bend is the anterior half of the vestibule. As The m astoid part (pars mastoidea) of the pe
the canal straightens after the first turn, and be trosum is the posterior portion and is the only
fore the second turn begins, it opens into the part to have an external surface. This surface lies
cavity of the middle ear lateral to the oval between the supram astoid foram en (foramen
window. The direction of the second bend of the supramastoideum) dorsally and the stylomastoid
canal is the reverse of that of the first, so that the foram en (foramen stylomastoideum) ventrally,
whole passage is S-shaped, but does not lie in both of which it helps to form. It articulates with
one plane. The fo ssa fo r the stapedius muscle the exoccipital medially and the squamosum
(fossa m. stapedius) is located on the dorsal wall laterally. The ventral part is slightly enlarged,
of the facial canal just before the canal opens into forming the m astoid process (processus mas-
the middle ear cavity from the cranium. After toideus), which serves for muscle attachment
completing its second arc the facial canal opens and articulates with the tympanohyoid cartilage.
to the outside by the deeply placed stylomastoid The facial canal, as it leaves the stylomastoid
foramen (foramen stylomastoideum). The small foramen, grooves the ventral surface of the pars
petrosal canal (canalis petrosus) leaves the mastoidea. The stylomastoid foramen is dorsal
facial canal at the genu by extending anteriorly, to the posterior part of the tympanic bulla.
dorsal to the fossa m. tensoris tympani. It runs The tympanic part (pars tympanica) of the
anteroventrally just within the wall of the fossa temporal bone, or tympanicum, is the ventral
to a small opening near the distal end of the portion and is easily identified by its largest
petrosquamous suture and lateral to the canal component, the smooth bulbous enlargement,
for the trigeminal nerve. If a dark bristle is in or bulla tympanica, which lies between the
serted in the canal its path can be seen through retroglenoid and jugular processes. In puppies
the wall of the fossa. The greater superficial its walls are not thicker than the shell of a hen’s
petrosal nerve passes through the petrosal canal. egg. The cavity it encloses is the fundic part of
The canaliculus chordae tympani carries the the tympanic cavity, which is delimited from
chorda tympani nerve from the facial canal to the dorsal part of the tympanic cavity proper
the cavity of the middle ear. It arises from the by a thin ledge of bone. In old animals this
peripheral turn of the facial canal. After the bony ledge has fine, knobbed spicules protrud
nerve has crossed the medial surface of the han ing from its free border. The osseous external
dle of the malleus it passes under a fine bridge of acoustic meatus (meatus acusticus extemus)
bone of the tympanic ring to continue in the di is the canal from the external ear to the
T h e Sk u l l 23
tympanic membrane. Its length increases with membrane, is the tympanic cavity proper, and
age but rarely exceeds 1 cm. even in old, large its most dorsal extension, for the incus, part of
skulls. It is piriform, with its greatest dimension the stapes, and head of the malleus, is the epi-
dorsoventrally and its smallest dimension trans tym panic recess (recessus epitympanicus).
versely. In carefully prepared skulls the malleus The squamous part (pars squamosa) of the
can be seen through the meatus, somewhat dis temporal bone, or squamosum, possesses a long,
placed but articulated with the incus. All but the curved, zygomatic process (processus zygomati-
dorsal part of the external acoustic meatus is cus) (Figs. 1-15, 1-16), which extends antero-
formed by the tympanicum. The tym panic laterally and overlies the posterior half of the
membrane (membrana tympani), or ear drum, is zygomatic bone in forming the zygomatic arch
a membranous diaphragm attached to the tym (arcus zygomaticus). The ventral part of the
panic ring (annulus tympanicus). If planes are base of the zygomatic process expands to form a
drawn through the ear drums they meet at the transversely elongated, smooth area, the m an
anterior end of the brain case. At the anterior dibular fo ssa (fossa mandibularis), which re
margin of the bulla, lateral to the occipitosphe- ceives the condyle of the mandible to form the
noidal suture, there are paired notches and two tem porom andibular joint (articulatio temporo-
large openings. The more medial of the two mandibularis). The retroglenoid process (proces
openings is the external carotid foram en (fo sus retroglenoideus) is a ventral extension of the
ramen caroticum externum). It is flanked on the squamosum. Its anterior surface forms part of
medial and lateral sides by sharp, pointed proc the mandibular fossa and its posterior surface is
esses of bone from the bulla wall. The medial grooved by an extension of the retroglenoid
process meets the lingula of the sphenoid bone foram en (foramen retroglenoideum). The dorsal
in separating the foramen from the lateral open part of the squamosum is a laterally arched, con
ing, the osseous auditory tube (tuba auditiva vex plate of bone which articulates with the
ossea). By means of the osseous auditory (eusta- parietal bone dorsally, the temporal wing of the
chian) tube, the tympanic cavity communicates basisphenoid bone anteriorly, the tympanicum
with the nasal pharynx. The carotid can al (ca ventrally, and the pars mastoidea and the supra
nalis caroticus) runs longitudinally through the occipital posteriorly. Near the posterolateral
medial wall of the osseous bulla where it articu border of the bone is the ventral part of the dor
lates with the basioccipital bone. It begins at the sal nuchal line. This line is continued anteriorly
posterior carotid foram en (foramen caroticum dorsal to the external acoustic meatus as the
posterius), which is hidden in the depths of the temporal crest (crista temporalis) of the zygoma
petrobasilar fissure. It runs anteriorly, makes a tic process. The smooth, rounded outer surface
ventral turn at a little more than a right angle, above the root of the zygomatic process is the
and opens to the outside at the external carotid fa cies temporalis. The tem poral meatus (meatus
foramen. At its sharp turn ventrad it concurs temporalis), or canal, seen on the inner postero-
with the posterior part of the sphenoid bone ventral surface of the squamosum, forms a pas
which here forms not only the anterior boundary sage for the retroglenoid vein which exits by
of the vertical parts of the carotid canal, but also means of the retroglenoid foramen.
the anterior boundary of an opening in the brain The squamous part of the temporal bone
case, the internal carotid foram en (foramen overlaps the parietal bone, forming a squam ous
caroticum internum). The carotid canal trans suture (sutura squamosa). It also extends over
mits the internal carotid artery. The lateral the caudal margin of the temporal wing of the
boundary of the petrobasilar canal (canalis pet- sphenoid bone, forming the sphenosquamosal
robasilaris) is formed by the tympanic bulla and suture (sutura sphenosquamosa). Anteriorly, the
petrosum. Medially the basioccipital bone zygomatic process of the squamosum meets the
bounds it. The petrobasilar canal contains the zygomatic bone at the temporozygomatic suture
ventral petrosal sinus which parallels the hori (sutura temporozygomatica). Ventrally, the
zontal part of the carotid canal and lies medial tympanic part of the temporal bone meets the
to it. basioccipital to form the anterior part of the
The tympanic cavity (cavum tympani) is the tym pano-occipital joint (synchondrosis tym-
cavity of the middle ear. It can be divided into pano-occipitalis). Posteriorly, the tympanicum
three parts: the largest, most ventral part is lo articulates with the jugular process of the ex-
cated entirely within the tympanic bulla and is occipital to form the posterior part of this joint.
the fundic part. The smaller, middle compart The petrobasilar fissure (fissura petrobasilaris) is
ment, which is located opposite the tympanic formed between these articulations. At the
24 Chapter 1. T h e S k e l e t a l S y s te m
depth of this fissure the petrous temporal bone ternal lamina; and a cribriform plate, to which
articulates with the occipital bone in forming the ethmoturbinates of the lateral masses attach.
the petro-occipital synchondrosis (synchondro The perpendicular plate (lamina perpendicu-
sis petro-occipitalis). laris), or mesethmoid, is a median vertical sheet
of bone which, by articulating with the vomer
below and the septal processes of the frontal and
Ethmoid Bone nasal bones above, forms the osseous nasal sep
The ethmoid bone (os ethmoidale) (Figs. 1- tum (septum nasi osseum). This bony septum is
17 to 1-21) is located between the cranial and prolonged anteriorly by the cartilaginous nasal
facial parts of the skull, both of which it helps septum. Posteriorly, it fuses with the cribriform
to form. It is completely hidden from view in plate, but usually does not extend through it to
the intact skull. Its complicated structure is best form a crista galli. It forms only the ventral half
studied from sections and disarticulated speci of the nasal septum as the septal plates of the
mens. Although unpaired, it develops from frontal and nasal bones extend down halfway
paired anlagen. It is situated between the walls and fuse with it. The perpendicular plate is
of the orbits, and is bounded dorsally by the roughly rectangular in outline, with a rounded
frontal, laterally by the maxillary, and ventrally anterior border and an inclined posterior one,
by the vomer and palatine bones. It consists of so that it is longer ventrally than it is dorsally.
four parts: a median perpendicular plate, or The turbinates of the lateral masses fill the na
lamina; two lateral masses covered by the ex sal cavities so completely that an inappreciable
POSTERIOR
I'M.
' Vomer
Darsal lamina
ANTERI OR
C r ib r if o r m plate
Wing of v o m e r il/ o me r
F ig . 1-18. Vomer and medial aspect of left ethmoid. (Perpendicular plate removed.)
Roman numerals indicate endoturbinates.
Arabic numerals indicate ectoturbinates.
T h e Sk u l l 25
common nasal meatus (meatus nasi communis) ally. It is the sievelike partition between the
remains between each lateral mass and the lat nasal and cranial cavities. Approximately 300
eral surface of the septum. The dorsal border foramina, some as large as 1.5 mm. in diameter,
does not follow the contour of the face, but par perforate the plate and serve for the transmis
allels the hard palate. The roof of the thin exter sion of olfactory nerve bundles. These cribri
nal lamina arises from the dorsal part of the form foram in a (foramina laminae cribrosae) are
perpendicular plate. grouped into tracts which surround the attach
The external lamina (lamina externa), or ments of the turbinates, the larger foramina be
papyraceous lamina, is developmentally the ing adjacent to these attachments as well as
osseous lining of the nasal fundus. It is extremely around the periphery of the bone. Extending
thin and in places it is deficient as it coats the anteromedially from the middle of the lateral
inner surfaces of the heavier bones that form border is a slightly raised, foramen-free ridge of
this part of the face. This lamina is divided into bone which is surrounded by large foramina.
dorsal, lateral, and ventral parts, commonly Caudal to this low ridge the ethmoid concurs
called the roof, side, and floor plates, respec with the presphenoid to form one of the double
tively, of the lateral masses. From its origin on ethm oidal foram in a (foramina ethmoidalia) on
the perpendicular plate the external lamina runs each side. These two foramina carry the ethmoi
dorsally in contact with frontal and nasal parts dal vessel and nerve. A crista galii, dividing the
of the septum, swings laterally over the top of cranial surface of the cribriform plate into right
the ethmoidal labyrinth, forming the ro o f plates and left fossae for the olfactory bulbs of the
or dorsal lam ina (lamina dorsalis), and laterally brain, is present only in old specimens. The
becomes the lateral lam ina (lamina lateralis) as most anterior limit of the single ethm oidal fossa
it partly covers the side of the ethmoturbinates. (fossa ethmoidalis) touches a transverse plane
This portion of the lamina is exceedingly thin, passing through the middle of the orbital open
incomplete in places, and porous throughout. ings. The cribriform plate is not transverse in
Its anterodorsolateral part is channeled to form position, the right and left halves lying in nearly
the uncinate process (processus uncinatus), sagittal planes, and meeting in front at an angle
which is a part of the first endoturbinate as well of about 45 degrees. Its cerebral surface forms
as of the lateral lamina. The uncinate notch (in the inside of a laterally compressed cone which
cisura uncinata), in the meatus between the first is curved in all directions.
two endoturbinates, is located dorsoposterior to The ethmoidal labyrinth (labyrinthus ethmoi
the uncinate process. The depressed area of the dalis) forms the bulk of the lateral mass. It is
lateral lamina, the maxillary fo ssa (fossa maxil- largely composed of delicate bony scrolls, or
laris), forms the medial wall of the maxillary ethmoturbinates (ethmoturbinalia), which at
sinus (sinus maxillaris). The external lamina is tach to the external lamina by basal laminae and
deficient posteriorly, occurring only as paper- attach posteriorly to the cribriform plate. Since
thin, irregular plaques which remain attached the cribriform plate does not extend to the body
to the basal laminae of the scrolls of bone. The of the presphenoid, but only to its inner table, a
individual turbinates arise from the roof and posteriorly extending space on each side of the
lateral portions of this delicate covering. The body is formed. Likewise, the cribriform plate
ventral or transverse lamina (lamina transversa), attaches dorsally to the inner table of the frontal
which forms the floor plate, can be isolated as a bone, which in old, long skulls is separated from
thin, smooth leaf fused to the medial surfaces of the outer table by over 2 cm. Into these spaces
the maxillae. It continues from the ventral part extend the ethmoturbinates. So completely is the
of the lateral plate medially to the vomer in a cavity of the presphenoid filled by the ethmo
transverse, dorsally convex arch. It is closely ap turbinates that the dog is usually regarded as
plied to the horizontal part of the vomer, the not possessing a sphen oidal sinus (sinus sphe-
two conjoined sheets in this manner forming a noidalis), although in every other respect a
partition which separates the ethmoturbinates sinus does exist. Dorsally, the uppermost turbi
in the nasal fundus from the nasopharyngeal nates grow upward and backward from the
duct. cribriform plate into the cavity of the fron tal
The cribriform plate (lamina cribrosa) (Figs. sinus. Usually all compartments of the medial
1-18, 1-19) is a deeply concave partition, pro part of the frontal sinus have secondary linings
truding anteriorly, which articulates with the formed by ethmoturbinates. The anterior end
ethmoidal notches of the frontal bones dorsally of the large lateral compartment contains the
and with the presphenoid ventrally and later end of an ethmoturbinal scroll that is always
26 Chapter 1. T h e S k e l e t a l S y s te m
O u t e r t ab l e o f f r o n t a l bone - L a t p a r t of f r o n t a l si nus
E ctoturbin ote'3 " ~ - - L o c a t i o n o f na s o f r o n t a l opening
I n n e r t a b l e of f r o n t a l bone —
Cribriform plate
E thm oidal forami
— S p h e n o i d a l f oss a
Perpend i c u l a r p l a t e of p a l a t i n e bone
— Me d i al p o r t o f f r o n t a l si nus
1st m o l a r
Hard p a l a t e 1 ' Vomer
F ig . 1-20. Scheme of the ethmoturbinates in cross section immediately anterior to cribriform plate.
Roman numerals indicate endoturbinates.
Arabic numerals indicate ectoturbinates.
- - -N
M id d le nasal meatus - _ j / t '' "
Endoturbi nate "E ’’
i$4 — Maxi I l o t u r b i n a t e c r e s t
Common nasal m e a t u s -----,
Moxilla
" C a r t i l a q i n o u s n as a l se p t u m
V e n t r a l n a s a l m e a t u s - - Typ/
' 2 n(^ p r e m o l a r
Vomer/ Har d p a l a t e
F ig . 1 -2 1 . Schem e of the m axilloturbinates in cross section.
T h e Sk u l l 27
open, allowing free interchange of air between the basal laminae of the endoturbinates and do
the nasal fossa and the sinus. The ethmoturbi not approach the nasal septum as closely as do
nates are surprisingly alike in different speci the endoturbinates. The first two protrude
mens. They may be divided into four long, through the floor of the frontal sinus. According
deeply lying endoturbinates (endoturbinalia I to to Maier (1928), the second ectoturbinate
IV) and six smaller, more superficially lying ecto pushes up into the medial compartment of the
turbinates (ectoturbinalia 1 to 6). The difference frontal sinus, whereas the third ectoturbinate
between these two groups of turbinates is in pushes up into the lateral compartment. Since
their location and not in their form. Each the form of any one turbinate changes so dras
ethmoidal element (turbinate) possesses a basal tically from level to level, these delicate bones
leaf which attaches to the external lamina. can best be studied from sagittally sectioned
Most of these scrolls come from the lateral part heads which have been decalcified.
of this lamina, but some arise from the roof plate The cribriform p late of the ethmoid articu
proper and others from the septal part. Most lates ventrally with the presphenoid to form the
turbinates also attach to the cribriform plate sphenoethm oid suture (sutura sphenoethmoi-
posteriorly. Each ethmoturbinate is rolled into dalis) and with the vomer to form the vomero-
one or more delicate scrolls of lVz to 2'A turns. ethm oid suture (sutura vomeroethmoidalis).
Those turbinates with a single scroll turn ven Laterally and dorsally the frontoethm oidal
trally, with the exception of the first endoturbi- suture (sutura frontoethmoidalis) is formed by
nate, which turns dorsally. The elements with the union of the cribriform plate with the
two scrolls usually turn toward each other, and medial surface of the frontal bone. The trans
thus toward their attachments. Variations are verse lamina and the anterior part of the lateral
common, as the illustrations show. The endo lamina attach to the maxilla, forming the eth-
turbinates nearly reach the nasal septum medi moideomaxillary suture (sutura ethmoideomax-
ally. The first endoturbinate is the longest and illaris). The caudal part of the lateral lamina as it
arises from the dorsal part of the cribriform meets the transverse lamina attaches to the pala
plate posteriorly as well as from the medial part tine bone, forming the palatoethm oid suture
of the roof plate. In the region dorsal to the in (sutura palatoethmoidalis). The lateral lamina
fraorbital foramen it passes from the roof plate attaches to the small lacrimal bone to form the
to the medial surface of the maxilla. Further for lacrimoethmoidal suture (sutura lacrimoethmoi-
ward it attaches to the medial wall of the nasal dalis). Dorsally the dorsal lamina of the eth
bone as the dorsal nasal concha, or nasal turbi moid articulates with the nasal bones to form
nate (nasoturbinale). The uncinate process is the nasoethm oidal suture (sutura nasoethmoi-
formed at the attachment of the basal lamina to dalis). The external lamina intimately fuses
the nasal bone. This process is coextensive with with the bones against which it lies so that in
the lateral lamina and extends posteroventrally a young, disarticulated skull lines and crests are
into the maxillary sinus. The posterior part of present on the inner surfaces of the bones
the first endoturbinate is represented by a dorso- against which the lateral mass articulates. The
medially rolled plate. The small, ventrally in more salient lines are for the attachment of the
folded first ectoturbinate is located dorsally. endoturbinates, and the smaller ones for the at
The second endoturbinate arises from its basal tachment of the ectoturbinates, since the exter
lamina near the middle of the lateral lamina. It nal lamina has largely fused to the bones against
divides into two or more scrolls, which become which it lies.
widened and flattened in a sagittal plane an
teriorly and rest against the posterodorsal part
of the ventral nasal concha, or maxilloturbinate B on es o f t h e F a c e and P alate
(maxilloturbinale). Viewed from the medial
side the third and fourth endoturbinates have Incisive Bone
the same general form as the second. They are
progressively shorter than the second, so that Each incisive bone (os incisivum), or pre
the wide anterior free end of the second over maxilla (Fig. 1-22), carries three upper incisor
laps the third as do shingles on a roof. The teeth. These teeth are anchored in the body
fourth element is the smallest, and lies dorsal to (corpus ossis incisivi) by deep, conical sockets
the wing of the vomer. Posteriorly it invades the (alveoli dentales), which increase in size from
sphenoidal fossa. the medial to the lateral position. The bony
The ectoturbinates are squeezed in between partitions between the alveoli are the inter-
28 Chapter 1. T h e S k e l e t a l S y s te m
alveolar septa (septa interalveolaria). A later posterior half. The division between the two
ally facing concavity on the posterior alveolar parts of this turbinate is arbitrary. The nasal
surface forms the anteromedial wall of the alve bone ends anteriorly in a concave border which,
olus for the canine tooth. The dorsoposterior with that of its fellow, forms the dorsal bound
part of the incisive bone is the curved, tapering ary of the piriform aperture. The lateral pointed
nasal process (processus nasalis), the free an part is more prominent than the medial and is
terior border of which bounds the piriform aper called the nasal process (processus nasalis). The
ture. A minute groove on the medial surface of posterior extremity of the bone is usually
each incisive bone concurs with its fellow to pointed near the midsagittal plane and is known
form the incisive canal (canalis incisivus) in the as the fron tal process (processus frontalis). The
interincisive suture (sutura interincisiva). This nasal bone articulates extensively with its fellow
canal varies in size and position and occasion on the median plane, forming the intem asal su
ally is absent. Extending posteriorly from the ture (sutura internasalis) externally and the
body is the laterally compressed, pointed pala nasoethmoidal suture (sutura nasoethmoidalis)
tine process (processus palatinus). This process, internally. Posteriorly it articulates with the
with that of the opposite bone, forms a dorsal frontal bone, forming the frontonasal suture
sulcus (sulcus septi nasi), in which the anterior (sutura frontonasalis). Laterally the nasal bone
part of the cartilaginous nasal septum fits. The articulates with the maxilla and the incisive bone,
oval space medial to the palatine process is the forming the nasomaxillary suture (sutura naso-
palatine fissure (fissura palatina), which is the maxillaris) and the nasoincisive suture (sutura
only large opening in each half of the bony nasoincisiva), respectively.
palate. The incisive bone articulates posteriorly
with the maxilla to form the incisivomaxillary M axilla
suture (sutura incisivomaxillaris). The postero-
dorsal parts of the right and left palatine proc The maxilla (Fig. 1-24) and the incisive bone
esses form the vom eroincisive suture (sutura of each side form the upper jaw. The maxilla is
vomeroincisiva) as they articulate with the divided grossly into a body, three processes, and
vomer. The medial surface of each nasal process four surfaces. It is the largest bone of the face,
articulates with the nasal bone to form the naso- and bears all of the upper cheek teeth. It is
incisive suture (sutura nasoincisiva). roughly pyramidal in form, with its apex an
teriorly and its wide base posteriorly. Like the
other facial bones, it shows great variation in
Nasal Bone size and form, depending on the skull type.
The smooth external surface (facies facialis)
The nasal bone (os nasale) (Fig. 1-23) is long, of the maxilla has as its most prominent feature
slender, and narrow posteriorly, but in large an elliptical infraorbital foram en (foramen infra-
dogs is almost 1 cm. wide anteriorly. The dorsal orbitale), for the passage of the infraorbital
or external surface (facies externa) of the nasal nerve and artery. The ventrolateral surface of
bone varies in size and shape, depending on the the bone which bears the teeth is the alveolar
breed. In brachycephalic types the nasal bone is process (processus alveolaris). The partitions
very short, whereas in dolichocephalic breeds of between adjacent teeth are the interalveolar
the same weight its length may exceed its width septa (septa interalveolaria), and the septa be
by 15 times. The external surface usually pre tween the roots of an individual tooth are the
sents a small foramen at its midlength for the interradicular septa (septa interradicularia).
transmission of a vein. The smooth elevations on the lower facial surface
The ventral or internal surface (facies in of the maxilla caused by the roots of the teeth
terna), in life, is covered by mucous membrane. are the alveolar juga (juga alveolaria), thejuga
It is deeply channeled throughout its anterior for the canine and the lateral roots of the shear
half, where it forms the dorsal nasal meatus ing tooth (fourth upper premolar) being the
(meatus nasi dorsalis) and bears the nasal turbi most prominent. The alveolar process contains
nate. The posterior half of the nasal surface is fifteen sockets (alveoli dentales) for the roots of
widened to form the shallow ethm oidal fossa, the seven teeth that it contains. Where the teeth
which bounds the dorsal part of the lateral mass are far apart the spaces between them are
of the ethmoid. The nasoturbinate crest (crista known as interdental spaces, and the margin of
nasoturbinalis) is a thin shelf of bone which gives the jaw at such places is called the interalveolar
rise to the nasal turbinate throughout its an margin. Interdental spaces are found between
terior half and to the first endoturbinate in its each of the four premolar teeth and posterior to
T he Sk u l l 29
- Nasal process
A l v e o l u s fo r c a n i n e tooth
m ed ia l side L o c a t i o n of p a l a t i n e f i s s u r e
---
W 2" -----P a l a t i n e p r o c e s s
N asoturbinate crest
A
Lat. s u r f a c e /\
Frontal p ro ce s s
Nasal p r o c e s s ^
- - E t h m o i d a l fossa
S e p ta l process
F ig . 1-23. Left nasal bone, ventral lateral aspect.
Entrance to i n f r a o r b i t a l c a n a l x D o r s a l e t h m o i da I c r e s t
A lv e o l a r foramina Crest f o r u n c i n a t e p r o c e s s
the canine tooth. The lateral border of the alveo posteriorly by the anterior border of the pala
lar process (margo alveolaris) is scalloped as a re tine process of the maxilla. The most posterior
sult of the presence of the tooth sockets, with process of the maxilla is a small pointed spur,
their interalveolar and interradicular septa. the pterygoid process o f the maxilla (processus
There are three alveoli for each of the last three pterygoideus), located posteromedial to the al
cheek teeth, two each for the next two an veolus for the last cheek tooth. This process
teriorly, and one for the first cheek tooth. In ad and the palatine bone form a notch, rarely a
dition to these alveoli the large posteriorly foramen, through which the minor palatine ves
curved alveolus for the canine tooth lies dorsal sels pass.
to those for the first two cheek teeth, or pre The nasal surface (facies nasalis) of the max
molars I and II. Lying dorsal to the three alveoli illa is its medial surface, and bears several crests.
for the shearing tooth is the short in fraorbital The maxilloturbinate crest (crista maxilloturbi-
can al (canalis infraorbitalis). This canal begins nalis) begins at or near the incisivomaxillary
posteriorly at the maxillary foram en (foramen suture, runs posteriorly, inclines ventrally, and
maxillare). Leading from the infraorbital canal terminates anterior to the opening of the maxil
to the individual roots of the premolar teeth lary sinus. The small ventral ethmoidal crest
(first four cheek teeth) are the alveolar canals (crista ethmoidalis ventralis) is a sagittofrontal
(canales alveolares), which open by numerous crest for the attachment of the floor plate or
alveolar fo ra m in a (foramina alveolaria) at the transverse lamina of the ethmoid bone. The
apex of each alveolus. The special incisivomax- dorsal ethm oidal crest (crista ethmoidalis dor
illary canal (canalis maxilloincisivus) carries the salis) limits the maxillary sinus dorsally and
nerves and blood vessels to the first three pre marks the line of attachment of the lateral
molar and the canine and incisor teeth. It leaves lamina of the ethmoid to the maxilla. An ob
the medial wall of the infraorbital canal within lique line passes from the nasoturbinate crest
the infraorbital foramen, passes dorsal to the posteroventrally and laterally to the mouth of
apex of the canine alveolus with which it com the maxillary sinus to which the uncinate proc
municates, and enters the incisive bone. It con ess of the ethmoid is attached. The lacrimal
tinues anteriorly and medially in the incisive canal (canalis lacrimalis) continues from the
bone, giving off branches to the incisor alveoli. lacrimal bone into the maxilla, where it opens
The frontal process (processus frontalis) ventral to the nasoturbinate crest. The medial
arches dorsally between the nasal bone and or wall of the canal is thin and may be incomplete.
bit to overlap the frontal bone in a squamous The maxillary sinus or recess (recessus maxil-
suture. The zygomatic process (processus zygo- laris) lies medial to the infraorbital and lacrimal
maticus) is largely hidden, in an articulated canals, both of which protrude slightly into it.
skull, by the laterally lying zygomatic bone The lateral wall of the maxillary sinus is formed
which is mitered into the maxilla both above largely by the maxilla with the addition of the
and below the bulk of the process. This type of palatine bone posteriorly. The floor of the ptery
articulation prevents dislocation at a place gopalatine fossa (fossa pterygopalatina) lies pos
where injury frequently occurs. The palatine terior to the maxillary foramen. The shelf of
process (processus palatinus) is a transverse bone which forms it is thicker anteriorly and
shelf of bone which, with its fellow, forms most contains many foramina which lead to the al
of the hard p a la te (palatum osseum) and sepa veoli for the last two cheek teeth. The thin pos
rates the respiratory from the digestive passage terior part, barely thick enough to cover the
way. The dorsal surface of the palatine process roots of the last molar tooth, is the maxillary
forms part of the floor of the ventral nasal tuberosity (tuber maxillae).
meatus. Its ventral su rface (facies palatina) is The maxilla articulates with the incisive bone
grooved on each side by the palatine sulcus anteriorly, forming the incisivomaxillary suture
(sulcus palatinus) and forms part of the roof of (sutura incisivomaxillaris). Dorsomedially, the
the oral cavity. Each sulcus extends anteriorly nasal bone meets the maxilla at the nasomaxil
from the major palatine foram en (foramen pal- lary suture (sutura nasomaxillaris). Dorsopos-
atinum majus), which is an oval, oblique open teriorly, the maxilla articulates with the frontal
ing in the suture between the palatine process bone, forming the frontomaxillary suture (sutura
of the maxilla and the palatine bone. In some frontomaxillaris) at its dorsocaudal angle. Ven
specimens the palatine sulcus may reach the tral to this suture the lacrimal bone and maxilla
palatine fissure (fissura palatina), which is a form the short lacrim om axillary suture (sutura
large, sagittally directed oval opening formed lacrimomaxillaris). The ventrolateral part of the
T he Sk u l l 31
maxilla forms the unusually stable zygom atico bony scrolls is posteroventral. Usually five pri
maxillary suture (sutura zygomati com axillaris), mary scrolls can be identified, and they are
as it articulates with the zygomatic bone. Ven- numbered, dorsoventrally, from 1 to 5. The first
troposteriorly, the maxilla forms the extensive primary unit leaves the dorsal surface of the
palatomaxillary suture (sutura palatomaxillaris) basal lamina and runs toward the nasoturbinate.
with the palatine bone. The m edian palatine It is displaced laterally in its posterior part by
suture (sutura palatina mediana) is formed by endoturbinates I and II. The second primary
the two palatine processes. The transitory joint unit arises several millimeters peripheral to the
between the ethmoid and maxilla is the ethmoi- first, and some of its subsequent leaves reach
deomaxillary suture (sutura ethmoideomaxil- nearly to the nasal septum. The third unit and
laris). The vom erom axillary suture (sutura its secondary and tertiary scrolls largely fill the
vomeromaxillaris) is formed in the median space formed by the union of the nasal septum
plane, within the nasal cavity. with the hard palate. The fourth unit at first runs
ventrally nearly to the palate and then inclines
medially, ventral to the third unit. The fifth, or
Nasal Turbinate
terminal, unit curves dorsally as a simple pos
teriorly closed scroll which runs under the
The nasal turbinate (os nasoturbinale) is the maxilloturbinate crest. It has fewer secondary
continuation of endoturbinate I of the ethmoid, scrolls than do the others. The secondary scrolls
which attaches by means of the nasoturbinate divide further, so that the whole nasal fossa is
crest (Fig. 1-23) to the nasal bone. Baum nearly filled with a labyrinthine mass of deli
and Zietzschmann (1936) regarded the first cately porous, bony plates. The larger the nasal
endoturbinate and the nasal turbinate as one cavity, the more numerous the bony scrolls.
structure. In this description, however, the nasal
turbinate is regarded as that turbinate which
Zygomatic Bone
begins at the uncinate process and continues an
teriorly, attached to the nasal bone. The unci The zygomatic (os zygomaticum), jugal, or
nate process and the posteriorly extending scroll
malar bone (Fig. 1-25) forms the anterior half of
constitute endoturbinate I of the ethmoid. The
the zygomatic arch (arcus zygomaticus). It is di
nasal turbinate, unlike the ethmoturbinates and
vided into two surfaces, four borders, and four
maxilloturbinate, is a simple, curved shelf of
processes. The lateral surface (facies lateralis) is
bone which is separated from the ventrally lying
convex longitudinally and transversely, although
maxilloturbinate by a small cleft, the middle
it is slightly dished ventral to the orbit. Usually
nasal meatus (meatus nasi medius). In life, the
a nutrient foramen is present near its middle.
scroll is continued anterior to the nasoturbinate
The medial or orbital surface (facies orbitalis) is
crest by a plica of mucosa which diminishes and
concave in all directions. The maxillary border
disappears in the vestibule of the nose.
articulates broadly with the maxilla and is re
cessed to form an unusually stable foliate type of
Maxilloturbinate Bone sutural joint. At the middle of this articular bor
der the zygomatic bone receives the zygomatic
The maxilloturbinate bone (os maxilloturbi- process of the maxilla, which it partly overlays.
nale) (see Fig. 1-21) is irregularly oval in shape, The temporal border forms a long harmonial
with its extremities anterior and posteroventral suture with the zygomatic process of the tem
in position. It is attached to the medial wall of poral bone. This suture is one of the last to close.
the maxilla by a single basal lamina, the maxillo- The orbital border (margo orbitalis) forms the
turbinate crest. The com m on nasal meatus ventral margin of the eye socket. It is thick and
(meatus nasi communis) is a small sagittal space beveled medially. The masseteric border
between the maxilloturbinate and the nasal sep (margo massetericus) is also thick, but is beveled
tum. The space dorsal to the maxilloturbinate is laterally. Both the thickness of the border and
the middle nasal m eatus (meatus nasi medius), the degree to which it is beveled decrease pos
and the space ventral to it is the ventral nasal teriorly. This border provides the origin for the
meatus (meatus nasi ventralis). The osseous strong masseter muscle. The lacrimal process
plates are continued as soft tissue folds which (processus lacrimalis) of the zygomatic bone is
converge anteriorly to form a single medially turned up and is of uniform width; it articulates
protruding ridge that ends in a clublike emi with the maxilla and lacrimal bones. The pos
nence in the vestibule. The direction of the teroventral margin of the zygomatic bone is
32 Chapter 1. T h e S k e l e t a l S y s te m
/ Frontal p r o c e s s
L a c r im a l process '
j - T e m p o r a l process
A r t i c u l a t i o n w ith m axilla-
Masseteric mar gin
M a x illa ry process
F ig . 1-25. Left zygomatic bone, lateral aspect.
of the horizontal part concurs with a similar, but pendicular part. The thin, irregularly convex
always deeper, notch in the maxilla to form the border of the most dorsal part of the palatine
major palatine foram en (foramen palatinum bone is the ethm oidal crest (crista ethmoidalis),
majus), which opens on the hard palate. Pos which conceals the medially lying ethmoidal
terior to this foramen there is usually one or, oc bone. The medial wall of the pterygopalatine
casionally, two or more minor palatine foram ina fossa is formed by the palatine bone. The
(foramina palatina minora). All of these open round dorsal opening is the sphenopalatine f o
ings lead into the palatin e can al (canalis pala ramen, and the oblong ventral one, about 1
tinus), which runs through the palatine bone mm. distant, is the posterior palatine foram en
from the pterygopalatine fossa. This canal trans (foramen palatinum posterius). The palatine
mits the major palatine artery, vein, and nerve. bone articulates posteriorly with the sphenoid
The perpendicular lamina (lamina perpen- and pterygoid bones, anteriorly with the max
dicularis) of the palatine bone leaves the pos illa and ethmoid, dorsolaterally with the lacri
terolateral border of the horizontal lamina at mal and frontal bones, dorsomedially with the
nearly a right angle. Medially it forms the vomer, and ventromedially with its fellow at
lateral wall (facies nasalis) of the nasopharyn the m edian palatine suture (sutura palatina
geal meatus, and laterally it forms the medial mediana). Anteriorly the palatine bones articu
wall (facies maxillaris) of the pterygopalatine late with the maxillae by a suture which crosses
fossa. The nasal surface is pardy divided by a the mid line, the transverse palatine suture
frontally protruding shelf, the lam ina spheno- (sutura palatina transversa). Dorsally, at the
ethmoidalis. This shelf parallels the horizontal anterior end of the median palatine suture, the
part of the bone as it lies dorsal and extends vomer articulates with the palatine bones, form
about half its length posterior to it. Dorsal to ing the vom eropalatine suture (sutura vomero-
the anterior end of the sphenoethmoid lamina palatina). In the medial part of the pterygo
is the sphenopalatine foram en (foramen sphe- palatine fossa the palatine bone articulates
nopalatinum), which lies dorsal to the posterior with the maxilla, forming the palatom axillary
palatine foramen and extends from the ptery suture (sutura palatomaxillaris), which is a con
gopalatine fossa to the nasal cavity. The nasal tinuation of the transverse palatine suture.
vessels and nerve of the sphenopalatine foramen Where the palatine bone articulates with the
groove the anterior end of the lamina spheno- pterygoid process of the sphenoid bone, as well
ethmoidalis. The area dorsal to this lamina is as with its orbital wing, the sphenopalatine
articular for the orbital wing of the sphenoid suture (sutura sphenopalatina) is formed. The
and the lateral lamina of the ethmoid bones. pterygopalatine suture (sutura pterygopalatina)
The anterior part of the nasal surface is exca is formed by the small pterygoid bone articu
vated to form the posterolateral part of the lating with the medial surfaces of the posterior
maxillary fossa (fossa maxillaris), which bounds part of the palatine bone as it unites with the
part of the maxillary sinus laterally. The small pterygoid process of the sphenoid. On the
ventral ethmoidal crest, located at the postero medial side of the orbit the fron topalatin e
ventral margin of the maxillary fossa, marks the suture (sutura frontopalatina) runs dorsoanteri-
line along which the lateral lamina of the eth orly. The deep surface of the palatine bone joins
moid articulates with the palatine bone to form anteriorly with the ethmoid bone to form the
the medial wall of the maxillary sinus. The part palatoethm oidal suture (sutura palatoethmoid-
of the palatine bone ventral to the sphenoeth alis).
moid crest is smooth, slightly concave, and
faces medially to form the anterior part of the
lateral wall of the nasopharyngeal meatus. The Lacrimal Bone
posterior border of the hard palate provides
attachment for the soft palate. The perpendic The lacrimal bone (os lacrimale) (Fig. 1-27),
ular lamina of the palatine bone has three proc located in the anterior margin of the orbit, is
esses. The posterior part between the pterygoid roughly triangular in outline and pyramidal in
bone medially and the sphenoid bone laterally shape. Its orbital face (facies orbitalis) is con
is the sphenoidal process (processus sphenoid- cave and free. Located in its center is the fossa
alis). fo r the lacrim al sac (fossa sacci lacrimalis),
The maxillary process (processus maxillaris) which is about 6 mm. in diameter. The two
of the palatine bone articulates with the maxilla lacrimal ducts, one from each eyelid, unite in
at the anteroventrolateral extremity of the per a slight dilatation to form the lacrimal sac. From
34 Chapter 1. T h e S k e l e t a l S y s te m
the lacrimal sac the soft nasolacrimal duct the pterygoid bone and the pterygoid process
courses to the vestibule of the nose. The osse of the sphenoid is the minute pterygoid canal
ous lacrimal canal (canalis lacrimalis), contain (canalis pterygoideus), which carries the auto
ing the nasolacrimal duct, begins in the lacrimal nomic nerve of the pterygoid canal. The ptery
bone at the fossa for the lacrimal sac, runs goid bone forms an extensive squamous suture
ventroanteriorly through the lacrimal bone, and with the pterygoid process of the sphenoid
leaves at the apex of the bone. It continues in bone posteriorly, the pterygosphenoid suture
a dorsally concave groove in the maxilla and (sutura pterygosphenoidalis), and with the pala
opens ventral to the posterior end of the max tine bone anteriorly, the pterygopalatine suture
illoturbinate crest. The orbital crest (crista (sutura pterygopalatina).
orbitalis) is that part of the lacrimal bone which
forms the margin of the orbit. The frontal Vomer
process (processus frontalis) is a narrow strip
of the orbital margin which projects dorsally. The vomer (Fig. 1-29) is an unpaired bone
The crest is formed by the facial surface (facies which forms the posteroventral part of the
facialis) meeting the orbital surface at an acute nasal septum and therefore forms the medial
angle. Only a small part of the facial surface is boundaries of the choan ae, or posterior nares,
free; most of it is covered by the maxilla and and most of the medial wall of the nasopharyn
zygomatic bones. In some specimens a free geal meatus. Since this bone runs obliquely
facial surface is lacking. The nasal surface from the base of the neurocranium to the nasal
(facies nasalis) forms a small portion of the surface of the hard palate, the choanae are
nasal cavity. It contains a small ethm oid crest located in oblique planes in such a way that
(crista ethmoidalis) for articulation with the the ventral parts of the choanae are anterior to
ethmoid bone. a transverse plane through the posterior border
The lacrimal bone articulates dorsoposteriorly of the hard palate. The choanae are the open
with the frontal bone, forming the frontolacri- ings whereby the right and left nasopharyngeal
m al suture (sutura frontolacrimalis); anteriorly meatuses are continued as the single basipha
with the maxilla, forming the lacrimomaxillary ryngeal canal, the skeletal base of the nasal
suture (sutura lacrimomaxillaris); and antero- pharynx. The vomer is divided into sagittal and
ventrally with the zygomatic bone, forming the horizontal parts.
zygomaticolacrimal suture (sutura zygomat- The sagittal part is formed of two thin, bony
icolacrimalis). Posteroventrally the palatola- leaves, laminae laterales, which unite ventrally
crimal suture (sutura palatolacrimalis) is formed to form a sulcus, sulcus septi nasi, which in
by the articulation between the palatine and turn receives the cartilaginous nasal septum
lacrimal bones. Medially, the ethmoid bone anteriorly and the bony nasal septum, or per
articulates with the lacrimal, forming the lacri- pendicular plate of the ethmoid, posteriorly. It
m oethm oidal suture (sutura lacrimoethmoidalis). articulates ventrally with the palatine processes
of the maxillae, with the posterior parts of the
Pterygoid Bone palatine processes of the incisive bones, and
with the anterior parts of the horizontal por
The pterygoid bone (os pterygoideum) (Fig. tions of the palatine bones. This posterior artic
1-28) is a small, thin, slightly curved, nearly ulation is at the palatine suture and the ventro-
four-sided plate of bone which articulates with anterior half of the base o f the vomer (basis
the bodies of both the presphenoid and the vomeris). The sagittal part of the vomer is
basisphenoid bone, but particularly with the sharply forked at each end. The anterior notch
medial surface of the pterygoid process of the is the incisive incisure (incisura incisiva); the
basisphenoid. It extends ventrally beyond this posterior is the sphenoidal incisure (incisura
process, to form the posterior part of the osse sphenoidalis).
ous lateral wall of the nasal pharynx, or the The horizontal part of the vomer, constitut
basipharyngeal canal. The posteroventral angle, ing the wings (alae vomeris), is at right angles
hamulus pterygoideus, is in the form of a pos to the sagittal part, flaring laterally and articu
teriorly protruding hook around which, in life, lating with the sphenoid, ethmoid, and palatine
the tendon of the m. tensor veli palatini plays. bones. The wings, with the transverse lamina
The smooth concave medial surface is the facies of the ethmoid, form a thin septum that sepa
nasopharyngea. Running in the suture between rates the dorsally lying nasal fundus, in which
T h e Sk u l l
. Frortal process
O r b i t a l c r e s t ------&
H a m u lu s pferijCjoideus
■ f ------I n c i s i v e i n c i s u r e
|
>
I
14 * r -
k
j ■.
A r t i c u l a t i o n w ith lamina- \
l a t e r a l i s of et hmoi d A
A r t i c u l a t i o n w i t h p a l a t i n e ----- M w
-------- Sphen o i d a l i n c i s u r e
F ig . 1-29. Vomer, ventral aspect.
36 Chapter 1. T he S k e l e t a l Sy stem
lie the ethmoturbinates, from the ventrally mandibulae), in which lies the tongue. On each
lying nasopharyngeal meatuses and the nasal side the body of the mandible presents a lateral
pharynx. su rface which faces to the cheek (facies buc-
The vomer articulates dorsally with the sphe calis) and lips (facies lingualis), and a lingual sur
noid bone, forming the vom erosphenoid suture fa c e (facies lingualis), which faces the tongue.
(sutura vomerosphenoidalis). Anterior to this The lingual surface may present a wide, smooth,
suture and hidden from external view is the longitudinal ridge, the m ylohyoid line (linea
vomeroethmoid suture (sutura vomeroethmoi- mylohyoidea), for the attachment of the mylo
dalis), for articulation with the ethmoid bone. hyoid muscle. Anteriorly, the lingual surface
Laterally the wings of the vomer articulate with gives way to the sym physeal surface, which
the palatine bones, forming the dorsal vomero- articulates extensively with its fellow. The
palatine suture (sutura vomeropalatina dorsalis). lateral surface is long, smooth, and of a uniform
The vomer articulates with the conjoined pala width posterior to the symphysis. It ends in the
tine crests to form the ventral vomer op alatine thick, convex ventral border, with which the
suture (sutura vomeropalatina ventralis). An lateral and lingual surfaces are confluent. An
terior to this suture the vomer articulates with teriorly it turns medially and presents the
the palatine processes of the maxillae and in anterior mental foram en (foramen mentale
cisive bones to form the vomeromaxillary su anterius near the symphysis, ventral to the
ture (sutura vomeromaxillaris) and the vomero- alveolus of the central incisor tooth. The larg
incisive suture (sutura vomeroincisiva), respec est of the mental foramina, the m iddle mental
tively. foramen (foramen mentale medium), is located
ventral to the septum between the first two
cheek teeth. The small posterior mental foram en
Mandible (foramen mentale posterius) is located about
1 cm. posterior to the middle opening.
The mandible (mandibula) (Fig. 1-30) of the The ramus of the mandible (ramus mandib
dog consists of right and left halves firmly united ulae) is the posterior non-tooth-bearing, verti
in life at the m andibular symphysis (symphysis cal part of the bone. It contains three salient
mandibulae), which is a strong, rough-surfaced, processes. The coronoid process (processus cor-
fibrous joint. Each half is divided into a hori onoideus), which forms the most dorsal part of
zontal part, or body, and a vertical part, or the mandible, extends upward and outward. It
ramus. is a large thin plate of bone with a thickened
The body of the mandible (corpus mandib anterior border. The condyloid or articular
ulae) can be further divided into the part that process (processus condylaris s. articularis) is a
bears the incisor teeth (pars incisiva), and the transversely elongated, sagittally convex articu
part that contains the molar teeth (pars molaris). lar process which forms the temporomandibular
The sockets, or alveoli (alveoli dentales), which joint by articulating with the temporal bone.
are conical cavities for the roots of the teeth, The mandibular notch (incisura mandibulae) is
indent the alveolar border (arcus alveolaris) of located between the condyloid and coronoid
the body of the mandible. There are single processes. The angle o f the m andible (angulus
alveoli for the roots of the three incisor teeth, mandibulae) is the posteroventral part of the
the canine, and the first and last cheek teeth. bone. It contains a salient hooked process in the
The five middle cheek teeth have two alveoli dog, the angular process (processus angularis).
each, with those for the first molar, or fifth The lateral surface of the ramus contains a
cheek tooth, being the largest, as this is the prominent, three-sided depression, the mas
shearing tooth of the mandible. The free dorsal seteric fo ssa (fossa masseterica), for the inser
border of the mandible between the canine and tion of the strong masseter muscle. This muscle
first cheek tooth (first premolar) is larger than attaches to the coronoid and condyloid proc
the others and is known as the interalveolar esses and is limited by the coronoid crest (crista
margin (margo interalveolaris). Similar but coronoidea) anteriorly and by the condyloid
smaller spaces are usually present between ad crest (crista condyloidea) posteriorly. The me
jacent premolar teeth, where the interalveolar dial surface of the ramus is slightly dished for
septa (septa interalveolaria) end in narrow the insertion of the temporal muscle. Directly
borders. From the symphysis, the bodies of ventral to this insertion is the m andibular f o
each half of the mandible diverge from each ram en (foramen mandibulae). This foramen is
other, forming the m andibular space (spatium approximately 8 mm. dorsal to the ventral
T he Skull 37
R a m u s of m a n d i b l e ^ p; Coronoid c r e s t
^ ' M a n d i b u l a r notch
M a n d i b u l a r foram en ^ ^M asseteric fossa
- Condyloid eres t
Mylohyoid c re s t— |
Ancjular process
x Mosseteric lin e
The basihyoid bone or body (os basihyoideum) Dorsal Surface of Skull (Fig. 1-3)
is a transverse unpaired element lying in the
musculature of the base of the tongue as a ven Cranial part. The dorsal surface of the cranial
trally bowed, dorsoventrally compressed rod. or neural part of the skull (neurocranium) is
Its extremities articulate with both the thyro nearly hemispherical in the newborn and is de
hyoid and the keratohyoid bones. void of prominent markings. On the other hand,
a skull from a heavily muscled adult possesses a
prominent external sagittal crest, a median lon
Thyrohyoid Bone gitudinal projection which is the most prominent
feature of the dorsal surface of the skull. Poste
The thyrohyoid bone (os thyrohyoideum), or riorly, the dorsal surface is limited by the dorsal
cornu majus of man, is a laterally bowed, sagit- nuchal line, a transverse, variably developed
tally compressed, slender element which ex crest which marks the transition between the
tends dorsocaudally from the basihyoid to ar dorsal and the posterior surface of the skull. An
ticulate with the cranial cornu of the thyroid teriorly, in brachycephalic skulls, the external
cartilage of the larynx. sagittal crest gives way to the right and left tem
poral lines, which diverge from the sagittal crest
at the anterior end of the interparietal process of
Keratohyoid Bone the occipital bone. In dolichocephalic skulls the
external sagittal crest ends about 1 cm. anterior
The keratohyoid bone (os keratohyoideum) is to the coronal suture, where it gives rise to the
a small, short, tapered rod having a distal extrem external frontal crests, which continue anteriorly
ity which is about twice as large as its proximal to the zygomatic processes of the frontal bones.
extremity. It articulates with the basihyoid and The convex surface on each side of the dorsum
the thyrohyoid. The proximal extremity, which of the skull is the temporal fossa, from which the
points nearly cranially in life, articulates with temporal muscle arises. Each is bounded by the
the epihyoid bone at a right angle. external frontal crest anteriorly, by the temporal
T he Skull 39
line and the external sagittal crest medially in ess is the bu lla tym pan ica, which can be seen
brachycephalic breeds, and by the dorsal nuchal best from the ventral aspect. The jugular process
line posteriorly in all breeds. This surface of the is a sturdy ventral projection posterior to the
skull is the parietal plane (planum parietale). bulla tympanica, and lateral to the occipital
The frontal plane (planum frontale) begins condyle.
anteriorly at a transverse plane through the pos The orbital region is formed by the orbit and
terior ends of the nasal bones. This plane passes the ventrally lying p terygopalatin e fo s sa . The
through the cribriform plate. The frontal plane orbital opening faces anterolaterally, and is
is a curved pentagon in shape, and of variable nearly circular in the brachycephalic breeds and
size, depending on the development of the fron irregularly oval in the dolichocephalic. Approx
tal sinuses. Its anterior boundary is imaginary, imately the posterior fourth of the orbital mar
but the two lateral limits are the concave dorsal gin is formed by the orbital ligam ent. A line
margins of the orbits and the convex to nearly from the center of the optic canal to the cen
straight external frontal crests. ter of the orbital opening is the axis of the orbit.
Facial part. The dorsal surface of the facial The eyeball and its associated muscles, nerves,
part of the skull is extremely variable, depend vessels, glands, and fascia are the structures of
ing on the breed, and is greatly foreshortened in the orbit. Only the medial wall of the orbit is en
brachycephalic dogs. It is formed by the dorsal tirely osseous. Its posterior part is marked by
surfaces of the nasal, incisive, and maxillary three large openings which are named, from an-
bones, and the nasal processes of the frontal terodorsal to posteroventral, the optic canal, or
bones. Its most prominent feature is the un bital fissure, and anterior alar foram en. In addi
paired external nasal opening or piriform aper tion to these there are usually two ethm oidal
ture (apertura nasi ossea s. piriformis). In foram ina, which are located anterodorsal to the
brachycephalic skulls this opening is not piri optic canal. Within the anterior orbital margin
form, since its transverse dimension is greater is the fossa fo r the lacrimal sac. The lacrimal
than its dorsoventral one. c a n a lle aves the fossa and extends anteroven-
The stop, or glabella, present only in brachy trally. Ventral to the medial surface of the orbit,
cephalic skulls, is a wide, smooth, transverse and separated from it by the dorsally arched
ridge which lies directly dorsal to the dish of the ventral orbital crest (crista orbitalis ventralis), is
face or in a transverse plane through the postero- the pterygopalatin e fo ssa . T h e anterior end of
dorsal parts of the frontomaxillary sutures. An this fossa funnels down to the maxillary foramen
unpaired midsagittal fossa, the frontal fossa, ex which is located dorsal to the posterior end of
tends forward on the nasal bones from the fron the shearing tooth. A small part of the medial
tal bones. wall of the fossa just posterior to the maxillary
foramen frequently presents a defect. Still
Lateral Surface of the Skull farther posteriorly are the more ventrally lo
(Figs. 1-32, 1-33) cated sphenopalatine foram en and the posterior
palatine foram en. The more dorsally located
Cranial part. The salient features of the lat sphenopalatine foramen is separated from the
eral surface of the cranial part of the skull are posterior palatine foramen by a narrow septum
the prominent zygomatic arch and the orbit. of bone. The ventral orbital crest marks the dor
The zygomatic arch is a heavy, laterodorsally sal boundary of the origin of the medial ptery
convex bridge of bone located between the facial goid muscle. The crest ends posteriorly in the
and neural parts of the skull; it is laterally com septum between the orbital fissure and the an
pressed anteriorly and laterally, and dorsoven- terior alar foramen. The posterior border of the
trally compressed posteriorly. It is composed of pterygoid bone also forms the posterior border
the zygomatic bone and the zygomatic proc of the pterygopalatine fossa.
esses of the temporal and maxillary bones. It Facial part. The lateral surface of the facial
serves three important functions: to protect the part of the skull is formed primarily by the max
eye, to give origin to the masseter and a part of illa. It is gently convex dorsoventrally, and has
the temporal muscle, and to provide an articu as its most prominent feature the vertically oval
lation for the mandible. The external acoustic infraorbital foram en , which lies dorsal to the
meatus is the opening of the external acoustic septum between the third and fourth cheek
process to which the external ear is attached. teeth. The alveolar juga of the shearing and ca
Ventral and medial to the external acoustic proc nine teeth are features of this surface.
40 Chapter 1. T he Sk el et a l S y stem
Tempor al w i ng of
Z yg o m a tic, , sphen oi d
Pa l a t i n e
L a crim a I Temporal
\
M a x i I la \
\
Nasa I \
- O ccip ita !
In c i s i ve\
" T y m p a n o h y o i d ca r t i l ag e
Stylohyoid
Ke r a t o h y o i d -
Basi hyoid/
T hyrohyoid1 I \ Trachea
T e m p o r a l f os s a
Ext. f r o n t a l c r e s t lExt. s o g i t t a i c r e s t
Z y g o ma t ic p ro c e s s of f r o n t a l bone ^ Do r s a l n u c h a l I i n e
F th mo i d a I f o r a r n i r a
i J n t e r p a r i e t a i process
Groove f o r a n g u l a r i s o c u l i v.
I /
i / Ext. occi pit al protuberance
O rb ita l m argin
Supramastoid f o r a me n
Ventral o r b i t a l crest-
M a s t o i d process
Fossa for l a c r i m a l sac
Dor sal c o n d y l oi d fossa
Fo r a me n m a g n u m
V e n t r a l c o n d y l o i d fosse
i - --- . J u g u l a r process
V \ ' v v
^ \^ \ \ X S t y l o m a s t o i d foramen
I nf raorbi tal foramen ' / / IV x i ^ Fxt. a c o u s t i c m e a t u s
------------------------------ / / / \ \ ' \ \
A lve o la rju g a / ’ / / ' ' \ ' y ' Tympanic bulla
Sphenopal at i ne foramen/ ^ t
/ / \ \ \ Siet roal p. noid f o r a m e n
Posterior p a i a t i n e . f o r a m e n ■
/' / I \ \ \ Retroglenoid p r o c e s s
P te r y g o i d process' j Po s t e r i or a l a r f o r a m e n
D p t ic c a n a l
/ / 1 ' i n t e r io r a la r foramen
i ' ------------------------------ —
/ i ' •Pterygoi d b o n e
i n t e r i o r o p e n i n g of p t e r y c j o i d canal
Ventral Surface of the Skull (Fig. 1 -3 4 ) pharynx, which starts anteriorly at the choanae
and ends posteriorly at the pharyngeal isthmus.
Cranial part. The ventral surface of the cra At the junction of the temporal wing with the
nial part of the skull extends from the foramen body of the basisphenoid is the short alar canal.
magnum to the hard palate. Posteriorly, it pre Running in the suture between the pterygoid
sents the rounded occip ital con dyles with the process of the sphenoid bone and the pterygoid
intercondyloid notch and the median basioccipi bone is the pterygoid canal. The minute ptery
tal which extends forward between the hemi goid groove leading to the posterior opening of
spherical tym panic bullae. The muscular tuber the canal will be seen in large skulls lying dorsal
cles are low, rough, sagittally elongated ridges to and in the same direction as the muscular
of the basioccipital bone which articulate with process of the temporal bone. The anterior open
the medial surfaces of the bullae. Between the ing of the canal is in the posterior part of the
bullae and the occipital condyle is the ventral pterygopalatine fossa in the vicinity of the sep
condyloid fossa, in which opens the small circu tum between the orbital fissure and the optic
lar hypoglossal foram en . Between this small canal. It conducts the nerve of the pterygoid
round opening and the tympanic bulla (in the canal.
petro-occipital suture) is the obliquely placed, Facial part. The ventral surface of the facial
oblong petrobasilar fissure, or foram en lacerum part of the skull is largely formed by the hori
caudale, into which open the jugular and poste zontal parts of the palatine, maxillary, and in
rior carotid foramina. Fused to the posterior sur cisive bones, which form the hard palate. Lat
face of the bulla is the jugular process. Immedi eral to the hard palate on each side lie the teeth
ately anterior to the bulla and guarded ventrally in their alveoli. There are three alveoli for each
by the sharp-pointed muscular process of the of the last three cheek teeth, two for each of the
temporal bone is the osseous auditory tube. The next two, anteriorly, and one for the first cheek
external carotid foram en lies medial to the osse tooth. The largest alveolus is at the anterior end
ous auditory tube and lateral to the anterior part of the maxilla, for the canine tooth. At the ante
of the basioccipital, where it is flanked by small rior end of the hard palate, in the incisive bones,
bony processes from the tympanic bulla. The are the six incisor teeth in individual alveoli. In
largest foramen of this region is the oval fora the puppy skull only nine alveoli are present in
men, which lies medial to the m andibular fossa. each maxilla. There is one for the canine tooth,
The mandibular fossa is the smooth articular two for the first cheek tooth, and three for each
area on the transverse posterior part of the zygo of the last two deciduous molar teeth. The first
matic arch. The minute opening medial to the permanent premolar has no deciduous prede
retroglenoid process is the p etrotym pan ic fis cessor. The medial alveoli for the last three
sure, through which passes the chorda tympani cheek teeth diverge from the lateral ones, and
nerve. Posterior dislocation of the mandible, the lateral alveoli of the shearing tooth diverge
which articulates in the mandibular fossa, is pre from each other.
vented by the curved, spadelike retroglenoid The features of the hard palate vary with age.
process. The posterior surface of this process The palatine sulcus extends to the palatine fis
contains a groove which helps to form the retro sure only in adult skulls. In old skulls, transverse
glenoid foram en. ridges and depressions may be present on the
The basipharyngeal canal of Jayne (1898), or hard palate. The m ajor palatine foram in a me
the ch oan al region, is the osseous part of the dial to the carnassial teeth lie anterior to the
nasal pharynx which extends from the choanae to minor palatine foram ina. The minor palatine
the posterior borders of the pterygoid bones. It foramina are usually two in number, located
is twice as long as it is wide, and its width ap close together ventral to the palatine canal. The
proximates its depth. The palatine and ptery major palatine vessels and a nerve leave the pal
goid bones form its lateral walls and part of the atine foramina, run forward in the palatine sul
roof. The median portion of its roof is formed by cus, and supply the hard palate and adjacent
the vomer, presphenoid, and basisphenoid. In soft structures. The posterior border of the hard
young skulls a small space exists between the palate forms the choanal border, and the me
presphenoid and vomer, which is later closed dian eminence, or posterior nasal spine, may be
by a posterior growth of the vomer. In the liv inconspicuous. The lateral posterior part of the
ing animal the soft palate completes the basi hard palate presents a distinct notch, which fol
pharyngeal canal and forms a tube, the nasal lows the palatomaxillary suture and is located
T he Sk u ll 43
sB alatine fissure.
P a l a t i n e process o f m a x i l l a
P a la t in e sulcus
A l v e o l i of f o u r t h p r e m o l a r t oot h
M a j o r p a l a t i n e f oramen \
P ost er i or nasal s p i n e of p a l o t i n e
M i n o r p a l a t i n e for amen%
A l v e o l i of f i r s t m o l a r tooth
Frontal foramer\ s P t e r y g o i d p r o c e s s of m a x i l l a
D p t i c canal ^ Zyc/ omat i c p r o c e s s of f r o n t a l
a l a r foramen^ _
X'
A n t e r io r .Post, foramen, pt er ygoi d c a n a l .
..Posterior a l a r f or amen s ^ Sp i n o u s f o r a m e n
Dva! foromen-^[ _M u s c u l a r p r o c e s s
Prom
moonn t o rry l u n n l n r fnrnmpn
i i
V e n t r a l c o n d y l o i d fossa i i J u g u l a r process
i i
Condijloid process1 / 'Hupoalossal for a m e n
F ig . 1-34. Skull, ventral aspect. (Right tympanic bulla removed. Left fourth premolar and left first molar removed.)
44 Chapter 1. T he Sk el et a l Sy st em
between the palatine bone and the pterygoid rior ends of the upper and the lower jaw, each
process of the maxilla. The minor palatine ves of which bears six incisor teeth. Its most promi
sels pass through it. The sagittal parts of the pal nent feature is the nearly circular nasal or piri
atine bones and the pterygoid bones project form aperture.
ventrally to a frontal plane through the hard pal
ate. The oval palatin e fissures between the ca
nine teeth are separated by the palatine proc C a v it ie s of th e Skull
esses of the premaxillary bones. Through them
the palatine vessels anastomose with the infra Cranial Cavity
orbital and nasal vessels. On the midline the two
halves of the hard palate join to form the p ala The cranial cavity (cavum cranii) (Figs. 1-35,
tine suture. On the premaxillary part of this su 1-36) contains the brain, with its coverings and
ture is located the small ventral opening of the vessels. Its capacity varies more with body size
incisive canal. than with head shape. The smallest crania have
capacities of about 40 cc., and are known as mi-
Posterior Surface of the Skull crocephalic; the largest have capacities of ap
proximately 140 cc., and are known as mega-
The posterior surface of the skull (planum cephalic. The boundaries of the cranial cavity
nuchale) is three-sided and irregular. It is may be considered as the roof, base, posterior
formed laterally by the exoccipitals, with their wall, anterior wall, and the side walls. The roof
condyles and jugular processes, dorsally by the of the skull (cranial vault or skull cap) is the cal-
supraoccipital, and midventrally by the basioc varium. It is formed by the parietal and frontal
cipital, with its intracondyloid incisure. The lat bones, although posteriorly the interparietal
eral sides of the posterior surface are separated process of the occipital bone contributes to its
from the temporal fossae by the crestlike dorsal formation. The anterior two-thirds of the base
nuchal line. About 1 cm. ventral to the mid-dor- of the cranium is formed by the sphenoid bones
sal point of the dorsal nuchal line and running and the posterior third by the basioccipital. The
from side to side is the ventral n uchal line. The posterior wall is formed by the occipitals, and
external occipital protuberance is the mid-dorsal the anterior wall by the cribriform plate of the
posterior end of the external sagittal crest. Lat ethmoid. The lateral wall on each side is formed
eral to the external occipital protuberance is a by the temporal, parietal, and frontal bones, al
rough area for the attachment of the m. semi- though ventrally the sphenoid and posteriorly
spinalis capitis. Between the external occipital the occipital bones contribute to its formation.
protuberance and the foramen magnum is the The base of the cranial cavity is divided into an
external occipital crest, which is frequently terior, middle, and posterior cranial fossae. The
bulged in its middle by the verm iform fossa. interior of the cranial cavity contains smooth
The foram en m agnum is the large, frequently digital impressions bounded by irregular eleva
asymmetrical, ventral, median opening for the tions, the cerebral and cerebellar juga. These
spinal cord and associated structures. Lateral markings are formed by the gyri and sulci, re
to the foramen magnum are the smooth, con spectively, of the brain.
vex occipital condyles. Each is separated from The anterior cranial fossa (fossa cranii an-
the jugular process by the ventral condyloid terius) supports the olfactory bulbs and tracts
fossa, in the anterior part of which is the hypo and the remaining parts of the frontal lobes of
glossal foramen. In the young skull the occipi the brain. It lies at a higher level and is much
tomastoid suture is present lateral to the jugu narrower than the part of the cranial floor pos
lar process. This suture fails to close dorsally, terior to it. It is continued anteriorly by the
forming the supram astoid foram en . The mas deep ethmoidal fossa. Only in old dogs is a
toid process is no more than the mastoid part crista galii present, and this vertical median
of the temporal bone dorsal to the stylomastoid crest is confined usually to the ventral half of
foram en, which is anterior to the jugular proc the ethmoidal fossa. In most specimens a line
ess and dorsal to the tympanic bulla. indicates the posterior edge of the perpendicular
plate of the ethmoid, which takes the place of
Apex of the Skull the crest. The cribriform p late is so deeply in
dented that its lateral walls are located more
The apex of the skull is formed by the ante nearly in sagittal planes than in a transverse
T he Sku ll 45
plane. It is perforated by the numerous cribri ventral petrosal venous sinus. The internal
form foram ina. At the junction of the ethmoid carotid fo ra m en is located anterolateral to the
with the frontal and sphenoid bones are located petrobasilar foramen, which it resembles in size
the double ethmoidal foramina. The transversely and shape. It lies directly under the apex of the
concave sphenoidal yoke of the presphenoid petrous temporal bone, where it is located ven
bone forms most of the floor of this fossa. The tral to the canal for the trigeminal nerve and
right and left optic canals diverge as they run dorsal to the external carotid foramen. It is the
rostrally through the cranial floor. The sulcus internal anterior opening of the carotid canal,
chiasmatis lies posterior to the optic canals, which conducts the internal carotid artery and a
and in young specimens its middle part forms vein. The canal fo r the trigeminal nerve runs al
a transverse groove connecting the internal most directly anteriorly and is nearly horizontal
portions of the two canals. The shelf of bone in direction as it perforates the petrous temporal
located above the anterior part of the sulcus bone.
chiasmatis is the orbitosphenoidal crest which Posterolateral to the canal for the trigeminal
bears the anterior clinoid processes. nerve is located the internal acoustic pore, which
The middle cranial fossa (fossa cranii media) leads into the short internal acoustic meatus.
is situated at a lower level than the anterior Dorsolateral to the pore is the variably developed
fossa. The body of the basisphenoid forms its cerebellar fossa. In the petro-occipital suture, at
floor. Posteriorly, it is limited by the antero- the posteroventral angle of the pyramid, is the
dorsal surfaces of the pyramids, which end jugular foram en. Posteromedial to this opening
medially in the sharp petrosal crests. The or is the small internal opening of the hypoglossal
bital fissures are large, diverging openings canal. Located within the medial part of the
on the lateral sides of the tuberculum sellae. lateral occipital bone is the large condyloid
Posterior and slightly lateral to the orbital fis canal, for the transmission of the condyloid vein.
sures are the round foram ina, which open into The anterior part of the canal frequendy bends
the alar canals. Posterolateral to the round foram dorsally, so that its opening faces anterodorsally.
ina are the larger oval foram ina. The complex The cranial fossae form the floor of the
of structures which surround the hypophysis is cranial cavity. The remaining portion of the
called the sella turcica. It consists of an anterior cranium is marked internally by smooth de
pommel, or tuberculum sellae, and a posterior pressions and elevations which are formed by
elevation, or dorsum sellae. The posterior clinoid the gyri and sulci of the brain. The impressions
processes, which are irregular in outline, form are called digital impressions. The elevations are
the sides of the flat but irregular top of the dor formed by the sulci of the cerebellum as well as
sum sellae. The hypophyseal fossa, in which the those of the cerebrum, but unless specifically
pituitary body lies, is a shallow oval excavation indicated by the name cerebellar juga, they are
of the basisphenoid bone, located between the all known by the more comprehensive term,
tuberculum sellae and the dorsum sellae. The cerebral juga. The vascular groove (sulcus vascu-
temporal lobes of the brain largely fill the lateral losus arteriae meningeae mediae), for the middle
parts of the middle cranial fossa. meningeal artery and vein, begins at the oval
The caudal cranial fossa (fossa cranii caudalis) foramen and ramifies dorsally. Its branches vary
is formed by the dorsal surface of the basioccipi greatly in their course and tortuosity, and in old
tal bone and is located posterior to the middle specimens parts of the groove may be bridged
cranial fossa. It is bounded in front by the by bone.
dorsum sellae; posteriorly, it ends at the foram en The edges of the petrosal crests and the ten
magnum. Its dorsal surface is concave where the torium ossium serve for the attachment of the
pons, medulla oblongata, and vessels rest upon tentorium cerebelli, which separates the cere
it. Laterally, a considerable cleft exists between brum from the cerebellum. Extending from the
the apical part of the petrous temporal and the tentorium ossium to the suture, between the
basioccipital bones. At the posteromedial part of petrosum and squamosum, is the groove of the
this cleft is located the petro-occipital foram en transverse sinus. This transverse groove connects
(foramen petro-occipitalis), by means of which the transverse canal in the internal occipital
the petrobasilar canal opens anteriorly. This protuberance with the tem poral canal, which
foramen is continued toward the dorsum sellae leads to the outside by the retroglenoid foramen.
by a groove. The foramen and canal conduct the The foram en im par is usually a single opening,
46 Chapter 1. T he Sk el et a l System
T e n t o r i u m osseum f i r o o v e for m i d m e ni n g e a l a.
Transverse g r o o v e , Lat. p a r t o f f r o n t a l s i nu s
i
F oramen i mpar^ 1 I nt er nal tabl e of frontal bone
T r a n s v e r s e canal, nus
C e r e b & l l a r fossa
Supramastaid
foramen l oturbinates
Condul oi d canal .
A n t e r i o r o p e n i n g " ~ ik
P o s t e r i o r opening
Jugular foram en-'I
H y p o g l o s s a l f o r ame n ' '
Dorsum s e l l a e 1 | \ 'Opf/p c a n a l
. \
1 \Orbital fissure
Foramen oval e
1Foramen rotundum.
F ig . 1-35. Sagittal section of skull. The position of the vomer is indicated by a dotted line.
Roman numerals indicate endoturbinates.
Arabic numerals indicate ectoturbinates.
T he Skull 47
P ir ifo rm aperfure
Palatine fissu re
I n f r a o r b i ta I f o r a m e n h
Fossa f o r l a c r i m a l sac
M a x i l l a r y f o r a men
L a t p a r t of f r o n t a l s i n
Alveolar foramina
Cribriform p la ti
Dorsum s e l la e -
Canal for trig e m in a l n
C r i s t a petrosa
Int. a c o u s t i c m e a t u s
Transverse g ro o v e ''''
^ J u g u l a r f o r ame n
C ereb ellar fossax
not necessarily median in position, which is lo The dorsal nasal meatus (meatus nasi dorsalis)
cated on the anterior surface of the internal is located between the dorsal turbinate and the
occipital protuberance dorsal to the tentorium nasal bone. The m iddle nasal meatus (meatus
ossium. The small internal sagittal crest is a nasi medius) is located between the nasal and
median, low, smooth ridge which runs a short the maxilloturbinate bones. The ventral nasal
distance forward from the internal occipital meatus (meatus nasi ventralis) is located be
protuberance and provides attachment for the tween the maxilloturbinates and the dorsum of
falx cerebri. No constant sulcus for the dorsal the hard palate. The com m on n asal meatus
sagittal sinus exists. Ventral to the internal oc (meatus nasi communis) is the median lon
cipital protuberance is the vermiform impression gitudinal space located between the turbinate
for the vermis of the cerebellum. The divided bones and the nasal septum.
internal occipital crest flanks it. The nasopharyngeal meatus (meatus naso-
pharyngeus) is the air passage extending from
Nasal Cavity the posterior ends of the middle, ventral, and
common nasal meatuses to the choana. In the
The nasal cavity (cavum nasi) is the facial part fresh state, it is continued by the nasal pharynx
of the respiratory tract. It is composed of two or by the basipharyngeal canal in the skull. It is
symmetrical halves, the n asal fo s s a e (fossae bounded by the sagittal part of the vomer me
nasales), which are separated from each other by dially and by the maxillary and palatine bones
the nasal septum (septum nasi). This median laterally and ventrally. The dorsal part is
partition is formed anteriorly by the septal carti bounded by the floor plate of the ethmoid bone.
lage, and posteriorly by the septal processes of The entire mass of bony scrolls of the maxillotur
the frontal and nasal bones, the perpendicular binates are so formed that numerous ventropos-
plate of the ethmoid, and the sagittal portion of teriorly directed air passages exist. The posterior
the vomer. The single osseous anterior nasal portion of the maxilloturbinates is overlapped
opening is the piriform aperture. Each nasal medially by endoturbinates II and III. Incoming
fossa is largely filled by the maxilloturbinates an air is directed by the maxilloturbinate scrolls to
teriorly and the ethmoturbinates posteriorly. ward the maxillary sinus and the nasopharyngeal
The longest ethmoturbinate is endoturbinate I. meatus.
The nasoturbinate (see Fig. 1-21) is a curved The ethm oidal labyrinth (see Figs. 1-17, 1-
shelf of bone which protrudes medially from the 18) forms the scrolls which lie largely in the nasal
nasoturbinate crest into the dorsal part of the fundus. Each ethmoidal labyrinth is composed
nasal fossa. It separates the relatively large un of four medially lying endoturbinates and six
obstructed dorsal nasal m eatus from the middle smaller, laterally lying ectoturbinates. The ecto
nasal meatus which is located between the naso turbinates are interdigitated between the basal
turbinate and the maxilloturbinate bones. laminae of the endoturbinates. The endoturbi
The maxilloturbinates (see Fig. 1-21) pro nates attach posteriorly to the cribriform plate.
trude into the anterior part of the nasal fossa By means of basal laminae both the endoturbi
from a single leaf of attachment, the maxillotur nates and ectoturbinates attach to the external
binate crest. The basal lamina of the maxillotur lam ina of the ethmoid bone. The external lam
binates curves inward and downward from this ina is a thin, imperfect, papyraceous osseous
crest. From the convex surface of the lamina coating of the ethmoidal labyrinth. It is fused
arise five or six accessory leaves which divide largely to adjacent bones around its periphery.
several times, forming a complicated but rela The most ventroposterior extension of the eth
tively constant pattern of delicate bony scrolls. moturbinates is endoturbinate IV, which fills the
The greatest number of subdivisions leave the sphen oidal fo ss a so that what would otherwise
first accessory leaf. Subsequent accessory leaves be a sphenoidal sinus is largely obliterated. The
have fewer subdivisions. The free ends of the most dorsoposterior extensions of the ethmotur
bony plates are flattened near the floor of the binates are the first two ectoturbinates, which
nasal septum and nasoturbinate. invade the fron tal sinus, completely lining the
In each nasal fossa the shelflike nasal turbi medial part and also, to some extent, the anterior
nate and the elaborate oblique scrolls of the portion of the lateral part. A posteroventrally
maxilloturbinates divide the nasal cavity into running canal exists between the maxilloturbi
four primary passages, known as meatuses. nates and ethmoturbinates. This canal lies
T he V ertebral C olum n 49
against the maxilla and directs incoming air past of ectoturbinate 3, the posterior extremity of
the opening of the maxillary sinus into the naso which flares peripherally and ends as a delicate
pharyngeal meatus. The area occupied by the free end closely applied to the heavier frontal
ethmoturbinates is the fundus o f the nasal fossa bone. Not only is the ectoturbinate covered by
(fundus nasi). It is separated from the nasopha mucosa, but the whole sinus is also lined with
ryngeal meatus by the floor plate of the ethmoid mucosa, because it is an open cavity in free
bone and the wings of the vomer. Anteriorly, the communication with the nasal fossa in and
floor of each nasal fossa contains the oblong pal around ectoturbinate 3. The m edial part of the
atine fissure. The nasolacrimal canal arises from frontal sinus is more irregular and subject to
the anterior part of the orbit and courses to the greater variations in size than is the lateral. The
concavity of the maxilloturbinate crest, where it inner table of the frontal bone here is largely de
opens. Its medial wall may be deficient in part. ficient, so that the ethmoturbinates completely
The sphenopalatine foram en is an opening into invade this compartment. Ectoturbinates 1 and
the nasopharyngeal meatus from the anterior 2 are the scrolls which are located in this com
part of the pterygopalatine fossa. partment. They are usually separated by a lat
eral shelf of bone, to which ectoturbinate 2 is at
tached in such a way that ectoturbinate 1 lies
Paranasal Sinuses
anterior to 2, although many variations occur.
The size and form of the frontal sinus are de
The maxillary sinus or recess (Fig. 1-37) is a
pendent on skull form and age. In heavily mus
large, lateral diverticulum of the nasal fossa. The
cled, dolichocephalic breeds, the lateral com
opening into the sinus, or aditus nasomaxillaris,
partment is particularly large. In brachycephalic
usually lies in a transverse plane through the an
breeds, the medial compartment is much re
terior roots of the upper shearing tooth; the sinus
duced in size or absent, and the lateral part is
runs posteriorly to a similar plane through the
small. All paranasal sinuses enlarge with age,
last cheek tooth. The posterior part of the sinus
and only the largest definitive diverticula are
forms a rounded fundus by a convergence of its
present at birth.
walls. The medial wall of the maxillary sinus is
formed by the lateral lamina of the ethmoid
bone, and the lateral wall is formed by the max THE VERTEBRAL COLUMN
illary, palatine, and lacrimal bones. The medial
and lateral walls meet dorsally and ventrally at General
acute angles. The osseous lateral wall of the
sinus may be deficient, as a result of which there The vertebral column (columna vertebralis),
may be a communication with the pterygopala or spine (see Fig. 1-1), consists of approximately
tine fossa. Although this excavation in the max 50 irregular bones, the vertebrae. (The three sep
illa is known as the maxillary sinus, it would be arate hemal arches to be described later are not
more appropriate, in the dog, to call it the maxil included in this number.) The vertebrae are ar
lary recess, because the circumference of its ranged in five groups: cervical, thoracic, lumbar,
opening into the nasal fossa is as great as that of sacral, and coccygeal. The first letter (or abbre
the main portion of the cavity. viation) of the word designating each group, fol
The frontal sinus (Fig. 1-37) is located chiefly lowed by a digit designating the number of ver
between the outer and inner tables of the frontal tebrae in the specific group, constitutes the verte
bone. It varies more in size than any other cavity bral formula. That of the dog is C 7T 13L 7S3Cy20.
of the skull. It is divided into lateral and medial The number 20 is arbitrary for the coccygeal
parts. The lateral part occupies the whole trun vertebrae; many dogs have less, and a few have
cated enlargement of the frontal bone which more. All vertebrae except the sacral vertebrae
forms the supraorbital process. It may be partly remain separate and articulate with contiguous
divided by osseous septa which extend into the vertebrae in forming movable joints. The three
cavity from its periphery. Anteriorly an uneven sacral vertebrae are fused to form a single bone,
transverse partition unites the two tables of the the sacrum (os sacrum). The vertebrae protect
frontal bone. This partition is deficient medially, the spinal cord and roots of the spinal nerves, aid
resulting in formation of the nasofrontal opening in the support of the head and the internal or
(apertura sinus frontalis) into the nasal fossa. gans, and furnish attachment for the muscles
Through the opening extends the delicate scroll governing body movements. Although the
50 Chapter 1. T he Sk el et a l Sy st em
amount of movement between any two verte of each transverse process, in the cervical region,
brae is limited, the vertebral column as a whole is the transverse foram en (foramen transver-
possesses considerable flexibility (Slijper 1946). sarium), which divides the process into dorsal
A typical vertebra consists of a body, or cen and ventral parts. The dorsal part is an intrinsic
trum; a vertebral arch, or neural arch, consist part of the transverse process. It is comparable
ing of right and left pedicles and laminae; and to the whole transverse process found in a tho
processes for muscular or articular connections, racic vertebra. The part ventral to the transverse
which include transverse, spinous, and articular foramen is serially homologous with a rib, a cos
processes. tal element which has become incorporated into
The body (corpus vertebrae) of a typical ver the transverse process. It is not unusual in the
tebra is constricted centrally. It has a slightly dog for this costal element to be free from the
convex cranial articular surface and a centrally seventh cervical vertebra on one or both sides.
depressed caudal articular surface. In life, the In such instances the separate bone is known as
intervertebral jibrocartilage or disc (discus in- a cervical rib. Paired articular processes are
tervertebralis) is located between adjacent ver present at both the cranial and the caudal sur
tebrae. Its center is composed of a pulpy nu face of a vertebra, at the junction of the root
cleus (nucleus pulposus), which bulges freely and lamina. The cranial process (processus ar-
when the confining pressure of the outer por ticularis cranialis), or prezygapophysis, faces
tion, or fibrous ring (annulus fibrosus), is re craniodorsally, whereas the caudal process
leased. The tough outer or laminar portion of (processus articularis caudalis), or postzyga-
the disc attaches firmly to adjacent vertebrae, pophysis, faces caudoventrally.
forming a formidable retaining wall for the
amorphous, gelatinous center. A pathologic-
anatomical study of disc degeneration in the dog Cervical Vertebrae
has been made by Hansen (1952).
The vertebral arch (arcus vertebralis), or neu The cervical vertebrae (vertebrae cervicales)
ral arch, consists of two pedicles (pediculi arcus (Figs. 1-38 to 1-42) are seven in number in most
vertebrae) and two lam inae (laminae arcus ver mammals. The first two, differing greatly from
tebrae). Together with the body, the arch forms each other and also from all the other vertebrae,
a short tube, the vertebral foram en (foramen can be readily recognized. The third, fourth, and
vertebrale). All the vertebral foram ina concur to fifth differ only slightly, and are difficult to dif
form the vertebral canal (canalis vertebralis). On ferentiate. The sixth and seventh cervical verte
each side the root of the vertebra extends dor brae present differences distinct enough to make
sally from the dorsolateral surface of the body, their identification possible.
presenting smooth-surfaced notches. The cranial The atlas (Fig. 1-38), or first cervical verte
vertebral notch (incisura vertebralis cranialis) is bra, is atypical in both structure and function. It
shallow; the caudal vertebral notch (incisura articulates with the skull cranially, and with the
vertebralis caudalis) is deep. When the vertebral second cervical vertebra caudally. Its chief pecu
column is articulated in the natural state, the liarities are the modified articular processes, lack
notches on either side of adjacent vertebrae, of a spinous process, and reduction of its body.
with the intervening fibrocartilage, form the As if to compensate for these deficiencies, the
right and left intervertebral foram in a (foramina lateral parts are thick and strong, forming the
intervertebralia). Through these pass the spinal lateral masses (massae laterales), which are
nerves, arteries, and veins. The dorsal part of the united by the dorsal and the ventral arch (arcus
vertebral arch is composed of right and left lam dorsalis et ventralis). The elliptical space be
inae which unite at the mid-dorsal line to form a tween the dorsal arch of the atlas and the occip
single spine, or spinous process (processus spi- ital bone is the spatium interarcuale atlanto-oc-
nosus), without leaving any trace of its paired cipitale. The shelflike transverse processes, or
origin. Most processes arise from the vertebral wings (alae atlantis), project from the lateral
arch. Each typical vertebra has, in addition to masses. Other eminences of the atlas are the dor
the single, unpaired, dorsally located spinous sal and the ventral tubercle (tuberculum dorsale
process, on either side an irregularly shaped et ventrale), located on the respective arches,
transverse process (processus transversus) which median in position and near their caudal bor
projects laterally from the region where the ders. Frequently the dorsal tubercle is bifid,
pedicle joins the vertebral body. At the root and the ventral tubercle may take the form of a
52 Chapter 1. T he Sk eleta l System
conical process. The cranial articular surface down by the transverse ligament. The cranial
(fovea articularis cranialis) consists of two coty articular surfaces of the axis are located laterally
loid cavities which sometimes meet ventrally. on the expanded cranial end of the vertebral
They articulate with the occipital condyles of body. The caudal articular processes are ventro
the skull, forming a joint of which the main lateral extensions of the vertebral arch which
movements are flexion and extension. Since the face ventrally. Through the pedicles of the ver
atlanto-occipital joint allows rather free up-and- tebra extend the short transverse canals. Two
down movement of the head, it may be remem deep fossae, separated by a median crest, mark
bered as the “yes joint.” The caudal articular the ventral surface of the body. The cranial ver
surface (fovea articularis caudalis) consists of tebral notches concur on either side with those
two shallow glenoid cavities which form a freely of the atlas to form the large intervertebral foram
movable articulation with the second cervical ina for the transmission of the second pair of
vertebra. This is sometimes spoken of as the “no cervical nerves and the intervertebral vessels.
joint,” since rotary movement of the head oc The caudal notches concur with those of the
curs at this articulation. The caudal part of the third cervical vertebra to form the third pair of
dorsal surface of the ventral arch contains the intervertebral foramina, through which pass the
odontoid fo v ea (fovea dentis), which is concave third pair of cervical nerves and the interverte
from side to side and articulates with the dens bral vessels.
of the second cervical vertebra. This articular The third, fourth, and fifth cervical vertebrae
area blends with the articular areas on the cau (Fig. 1-41) differ slightly from each other. The
dal surface of the lateral masses. Besides the spinous processes increase in length from the
large vertebral foram en, through which the third to the fifth vertebra. The laminae are par
spinal cord passes, there are two pairs of foram ticularly strong on the third cervical vertebra,
ina in the atlas. The transverse foram ina are but gradually become shorter and narrower on
short canals passing obliquely through the trans the remaining vertebrae of the series. Tuber
verse processes, or wings, of the atlas, whereas cles are present on the caudal articular proc
the intervertebral foram ina perforate the cranio- esses, decreasing in prominence from the third
dorsal part of the dorsal arch. Cranial and caudal to seventh cervical segment. The transverse
notches are located at the origin of the trans processes are two-pronged and slightly twisted
verse processes. The atlantal fo s s a e (fossae at- in such a manner that the caudal prong lies at a
lantis) are depressions ventral to the wings. In more dorsal level than the cranial. The trans
some specimens there is an intraosseous canal verse processes of the fifth cervical vertebra are
running from the atlantal fossa into the lateral the shortest. On each vertebra there is a pair of
mass. The vertebral vein and artery traverse transverse foram ina, which extend through the
the atlantal fossa. The vein extends through the pedicles of the vertebral arches.
transverse foramen caudally and anastomoses The sixth cervical vertebra possesses a higher
with the internal jugular vein in the ventral con spine than the third, fourth, or fifth, but its main
dyloid fossa rostrally. A venous branch runs dor peculiarity is the expanded platelike transverse
sally through the cranial notch in the wing and processes. These plates, which extend down
aids in forming the external vertebral venous ward and outward, represent only the caudal
plexus. The vertebral artery enters the vertebral portion of the transverse processes. The remain
canal through the first intervertebral foramen, ing cranial portion is in the form of a conical pro
after first having run through the transverse fora jection ventrolateral to the transverse foramen.
men of the atlas. In contrast to all other vertebrae, the first six
The axis (Figs. 1-39, 1-40), or second cervi cervical vertebrae are characterized by trans
cal vertebra, presents an elongated, dorsal spi verse foramina.
nous process, which is bladelike cranially and ex The seventh, or last, cervical vertebra (Fig.
panded caudally. The spinous process overhangs 1-42) lacks transverse foramina. Cervical ribs,
the cranial and caudal articular surfaces of the when these are present, articulate with the ends
vertebral body. The axis is further characterized of the single-pronged transverse processes of this
by a cranioventral peglike eminence, the dens, vertebra. The spine of this vertebra is the high
or odontoid process. This process is morphologi est of all those on the cervical vertebrae. Some
cally the caudal part of the body of the atlas, al times rib foveae appear caudoventral to the cau
though definitively it lies on the ventral floor dal vertebral notches. In these instances the
within the vertebral foramen of the atlas, held heads of the first pair of true ribs articulate here.
T he V ertebral C olum n 53
Dorsal arch , , D o r s a i - t ub e r c l e
c -Spinous p r o c e s s
Intervertebral <
imen A lar notch
Cauda! a r tic u la r
w / n g of a t l a s su r f a c e
Crania l a r t i c u l a r
surface
i ' Dens
Transverse process
i V e n t r a I tubercle F ig . 1 -3 9 . Axis, cranial aspect.
| 'Caudal a r t i c u l a r s u r f a c e
Tr ansver se f o r a m e n
F ig . 1-38. Atlas, caudal lateral aspect.
-S pinous process
S pi no us p r o c e s s
- Transverse f o r a m e n T ransverse
process
Tr a nsi/erse
~p r o c e s s
Badtj Fovea f o r f i r s t r i b
F ig. 1-41. Fifth cervical vertebra, cranial lateral aspect. F ig . 1-42. Seventh cervical vertebra, caudal aspect.
54 Chapter 1. T he Sk e l e t a l Sy stem
Thoracic Vertebrae with the first thoracic and sometimes with the
last cervical vertebra. The first ribs therefore
articulate usually with the cranial part of the
There are thirteen thoracic vertebrae (verte body of the first thoracic vertebra and with the
brae thoracicae) (Figs. 1-43 to 1-45). The first fibrocartilage which forms the joint between
nine are similar; the last four present minor dif the last cervical and the first thoracic segment.
ferences from each other and from the preced The tubercles of the ribs articulate with the trans
ing nine. The bodies of the thoracic vertebrae verse processes of the thoracic vertebrae of the
are shorter than those of the cervical or lumbar same number in all instances. The last three tho
region. Although there are about twice as many racic vertebrae usually possess only one pair of
thoracic as there are lumbar vertebrae, the tho costal fovea on their bodies, owing to a gradual
racic region is slightly less than a third longer caudal shifting of the heads of each successive
than the lumbar region. The body of each tho pair of ribs.
racic vertebra possesses a cranial and a caudal The transverse processes are short, blunt, and
costal fovea or d em ifacet (fovea costalis crani irregular. All contain fo v e a e (foveae costales
alis et caudalis) on each side as far caudally as transversales) for articulation with the tubercles
the eleventh. The body of the eleventh fre of the ribs. These foveae decrease in size and con
quently lacks the caudal demifacets, and the vexity from the first to the last thoracic vertebra.
twelfth and thirteenth thoracic vertebrae always The m am m illary processes (processus mam-
have one complete fovea on each side. The millares), or metapophyses, start at the second
foveae on the bodies of the thoracic vertebrae or third thoracic vertebra and continue as paired
are for articulation with the heads of the ribs. projections through the remaining part of the
The bodies of most of the thoracic vertebrae thoracic and through the lumbar, sacral, and
have a pair of nutrient foramina entering the coccygeal regions. They are small knoblike emi
middle of the ventral surface. All show paired nences which project dorsally from the trans
vascular foramina on the flattened dorsal sur verse processes. At the eleventh thoracic verte
face of the body. The pedicles of the vertebral bra they become associated with the cranial
arches are short. The caudal vertebral notches articular processes and continue as laterally
are deep, but the cranial notches are frequently compressed tubercles throughout the remaining
absent. The laminae give rise to a spinous proc vertebrae of the thoracic and those of the lum
ess, which is the most conspicuous feature of the bar region.
first nine thoracic vertebrae. The spine of the The accessory processes (processus accessorii),
first thoracic vertebra is more massive than or anapophyses, appear first in the mid-thoracic
the others, but is of about the same length. The region and are located on succeeding segments
massiveness gradually decreases with successive as far caudally as the fifth or sixth lumbar verte
vertebrae, but there is little change in the length bra. They leave the caudal borders of the pedi
and direction of the spines until the seventh or cles and, when well developed, form a notch
eighth thoracic is reached. The spines then be lateral to the caudal articular process which re
come progressively shorter and are inclined in ceives the cranial articular process of the verte
creasingly caudad through the ninth and tenth bra behind.
segments. The spine of the eleventh thoracic The articular processes are located at the
vertebra is nearly perpendicular to the long axis junctions of the pedicles and the laminae. The
of that bone. This vertebra, the anticlinal verte cranial pair of facets are widely separated on
bra (vertebra anticlinalis), is the transitional seg the first and second thoracic vertebrae, and
ment of the thoracolumbar region. All spines nearly confluent at the median plane on thoracic
caudal to those of the twelfth and thirteenth tho vertebrae 3 to 10. On thoracic vertebrae 11,12,
racic vertebrae are directed cranially, whereas and 13, the right and left facets face each other
the spines of all vertebrae cranial to the eleventh across the median plane and are located at the
thoracic are directed caudally. In an articulated base of the mammillary processes. The cranial
vertebral column the palpable tips of the spines articular facets, with the exception of those on
of the sixth and seventh thoracic vertebrae lie the last three thoracic vertebrae, face forward
dorsal to the cranial parts of the bodies of the and upward. The caudal facets articulate with
eighth and ninth; the tips of the spines of the the cranial ones of the vertebra behind, are simi
eighth to tenth thoracic vertebrae lie dorsal to lar in shape, and face downward and backward
the bodies of the vertebrae behind them. on thoracic vertebrae 1 to 9. The joints between
The heads of the first pair of ribs articulate thoracic vertebrae 10 to 13 are conspicuously
T he V ertebra l C olum n 55
Spmous process -
S pino us process
F ig . 1 -4 3 . First thoracic vertebra, left lateral aspect. F ig . 1 -4 4 . Sixth th oracic vertebra, cranial lateral aspect.
56 Chapter 1. T he Sk el et a l S ystem
modified, since the caudal articular facets are lo Sacral Vertebrae
cated on the lateral surfaces of dorsocaudally
projecting processes. This type of interlocking The bodies and processes of the three sacral
articulation allows flexion and extension of the vertebrae (vertebrae sacrales) fuse in the adult
caudal thoracic and the lumbar region, while to form the sacrum (os sacrum) (Figs. 1-49 to 1-
limiting sagittal movement. Foveae on the trans 52). The bulk of this four-sided, wedge-shaped
verse processes and demifacets on the centra for complex lies between the ilia and articulates
articulation with the ribs characterize the thora with them. The body of the first segment is
cic vertebrae. larger than the bodies of the other two segments
combined. The three are united to form an
Lumbar Vertebrae arched, bony mass with a concave ventral, or
pelvic surface, a feature of obstetrical impor
The lumbar vertebrae (vertebrae lumbales) tance.
(Figs. 1-46 to 1-48), seven in number, have The dorsal surface (facies dorsalis) (Fig. 1-
longer bodies than do the thoracic vertebrae. 50) presents the m edian sacral crest (crista
They gradually increase in width throughout the sacralis mediana), which represents the fusion of
series, and in length through the first five or six the three spinous processes. Two indentations on
segments. The body of the seventh lumbar ver the crest indicate the areas of fusion. The dorsal
tebra is approximately the same length as the surface also bears two pairs of dorsal sacral fo
first. The ventral foramina of each body are not ramina (foramina sacralia dorsalia), which trans
always paired or present. The dorsal foramina mit the dorsal divisions of the sacral nerves and
are paired and resemble those of the thoracic vessels. Medial to these processes are low projec
vertebrae. Although longer and more massive, tions representing the fused mammillo-articular
the pedicles and laminae of the lumbar vertebrae processes of adjacent segments. In some speci
resemble those of typical vertebrae of the other mens the three mammillo-articular processes on
regions. each side are united by intervening ridges. The
The spinous processes are highest and most aggregate of the processes and the connecting
massive in the mid-lumbar region. The spines ridges then forms the intermediate sacral crest
are about half as long, and the dorsal borders (crista sacralis intermedia). The caudal articular
are approximately twice as wide as those of the processes are small and articulate with the first
vertebrae at the cranial end of the thoracic re coccygeal vertebra. The cranial articular proc
gion. esses are large, face dorsomedially, and form
The transverse processes are directed crani- joints with the seventh lumbar vertebra.
ally and slightly ventrally. They are longest in The pelvic surface (facies pelvina) (Fig. 1-51)
the mid-lumbar region. In emaciated animals of the sacrum is variable in its degree of con
the broad extremities of the transverse proc cavity. During the first six postnatal months two
esses can be palpated. intervertebral fibrocartilages mark the separa
The accessory processes are well developed tion of the vertebral bodies. These persist in the
on the first three or four lumbar vertebrae, and adult as two transverse lines (lineae transversae).
absent on the fifth or sixth. They overlie the Two pairs of pelvic sacral foram ina (foramina
caudal vertebral notches and extend caudad sacralia pelvina), situated just lateral to the fused
lateral to the articular processes of the succeed sacral bodies, are larger than the corresponding
ing vertebrae. dorsal foramina. In addition to blood vessels,
The articular processes lie mainly in sagittal they transmit the ventral branches of the first
planes. The caudal processes lie between the two sacral nerves. Lateral to the pelvic sacral
cranial processes of succeeding vertebrae and foramina are the fused transverse processes.
restrict lateral flexion. All cranial articular proc Those of the first and part of the second segment
esses bear mammillary processes. are greatly enlarged and modified for articula
There are 20 vertebrae in the thoracolumbar tion with the ilium. The transverse processes of
region. This number is quite constant. Iwanoff the third segment and part of the second form
(1935) found only one specimen out of 300 with the narrow, thin lateral sacral crest (crista sac
21 thoracolumbar vertebrae; all of the remain ralis lateralis), which terminates caudally in a
ing had 20. Among the specimens he studied the flattened, pointed process, the caudolateral
last lumbar segment was sacralized (fused to the angle. This angle frequently articulates with the
sacrum) in three, and the first sacral vertebra adjacent transverse process of the first coccygeal
was free in two. vertebra.
■ Spi n ou s process
in ous process
Mam m illary process
Cr a ni a l o r t i c u l a r s u r f a c e
Ca u d a l a r t i c u i o r
i
-process - M a m m i l l a r y pr ocess
CraniaI a r t i c u l a r
■surface
- A c c es s a r y
Cranial a r t i c u l a r ' p r o cess Caud, a r t i c --Caudal a rtic u la r
process surface process
- V e r t e b r al
foramen Dorsal foram ina
F ig . 1-47. F if th lu m b a r v e r te b ra , c a u d a l la te ra l a s p e c t.
Vertebral
Mammi l l o- ar t i cul ar
Caudal foramen D orsal sacroi V processes
articu foram ina (Intermediate
surf a sacral crest)
W in S pi nous
process
A rticu l a r -
surface
Transverse
T r ans v e r s e p r o c e s s Body process
Caudal a r t i c u l a r process
F ig . 1 -4 8 . Seventh lumbar vertebra, caudal aspect. F ig . 1-49. Sacrum, caudal lateral aspect.
Cronial articu la r
surface Sacra! canal
Promontoru
-W iny
Pel v ic
——J-? sacral
Mammillo - a r t i c u l a r
S ' ' f o r a mina
processes
Transverse
processes
(Lot sacral crest)
Caudal a r t i c u l a r — Apex
process ^ Spino us processes
(Median s a c r a l crest) Ca u d al a r t i c u l a r s u r f ace
F ig . 1-50. Sacrum, dorsal aspect. F ig . 1 -5 1 . Sacrum, ventral aspect.
57
58 Chapter 1. T he Sk e l e t a l System
M am m illo-articular p r a n i a l a r t i c u l a r process
processes Median -s ac ra l crest Mammillary process
Dorsa I surface
Tro n sv e rs e
Pranial a r tic u la r process
process
1C a u d . a r t i c u l a r p r o c e s s
-Caudal arfic.
process F ig . 1 -5 3 . F irs t c o c c y g e a l v e rte b ra , dorsal asp ect.
T r an s v e r se proc.
\ Lat. s a c r a l c r e s t
A u ric u la r surface xPelvic surface
F ig . 1-52. Sacrum and first coccygeal vertebra, lateral aspect.
T he V ertebra l C olum n 59
Body
F ig . 1 -5 5 . Fou rth coccygeal vertebra, cranial aspect.
16
Cr an. a r t i c u l a r process
uli
Mammillany
process 19
Caudal
a r t i c u l a r p r o c es s '
s
/
Transverse process' do r s al L a te ra l
F ig . 1-57. Sixth coccygeal vertebra, dorsal and lateral aspects. F ig . 1 -5 8 . Representative coccygeal vertebrae.
60 Chapter 1. T he Sk eleta l System
processes project from the caudal border of the powerful abdominal muscles, which act for this
arch and are frequently asymmetrical. They purpose as a complete elastic apron.
gradually disappear in a craniocaudal sequence, In the fetus the vertebral column is uniformly
so that at the twelfth coccygeal vertebra both dorsally arched from the head to the tip of the
caudal and cranial articular processes are no tail. In the adult standing position the head is
longer present. The spinous processes are small elevated, resulting in a secondary cervical curva
and disappear early in the series—at about the ture, which extends the joints between the
seventh coccygeal vertebra. The first four or five caudal cervical vertebrae. It is interesting to
pairs of transverse processes are well developed note that the greatest movement of the vertebral
and typical. Caudal to the fifth coccygeal verte column takes place near one or both ends of the
bra they are reduced in size, and they disappear several regions into which it is divided: at both
at about the fifteenth segment. ends of the cervical region, near the caudal end
H em al arches (arcus hemales) (Fig. 1-55) are of the thoracic region, at the lumbosacral junc
present as separate bones which articulate with tion, and in the cranial part of the coccygeal
the ventral surfaces of the caudal ends of the region.
bodies of the fourth, fifth, and sixth coccygeal The total length of a freshly isolated vertebral
vertebrae. They slope caudally and are shaped column of a shepherd-type, medium-propor-
like a V or Y. In life, they protect the me tioned mongrel dog weighing 45 pounds was
dian coccygeal artery, which passes through found to be 109 cm. The lengths of the various
them. Caudal to the hemal arches, and in corre regions as measured along the ventral surface of
sponding positions on succeeding vertebrae, are the articulated vertebral column are shown in
the paired hem al processes (processus hemales). Table 4.
Hemal processes are the last processes to disap
pear, and remnants of them can still be identified Table 4. Length of Various Regions of a Freshly
as far caudally as the seventeenth or eighteenth Isolated Vertebral Column
coccygeal vertebra. W IT H W IT H O U T
IN T E R V E R T E B R A L IN T E R V E R T E B R A L
R E G IO N F IB R O C A R T IL A G E S F IB R O C A R T IL A G E S
the width increases, so that the canal becomes vertebrae and the intervening fibrocartilage.
transversely oval. The shape of the canal does The articular areas, corresponding to those of
not grossly change in the last two lumbar verte the vertebrae with which they articulate, are of
brae, where it is larger than in any other verte about equal size and convex, and face cranially
bra caudal to the first thoracic. The lumbar en (facies articularis capitis costae cranialis) and
largement of the vertebral canal accommodates caudally (facies articularis capitis costae cau
the lumbosacral enlargement of the spinal cord. dalis). At the eleventh or twelfth thoracic ver
Possibly the small lumbar subarachnoid cistern tebra the caudal pair of demifacets or articular
and epidural fat contribute to this enlargement, areas on the head of the rib disappear, as the
as the spinal cord usually ends opposite the fibro- last two or three ribs articulate only with their
cartilage between the last two lumbar vertebrae. corresponding vertebrae. The heads of these ribs
are modified accordingly, and each lacks the
THE R IB S crest (crista capitis costae), or transverse ridge,
which separates the two articular areas when
The ribs (costae) (Figs. 1-59, 1-60) form all of they are present. The tubercle of the rib bears an
the thoracic skeleton, except for narrow mid articular f a c e t (facies articularis tuberculi cos
dorsal and mid-ventral strips formed by the tae) for articulation with the transverse process
vertebral column and the sternum, respectively. of the vertebra of the same number. The space
There are usually 13 pairs of ribs in the dog. between the neck and tubercle of the rib and
Each rib is divided into a laterally and caudally the body of the vertebra is known as the costo
convex dorsal bony part, the os costale, and a transverse foram en (foramen costotransversa-
ventral cartilaginous part, the costal cartilage rium), which is homologous to the transverse
(cartilago costalis). The first nine ribs articulate foramen of a cervical vertebra. In the last two
with the sternum and are called the true ribs or three ribs the articular areas of the head and
(costae verae); the last four are called the f a k e that of the tubercle become confluent, but the
ribs (costae spuriae). The costal cartilages of the tubercle remains for muscular attachment.
tenth, eleventh, and twelfth ribs unite with the The body of the rib (corpus costae), in general,
cartilage of the rib above to form the costal arch is cylindrical and slightly enlarged at the costo
(arcus costalis) on each side. Since the cartilages chondral junction. The third, fourth, and fifth
of the last (thirteenth) pair of ribs end freely in ribs show some lateral compression of the distal
the musculature, these ribs are known as floating halves of the bony parts. In the large breeds the
ribs (costae fluctuantes). The ninth ribs are the ribs are flatter than they are in the small breeds.
longest, with the longest costal cartilages. Pass In all breeds the vertebral portions of the ribs
ing both caudally and cranially from the ninth are slightly thicker from side to side than they
rib, both the bony and the cartilaginous parts of are from front to back. The angle (angulus cos
the other ribs become progressively shorter. The tae) is an indistinct lateral eminence about 2 cm.
costochondral junctions, or symphyses, of the distal to the tubercle. The costal groove (sulcus
third through eighth ribs lie nearly in the same costae) on the inner surface, for the intercostal
horizontal plane. Since the sternum and thoracic vessels and nerve, is not distinct on any of the
spine diverge from the thoracic inlet and the ribs.
successive ribs become progressively more lat The costal cartilage is the cartilaginous cylin
erally arched, the caudal part of the rib cage is drical distad continuation of the bony rib. It is
much more capacious than the cranial part. The smaller in diameter than the bony rib and, in ma
space between adjacent ribs is known as the in ture dogs, may be calcified. Near the costochon
tercostal space (spatium intercostale). These dral junctions the cartilages incline cranially.
spaces are two or three times as wide as the ad This is most marked in the first and twelfth ribs.
jacent ribs. The first rib articulates with the first sternebra, or
A typical rib (os costale) as exemplified by the manubrium sterni. Succeeding true rib cartilages
seventh, presents a vertebral extremity, a ster articulate with successive intersternebral carti
nal extremity, and an intermediate shaft, or lages. However, the eighth and ninth costal car
body. The vertebral extremity consists of a head tilages articulate with the cartilage between the
(caput costae), a neck (collum costae), and a tu seventh sternebra and the last sternebra, or
bercle (tuberculum costae). The head of the rib xiphoid process. The costal cartilages of the
has a wedge-shaped articular surface which artic tenth, eleventh, and twelfth ribs are long, slen
ulates with adjacent costal foveae of contiguous der rods each joined to the one above by connec-
62 Chapter 1. T he Sk el et a l System
Manubr i um of s t e r nu m
V i_ _ _ _ F i r s t r i b
~ Intersternebral cartilacje
Second s t e r n e b r a
Costal c a r t il a c je
- -Costochondral junction
------B o d y of r i b
True r i b s <
- Seventh sternebra
- '- i'------ X i p h o i d p r o c e s s
E ig h t h i n t e r c o s t a l s p a c e
' I n f r a s t e r n a l ancjle
" ^ X i p h o i d cariilacje
Fa I s e r i b s {
‘ Thirteenth rib
ruberc!e°f r l f rit>
T ,Nec^° .. d0f r u
•Heo'-
Angl ed n b '
64 Chapter 1. T he Sk eleta l S ystem
tive tissue to form the costal arch. The costal pressed, so that its width is in a vertical plane
cartilage of the thirteenth rib, shorter and more and its thickness is in a horizontal one. The first
rudimentary than those of the adjacent ribs, en and last sternebrae are specialized. The cranial
ters the musculature of the flank, in which it ter half of the first sternebra is expanded and bears
minates. lateral projections for the attachment of the first
costal cartilages. The first sternebra is longer
THE STERNUM than the others and is known as the manubrium
The sternum (see Figs. 1-59, 1-60) is an un (manubrium sterni).
paired segmental series of eight bones (sterne- The last sternebra, called the xiphoid process
brae) which forms the floor of the thorax. It is (processus xiphoideus), is wide horizontally and
slightly turned up in front and turned down be thin vertically. Its length is about three times its
hind. The consecutive sternebrae are joined by width. It is roughly rectangular, and may have
short blocks of cartilage, the interstemebral an elliptical foramen in its caudal half. A thin
cartilages (cartilago intersternebralis). The ster cartilaginous plate (cartilago xiphoidea) prolongs
nal ends of the ribs articulate with the inter- the xiphoid process caudally.
sternebral cartilages, with the exception of the The firm cartilaginous joints between the
first pair, which articulate with the first sterne sternebrae (synchondroses sternales) may ossify
bra. The sternum of the dog is laterally com in old individuals.
A PPEN D IC U LA R SK ELETO N
The development of the limbs and epiphyseal to which it is related. The clavicle of the dog
fusion in the dog has been studied by Schaeffer does not usually appear on radiographs.
(1934), Pomriaskinsky-Kobozieff and Kobozieff
(1954), Bressou et al. (1957), Hare (1961), Smith Scapula
(I960), and Smith and Allcock (1960).
The scapula (Figs. 1-62 to 1-64) is the large,
BO N ES OF THE THORACIC LIM B flat bone of the shoulder. Its highest part lies
just below the level of the free end of the spine
Each pectoral or thoracic limb (membrum of the first or second thoracic vertebra. Longi
thoracicum) consists of its half of the pectoral tudinally, it extends from a transverse plane
girdle (cingulum membri thoracici), composed cranial to the manubrium sterni to one through
of the clavicle and scapula; the arm or brachium, the body of the fourth or fifth thoracic verte
represented by the humerus; the forearm bra. Since the pectoral limb has no articulation
or antebrachium, consisting of the radius and with the axial skeleton and supports the trunk
ulna; and the forepaw, or manus. The manus by muscles only, the normal position of the
includes the wrist or carpus, with its digits, scapula may vary by the length of one vertebra.
consisting of metacarpals, phalanges, and dor In outline it forms an imperfect triangle having
sal as well as palmar sesam oid bones. two surfaces, three borders, and three angles.
The lateral surface (facies lateralis) (Fig. 1-
62) is divided into two nearly equal fossae by a
Clavicle shelf of bone, the spine o f the scapula (spina
scapulae). The spine is the most prominent fea
The clavicle (clavicula) (Fig. 1-61) is a vestig ture of the lateral surface of the bone. It begins
ial bone not articulated with the skeleton. It is at the junction of the cranial and middle thirds
located near the medial end of the clavicular of the vertebral border as a thick, low ridge,
tendon of the brachiocephalicus muscle, and which gradually becomes wider but thinner as it
rarely it may be absent. A large clavicle may be is traced distally, so that it presents definite cra
over 1 cm. long, and a third as wide. It is thin nial and caudal surfaces throughout most of its
and slightly concave both longitudinally and length, and near its distal end there is a definite
transversely. Its medial half may be twice as caudal protrusion. The free border or crest of the
wide as its lateral half. The clavicle is more spine is slightly thickened and rolled caudally in
closely united to the clavicular tendon between heavily muscled specimens. The widened trun
the m. cleidomastoideus and the m. cleido- cate distal end of the spine of the scapula is
brachialis than to the underlying axillary fascia called the acromion. Its broadened superficial
B o n es o f t h e T h o r a c ic L im b 65
Supraspinous fossa
In fra s p in o u s fossa
Caudal angle
Cranial border -
S pine -
Facies s e r r a t a
Acromion-- _
Scapular n ofch
S ca p u la r
tuberositu ----- J ,
J A r f i c u l a r a ngl e •M u s c u l a r lines
F ig u r e 1-62
-S u b sca p u la r
fossa
1— i.o cm.—1
F ig u re 1-61
I n f r a - a r t i c u i a r tuberosity^
Corocoid process
Glenoid S ca p u la r
cavity - tu berosifij
A c r o m i on - /I'
fis F ig u r e 1-63
Suprospinous fosso
Scapul ar tuberosity
Pa-f iSarrocj/
Coracoi d p r o c e s s -
Infraspinous fossa
1C a u d a l b o r d e r
Gl enoi d c a v i t y
Infra-articular tuberosity
F ig u r e 1-64
portion is subcutaneous and easily palpated in it becomes rougher and thicker, as it runs into
the living animal. A nutrient foramen is fre the vertebral border at the cranial angle.
quently present at the junction of the ventral The vertebral border (margo vertebralis),
border of the spine and the scapula proper. The sometimes called the base, extends between the
acromial part of the m. deltoideus arises from the cranial and caudal angles. In life it is capped by
acromion and extends distally. The m. omotrans- a narrow band of scapular cartilage (cartilago
versarius arises from the distal end of the spine scapulae), which represents the unossified part
adjacent to the acromion and extends cranially. of the bone. The m. rhomboideus attaches to the
The m. trapezius inserts on, and the spinous part vertebral border of the scapula.
of the m. deltoideus arises from, the whole crest The caudal border (margo caudalis) (Fig. 1-
of the spine dorsal to the origin of the m. omo- 64) is the thickest of the three borders and bears,
transversarius. (Fig. 3-43). just dorsal to the articular angle, the infraglenoid
The supraspinous fo ssa (fossa supraspinata) is tuberosity (tuberculum infraglenoidale). This
bounded by the cranial surface of the scapular tuberosity is much thicker than the border, and
spine and the adjacent lateral surface of the scap is located largely on the costal surface of the
ula. It is widest in the middle because the cranial bone. Parts of the mm. triceps-caput longum and
border of the scapula extends in an arc from the teres minor arise from the infraglenoid tuberos
scapular notch to the cranial angle. The whole ity. The ventral third of the thick caudal border
thin plate of bone which forms the supraspinous contains two muscular lines which diverge dis
fossa is sinuous, possessing at its greatest undu tally; the more cranially located line extends
lation a lateral projection involving the middle nearly to the lip of the glenoid cavity, and the
of the fossa. The m. supraspinatus arises from all more caudal one ends in the infraglenoid tuber
but the distal part of the supraspinous fossa. osity. The more cranial line and adjacent cau
The infraspinous fo ssa (fossa infraspinata) is dal border of the scapula give origin to the m.
in general triangular. Since the caudal and ver teres minor; the more caudal line and adjacent
tebral borders are thick and the spine leaves the caudal border give origin to the m. triceps-
lateral surface at nearly a right angle, this fossa caput longum. The middle third of the caudal
is well defined. The m. infraspinatus arises from border of the scapula is broad and smooth; the
the infraspinous fossa. m. subscapularis curves laterally from the me
The medial or costal surface (facies costalis) dial side and arises from it here. Approximately
(Fig. 1-63) of the scapula lies opposite the first the dorsal fourth of the caudal border is sur
five ribs and the adjacent four or five thoracic rounded by a lip in the heavily muscled breeds.
vertebrae. Two areas are recognized: a small From this part arises the m. teres major.
dorsocranial rectangular area, fa c ies serrata, The caudal angle (angulus caudalis) is obtuse
from which arises the powerful m. serratus ven as it unites the adjacent thick caudal border with
tralis, and the large remaining part of the costal the thinner, rougher, gently convex vertebral
surface, or the subscapular fo ssa (fossa subscap- border. The m. teres major arises from the cau
ularis). It is nearly flat and usually presents three dal angle and the adjacent caudal border of the
relatively straight muscular lines which converge scapula.
toward the distal end of the bone. Between the The cranial angle (angulus cranialis) imper
lines the bone is smooth, and in some places it is ceptibly unites the thin, convex cranial border
concave. The largest concavity lies opposite the to the rough, convex, thick vertebral border. No
spine. From the whole subscapular fossa, and muscles attach directly to the cranial angle.
particularly from the muscular lines, arises the The articular angle (angularis articularis)
m. subscapularis. forms the expanded distal end of the scapula.
The cranial border (margo cranialis) is thin ex Clinically, the articular angle is the most impor
cept at its extremities. Distally, it forms a con tant part of the bone, since it contains the gle
cavity, the scapular notch (incisura scapulae) noid cavity (cavitas glenoidalis), which receives
which marks the position of the constricted part the head of the humerus in forming the shoulder
of the bone. The border undulates as it reflects joint. The glenoid cavity is very shallow; its lat
the warped nature of the supraspinous fossa. In eral border is flattened, and cranially it extends
the working breeds, the cranial border forms an out on the tuber scapulae. The medial border
arc, whereas in dogs with slender extremities, forms a larger arc than does the caudal border.
the border is nearly straight. Distally, the bor The scapular or supraglenoid tuberosity (tu
der becomes smoother and thicker; proximally, ber scapulae s. tuberculum supraglenoidale) is
B on es o f t h e T h o r a c ic L im b 67
the largest tuberosity of the scapula. For the tubercle meet at about a right angle cranially.
most part it projects cranially, with a medial in The two tubercles are separated craniomedially
clination. From it arises the single tendon of the by the intertuberal groove, and caudolaterally
m. biceps brachii. The small beaklike process by the head of the humerus.
which leaves the medial side of the scapular tu The neck of the humerus (collum humeri) is
berosity is the coracoid process (processus cora- distinct only caudally and laterally. It indicates
coideus), from which the m. coracobrachialis the line along which the head and parts of the
arises. The coracoid process is a remnant of the tubercles have fused with the shaft.
coracoid bone, which is still distinct in mono- The body of the humerus (corpus humeri),
tremes. Although in dogs this osseous element is or shaft, is the long, slightly sigmoid-shaped
no longer a separate bone, it still retains its own part of the humerus which unites the two ex
center of ossification. tremities. It varies greatly in shape and size, de
pending on the breed. Usually it is laterally com
Humerus pressed, possessing medial and lateral surfaces,
and cranial and caudal borders which are not
The humerus (Figs. 1-65 to 1-67) is the bone distinct throughout their length.
of the true arm, or brachium, and is the largest The lateral surface (facies lateralis) (Fig. 1-66)
bone of the thoracic limb. Proximally it articu is marked proximally by the an coneal line (linea
lates with the scapula in forming the shoulder anconea), and a tuberosity which divides it into
joint; distally it articulates with the radius and a narrow, slightly convex area, which faces
ulna in forming the elbow joint. Developmen- craniolaterally, and a wider, smoother surface,
tally it is divided into a shaft and two extremities; which is slightly concave and faces caudolater
definitively it is divided into a head, neck, and ally. The anconeal line begins at the head of the
body. humerus caudal to the greater tubercle, and in
The head (caput humeri) is oval, being elon an uneven, cranially protruding arc extends dis
gated in a sagittal plane. The articular area it tally to the elongated deltoid tuberosity (tuberos
presents is about twice the size of that of the gle itas deltoides). On the crest, below the head, is
noid cavity of the scapula with which it articu a small enlargement for the insertion of the m.
lates. Although it is rounded in all planes, it does teres minor. The remaining distal part of the
not form a perfect arc in any plane, as the cra crest serves for the origin of the m. triceps-caput
nial part is much flatter than the caudal part. laterale. The deltoid tuberosity is the most prom
The articular area of the head is continued dis inent feature of the lateral surface of the hu
tally by the intertubercular groove (sulcus inter- merus and serves for the insertion of the m.
tubercularis), which ridges the craniomedial deltoideus. The musculospiral groove (sulcus
part of the proximal extremity of the bone. The spiralis) (Hughes and Dransfield 1953) forms the
extension of the shoulder joint capsule into the smooth, flat to convex, lateral surface of most of
groove lubricates the bicipital tendon which the humerus. It begins at the neck caudally and
lies in it (Fig. 2-9). extends laterally and finally cranially as it twists
The greater tubercle (tuberculum majus), or to the distal extremity of the bone. Although the
large craniolateral projection of the proximal ex m. brachialis lies in the whole groove, it arises
tremity of the humerus, has a smooth convex from the proximal part only. Both the proximal
summit which in most breeds extends higher and the distal part of the lateral surface incline
than the head. It serves for the total insertion of cranially. The proximal part lies between the
the m. supraspinatus and the partial insertion of crest of the major tubercle medially and the
the m. pectoralis profundus. Between the head anconeal crest laterally.
and greater tubercle are several small foramina The m edial su rface (facies medialis) is
for the transmission of veins. The relatively rounded transversely, except for a nearly flat tri
smooth facet distal to the summit of the greater angular area in its proximal fourth. Caudally, this
tubercle serves for the insertion of the m. infra area is bounded by the crest o f the lesser tuber
spinatus. The lesser tubercle (tuberculum minus) cle (crista tuberculi minoris), which ends distally
is a medially flattened enlargement of the proxi in an inconspicuous eminence, and the teres tu
mal medial part of the humerus, the convex bor berosity (tuberositas teres), which lies in the
der of which does not extend as high as the head. same transverse plane as the laterally located
To this convex border attaches the m. subscapu deltoid tuberosity. The m. coracobrachialis in
laris. Planes through the lesser and the greater serts on the crest of the lesser tubercle adjacent
G r e a t e r t u be rc le _ / t
In t e r t u b ercul ar_\
groove
C rest o f - ■H e a d
G r e a t e r tubercle
j - - A n c o n e a l crest
Teres fu beros i ftj -
- - De l t o l d
tuberosity
-M u s c u l o s piral
cj roove
G reate r tubercle
Lat. epi condyl oi d
crest N 1 Head
i
Lesser t u b e r c l e
Radial fossa- H u me r a l
j>condyle
Supratrochlear
foram en
-N u trient
foramen
L a t e r a l epi c o n d y l o i d '
crest
Olecranon fossa -
'r'afSamiS-
68
B on es of the T h o r a c ic L im b 69
to the teres tuberosity. Cranial to this insertion (Fig. 1-67) is also known as the flexor epicon
the m. triceps-caput mediale arises from the dyle. Larger than the lateral epicondyle, it gives
crest by a small aponeurosis. The mm. teres ma origin to the m. flexor digitorum superficialis
jor and the latissimus dorsi insert on the teres and the humeral heads of the mm. flexor digi
tuberosity. The medial surface of the humerus is torum profundus, flexor carpi ulnaris, and flexor
loosely covered by the m. biceps brachii. carpi radialis. The proximal end of the medial
The cranial surface (facies cranialis) of the hu ligament of the elbow joint attaches to the artic
merus begins proximally at the crest of the ular margin and adjacent surface of the medial
greater tubercle. This crest swings laterally just epicondyle.
medial to the deltoid tuberosity, where it The olecranon fo ssa (fossa olecrani) is a deep
reaches the cranial edge of the spiral groove. excavation of the caudal part of the distal ex
The entire m. pectoralis superficialis attaches to tremity of the humerus. It receives the anconeal
the crest of the greater tubercle, and a portion of process (processus anconeus) of the ulna when
the m. pectoralis profundus attaches to its prox the elbow joint is extended. The olecranon fossa,
imal part (Figs. 3-47 and 3-48). in life, is covered by the m. anconeus, which
The caudal surface (facies caudalis) (Fig. 1- arises from its margin. Opposite the olecranon
67) begins at the neck of the humerus where the fossa is the radial fo ssa (fossa radialis), which is
m. triceps-caput accessorium arises. As a trans also called the coronoid fo ssa (fossa coronoidea)
versely rounded margin, it extends to the distal by many veterinary anatomists (Sisson and
fourth of the bone, where it is continued by the Grossman 1953, Baum and Zietzschmann 1936,
lateral epicondyloid crest. The caudal border is and Hughes and Dransfield 1953). The dog has
perforated below its middle by the distally di no coronoid fossa, since only the head of the ra
rected nutrient foram en. dius enters this depression when the elbow joint
The sagittally rounded distal end of the hu is flexed, and not the coronoid process of the
merus may be divided into a small, lateral articu ulna. The radial and olecranon fossae commu
lar area, the capitulum humeri, for articulation nicate with each other by means of the supra
with the head of the radius, and the trochlea hu trochlear foram en (foramen supratrochleare).
meri, a much larger, medially located pulley The foramen may be absent when the humerus
shaped part which extends proximally into the is small.
adjacent fossae (Fig. 1-65). The trochlea articu
lates extensively with the trochlear notch of the Radius
ulna in forming one of the most stable hinge
joints in the body. The distal end of the hu The radius (Figs. 1-68, 1-69) is the main
merus, including its articular areas and the adja weight-supporting bone of the forearm; it is
cent fossae, may be regarded as the humeral shorter than the ulna, which parallels it and
condyle (condylus humeri) (Fig. 1-66). serves primarily for muscle attachment. The ra
The lateral epicondyle (epicondylus lateralis) dius articulates with the humerus proximally in
(Figs. 1-65,1-67) is the enlarged distolateral end forming the elbow joint and with the carpal
of the humerus. It lies caudoproximal to the lat bones distally in forming the main joint of the
eral articular margin of the capitulum. It gives carpus. It also articulates with the ulna proxi
origin to the mm. extensor digitorum communis, mally by its caudal surface and distally by its
extensor digitorum lateralis, and extensor carpi lateral border. The radius, like the humerus, is
ulnaris. Functionally, it is known as the extensor divided into proximal and distal extremities,
epicondyle of the humerus. The proximal end of with an intervening shaft or body.
the lateral ligament of the elbow joint attaches The head (caput radii) is irregularly oval in
to the articular margin and adjacent surface of outline as it extends transversely across the prox
the lateral epicondyle. The lateral epicondyloid imal end of the bone. It is concave, articulates
crest (crista epicondylica lateralis) extends prox with the capitulum of the humerus, and bears
imally from the lateral epicondyle. It is a thick practically all the weight transmitted from the
rounded crest which ends by blending with the arm to the forearm. The articular circumference
caudal border at the beginning of the distal (circumferentia articularis s. articularis ulnaris
fourth of the humerus. The m. brachioradialis radii proximalis) is a caudal, smooth, osseous
arises from the proximal part of the crest, and band on the head for articulation with the radial
the m. extensor carpi radialis arises from the re notch of the ulna. The articular circumference
maining part. is longer than the corresponding notch in the
The medial epicondyle (epicondylus mediate) ulna, so that a limited amount of rotation of the
70 Chapter 1. T h e S k e l e t a l S y ste m
\-Olecranon
- Anco n e a i process
C a p i t u l a r depression
A rticular i
circumference'' s -T rochleor notch
Head - -
~Coronoid process
Neck -
" Radial notch
R a dia l ■
tuberosity •Ulnar t u b e r o s i t y
Nutrient foram en
Nu trie n t —
forom en
M e d i a l -----
border
Body- -\|-
L a t e r a l ------
border ■Int erosseous
ci'est
Interosseous -
crest
RA D I U S --U L N A
Ulnar
notchi
A rtic u la r
Carpal -- c ir c u m fe r e n c e
articular
surface- -S tyloid process
Styloid p r o c e s s
Groove for
E x t d i g i t a l i s communis^
Groove for
E x t c a r p i r a d i a l i s •-
G roove f o r
--U lnar notch- _
Abd. po llic ls lo n y u s -
- S t y l o i d process-
S t y lo id process- -
forearm is possible. The bulbous eminence on caudal surface of the radius is the proximally di
the lateral surface of the head does not serve for rected nutrient foramen. Distally, the caudal
muscular attachment; the m. supinator plays surface becomes smoother, wider, and more con
over it in its course to a more distal attachment. vex, as it blends with the caudal surface of the
The neck (collum radii) is the constricted seg distal extremity of the bone.
ment of the radius which joins the head to the The m edial and lateral borders of the radius
body. The constriction is more distinct laterally present no special features. They are smooth
and cranially than it is elsewhere. and acutely rounded as they form the margins of
The body (corpus radii), or shaft, is com the two surfaces of the bone. In large specimens,
pressed so that it presents two surfaces and two the rough area just above the middle of the bone
borders. Its width is two or three times its thick on the caudal surface, for the attachment of the
ness. The radial tuberosity (tuberositas radii) is interosseous ligament, encroaches on the lateral
a small projection which lies distal to the neck border.
on the medial border and adjacent caudal sur The distal extremity of the radius is the most
face of the bone. It is particularly variable in de massive part of the bone. It is irregularly quad
velopment, depending on the breed. This tuber rilateral in shape. Its distal surface (facies articu
osity serves for the lesser insertions of the mm. laris carpea) articulates primarily with the radial
biceps brachii and brachialis. A large eminence carpal bone and to a lesser extent with the ulnar
lies proximal to the radial tuberosity on the lat carpal. This surface is concave, both transversely
eral border of the radius just distal to the neck and longitudinally, except for a caudomedial
and serves for the distal attachment of the cra projection which lies in the groove of the radial
nial crus of the lateral ligament of the elbow carpal bone. The lateral surface of the distal ex
joint (Fig. 2-13). tremity is slightly concave and lipped, forming
The cranial surface (facies cranialis) is convex the ulnar notch (incisura ulnaris), which articu
both transversely and vertically. At the junction lates with a facet near the distal end of the ulna.
of the proximal and middle thirds, on the medial Medially, the styloid process (processus styloi-
border, there frequently is an obliquely placed deus) extends distal to the main carpal articular
rough line or ridge to which the m. pronator surface in the form of a sharp, wedge-shaped
teres attaches. The m. supinator attaches to most projection. The lateral surface of the styloid
of the cranial surface of the radius proximal to process enters into the formation of the carpal
the insertion of the m. pronator teres. In large articular surface; the medial portion is some
specimens, starting at the middle of the lateral what flattened for the proximal attachment of
border, and continuing distally from this border, the medial ligament of the carpal joint. The dor
there are alternating smooth ridges and grooves sal surface of the distal extremity of the radius
which run across the cranial surface of the ra presents three distinct grooves. The most medial
dius; these markings converge toward a short, groove, which is short, distinct, and obliquely
but distinct, oblique groove on the medial part placed, lodges the tendon of the m. abductor
of the distal extremity of the bone. The m. ab pollicis longus. The middle groove, which is the
ductor pollicis longus, which arises on the ulna, largest, contains the tendon of the m. extensor
as it courses distally, crosses the cranial surface carpi radialis. The most lateral groove, which is
of the radius obliquely and accounts for these wider but occasionally less distinct than the
markings. others, contains the tendon of the m. extensor
The caudal surface (facies caudalis) is di digitorum communis. The dorsal carpal ligament
vided into two flat to concave areas by the ver blends with the periosteum on the Up of the car
tical interosseous crest (crista interossea). The pal articular surface. The caudal surface of the
crest, which does not extend to either extrem distal extremity is rough-ended and tuberculate.
ity of the radius, divides the surface into a me It contains many foramina for the passage of
dial two-thirds and a lateral one-third. The in veins from the bone. The palmar carpal ligament
terosseous membrane attaches to it. The larger, blends with the periosteum on this surface.
flat, rough area medial to the crest gives attach
ment to the m. pronator quadratus. A prominent Ulna
rough area extends from the proximal part of the
interosseous crest distally to the lateral border. The ulna (see Figs. 1-68, 1-69), for descrip
The heavy, short, interosseous ligament which tive purposes, is divided into a body, or shaft,
unites the radius and ulna attaches to this raised, and two extremities. Located largely in the post-
roughened area. Slightly above the middle of the axial part of the forearm, it exceeds the radius
72 Chapter 1. T h e S k e l e t a l S y ste m
in length, and is, in fact, the longest bone in the interosseous ligament that attaches to the radius.
body. Proximally it articulates with the humerus The interosseous crest extends proximally from
by the trochlear notch (incisura trochlearis) and the notch which separates the distal extremity
with the articular circumference of the radius by from the body of the ulna. The interosseous
the rad ial n otch (incisura radialis) of the ulna. membrane attaches to the interosseous crest.
Distally it articulates with the ulnar notch of the Medial to the crest a faint vascular groove indi
radius, and with the ulnar carpal and accessory cates the position, in life, of the palmar interos
carpal bones by means of two confluent facets seous artery. The largest nutrient foramen is
on the knoblike distal end. directed proximally and is usually located proxi
The proximal extremity of the ulna is the ole mal to the rough area for the attachment of the
cranon, which serves as a lever arm, or tension interosseous ligament, near the interosseous
process, for the powerful extensor muscles of the crest. Other smaller nutrient foramina are located
elbow joint. It is four-sided, laterally com along the course of the vascular groove in the
pressed, and medially inclined; its proximal end middle third of the body. The m. pronator
is grooved cranially and enlarged and rounded quadratus attaches to the cranial surface of the
caudally. The mm. triceps brachii, anconeus, ulna medial and adjacent to the interosseous
and tensor fasciae antebrachii attach to the cau crest. The mm. abductor pollicis longus and
dal part of the olecranon; the mm. flexor carpi extensor pollicis longus et indicus proprius arise
ulnaris-caput ulnare and flexor digitorum pro- in that order from the cranial surface of the body
fundus-caput ulnare arise from the medial sur of the ulna, progressing from the interosseous
face of the olecranon (Figs. 3-53 and 3-54). crest to the lateral border. The caudal surface
The trochlear notch, is known, in some texts, (facies caudalis) of the ulna, unlike the cranial
as the semilunar notch. It is a smooth, vertical, surface, is smooth and concave throughout. It
half-moon-shaped concavity which faces crani gradually tapers toward the distal end. The m.
ally. A transverse plane through the middle of flexor digitorum profundus-caput ulnare arises
the trochlear notch separates the proximal ex largely from this surface lateral to the radius. The
tremity from the shaft of the ulna. The semilu mm. biceps brachii and brachialis insert mainly
nar outline of this salient notch is formed by a on the roughened ulnar tuberosity (tuberositas
sagittally placed ridge which divides its articular ulnae), which is located near the proximal end of
area into two nearly equal parts. The whole the caudal surface just distal to the trochlear
trochlear notch articulates with the trochlea of notch. The m edial border (margo medialis) is
the humerus so that the sharp-edged, slightiy sharper and straighter than the lateral one. The
hooked anconeal process (processus anconeus), lateral border (margo lateralis) continues the
at its proximal end, fits in the olecranon fossa wide, rounded, caudal border of the olecranon
of the humerus when the elbow is extended. distally and laterally to the distal extremity of
The coronoid process (processus coronoideus), the bone. The foregoing description of the body
distal to the trochlear notch, is divided into a of the ulna does not apply to some specimens, in
prominent, medial projection, and a less prom which the middle third is more prismatic than
inent, lateral one. Both of these eminences are flat. When the middle third is definitely three
articular, facing cranially and proximally, where sided, this feature continues distally, transform
they articulate with the radius and humerus, re ing the usually rodlike distal third to one which
spectively. They increase the surface area of the is three-sided.
elbow joint without contributing materially to The distal extremity of the ulna is separated
its weight-bearing function. from the body of the bone by a notch in its
The body (corpus ulnae), or shaft, in the cranial border. An oval, slightly raised facet
larger working breeds is typically compressed (circumferentia articularis s. facies articularis
laterally in its proximal third, three-sided radialis ulnae distalis) is located in the distal part
throughout its middle third, and cylindrical in of the notch for articulation with the ulnar notch
its distal third. Great variation exists, however, of the radius. The ulna of the dog possesses no
and in long-limbed breeds the body is some head. The pointed, enlarged distal extremity of
what flattened throughout its length. The cranial the ulna is the styloid process (processus styloi-
su rface (facies cranialis) is rough and convex, deus); on its distomedial part there are two con
both longitudinally and transversely. Its most fluent facets. The one which faces cranially is
prominent feature is a slightly raised, oval, rough concave and articulates with the ulnar carpal
area on the middle third of the bone. It serves bone; the smaller, convex, medial facet articu
for the ulnar attachment of the short, but strong, lates with the accessory carpal bone. The styloid
B on es of t h e T h o r a c ic L im b 73
process of the ulna projects slightly farther dis ments. It represents a fusion of the primitive
tally than the styloid process of the radius. radial carpal bone with the central and inter
mediate carpal bones. The proximal surface of
the bone is largely articular for the distal end of
T he F o repa w the radius. The distal surface of the radial carpal
bone articulates with all four distal carpal
The skeleton of the forepaw (manus) in bones. Laterally it articulates extensively with
cludes the carpus, metacarpus, phalanges, and the ulnar carpal. Its transverse dimension is
certain sesamoid bones associated with them. about twice its width.
The carpus, or wrist, is composed of seven bones The ulnar carpal bone (os carpi ulnare s. os
arranged in two transverse rows, plus a small triquetrum) is the lateral bone of the proximal
medial sesamoid bone. Articulating with the row. It is shaped somewhat like the radial carpal,
distal row of carpal bones are the five metacarpal but is smaller. It articulates proximally with the
bones which lie alongside one another and are ulna and radius, distally with the fourth carpal
enclosed in a common integument. Each of the and the fifth metacarpal, medially with the radial
lateral four metacarpal bones bears three phalan carpal, and on the palmar side with the acces
ges which, with their associated sesamoid bones, sory carpal. It possesses a small lateral process
form the skeleton of the four main digits. The and a larger palmar one for articulation with the
small, medially located, first metacarpal bone accessory carpal and metacarpal V. This latter
bears only two, which form the skeleton of the process is separated from the main part of the
rudimentary first digit. The bones of a typical bone on the lateral side by a concave articular
tetrapod manus are serially homologous with area for articulation with the styloid process of
those of the pes. In the lower vertebrate forms the ulna.
three groupings of the carpal and tarsal bones The accessory carpal bone (os carpi acces-
are made. The proximal grouping includes the sorium s. os pisiforme) is a truncated rod of bone
radial, intermediate, and ulnar carpal bones for located on the caudal or palmar side of the ulnar
the manus, and the tibial, intermediate, and carpal. Both ends of this bone are enlarged. The
fibular tarsal bones for the pes. The middle basal enlargement bears a slightly saddle-shaped
grouping includes the central elements, of which articular surface for the ulnar carpal which is
there are three or four in each extremity. The separated by an acute angle from a smaller,
distal grouping comprises a row of five small transversely concave, proximally directed artic
bones which articulate distally with the five ular area for the styloid process of the ulna. The
metacarpal or metatarsal bones. There has been free end is thickened and overhangs slightly.
considerable modification of this primitive ar The accessory carpal bone is not a true carpal
rangement in mammals, with the fusion or loss bone phylogenetically, but is rather a relatively
of various elements. new acquisition found in reptiles and mammals
(Romer 1962). The m. flexor carpi ulnaris inserts
on it.
Carpus The first carpal bone (os carpale primum s. os
trapezium) is the smallest carpal bone. It is
The carpus (Figs. 1-70 to 1-72, 75, and 76), or somewhat flattened as it articulates with the
wrist, includes the carpal bon es (ossa carpi) palmaromedial surfaces of the second carpal and
and the associated sesamoid bones. The term the base of metacarpal II. It articulates proxi
carpus also designates the compound joint mally with the radial carpal and distally with
formed by these bones, as well as the region be metacarpal I.
tween the forearm and metacarpus. The carpal The second carpal bone (os carpale secundum
bones of the dog are arranged in a proximal and s. os trapezoideum) is a small, wedge-shaped,
a distal row so that they form a transversely con proximodistally compressed bone which articu
vex cranial outline and a concave caudal one. lates proximally with the radial carpal, distally
The bones of the proximal row are the radial, with metacarpal II, laterally with the third
ulnar, and accessory carpal bones. Those of the carpal, and medially with the first carpal.
distal row are the first, second, third, and fourth The third carpal bone (os carpale tertium s.
carpal bones. os capitatum) is larger than the second carpal.
The radial carpal bone (os carpi radiale s. os It has a large palmar projection, which articulates
scaphoideum), located on the medial side of the with the three middle metacarpal bones. It ar
proximal row, is the largest of the carpal ele ticulates medially with the second carpal, later
74 Chapter 1. T h e S k e l e t a l S y s te m
ally with the fourth carpal, proximally with the phalanx of the first digit and a single palmar
radial carpal, and distally with metacarpal III. sesamoid bone.
The fourth carpal bone (os carpale quartum M etacarpal bones II to V are the main meta
s. os hamatum) is the largest bone of the distal carpal bones. They are irregular rods with a
row. It presents a caudal enlargement and is uniform diameter. Metacarpals II and V are
wedge-shaped in both cranial and proximal shorter than III and IV and are four-sided, par
views. It articulates distally with metacarpals ticularly at their proximal ends, whereas meta
IV and V, medially with the third carpal, and carpals III and IV are more triangular proxi
proximomedially with the radial carpal. mally. Distally, the bones diverge, forming the
According to Baum and Zietzschmann (1936), intermetacarpal spaces. The heads of the main
each true carpal element arises from a single metacarpal bones possess roller-like cranial parts
center of ossification. The intermediate carpal which are undivided and are separated from the
element fuses with the radial carpal, and then bodies dorsally by sesam oid fo ss a e (fossae sesa-
the two in turn fuse with the separate root of moidales). Between the heads and bodies of the
origin of the central carpal. The accessory carpal metacarpal bones on the palmar side are the
bone has an epiphyseal center of ossification sesamoid impressions (impressiones sesamoi-
which elaborates the cap of the enlarged caudal dales). The caudal parts of the heads possess
end of the bone. prominent, sharp-edged sagittal crests (cristae
The smallest bone of the carpus is a spherical sagittales), which effectively prevent lateral lux
sesamoid bone, about the size of a radish seed, ation of the two crescent-shaped sesamoid bones
which is located in the tendon of insertion of which articulate with these heads. The base of
the m. abductor pollicis longus on the medial metacarpal II extends farther proximally than do
side of the proximal end of the first metacarpal. the other metacarpal bones. It articulates with
According to Baum and Zietzschmann (1936), the first, second, and third carpals, as well as
on the palmar side of the carpus between the with metacarpals I and III. Besides articulating
two rows of bones there are two flat bones with adjacent metacarpals, the base of meta
similar in size to this small sesamoid bone. carpal III articulates with the third and fourth
carpals; the base of metacarpal IV articulates
Metacarpus with the fourth carpal; the base of metacarpal V
articulates with the fourth carpal and the disto-
caudal extension of the ulnar carpal. The interos
The term metacarpus refers to the region of seous muscles arise from the palmar surfaces of
the manus, or forepaw, located between the the bases of all of the main metacarpal bones.
carpus and the digits. The metacarpal bones The proximal palmar surfaces of the bodies of
(ossa metacarpalia I-V) (Figs. 1-73, 1-77) metacarpals II and III provide insertion for the
are typically five in number in primitive mam m. flexor carpi radialis, and the dorsal surfaces of
mals, although supernumerary metacarpal bones the bases provide insertion for the m. extensor
and digits may appear. In many mammals carpi radialis. The small m. adductor digiti
some of the abaxial metacarpal bones and their quinti inserts on the medial surfaces of the distal
accompanying digits have been lost. Like the parts of metacarpals IV and V, and on the lateral
distal row of carpal bones, the metacarpal surface of metacarpal V near the base of the
bones are numbered from the medial to the bone. The accessory carpal bone provides inser
lateral side. The five metacarpal bones are tion for the m. extensor carpi ulnaris, the m.
each cylindrically shaped and enlarged at each abductor pollicis longus inserts on the proximal
end, proximally to form the base, and distally medial part of metacarpal I, and the m. exten
to form the head. The middle portion, or shaft, sor pollicis longus inserts on the distal medial
of each metacarpal bone is known as the part of metacarpal I.
body. Unlike the first metatarsal bone of the The middle parts of the bodies of the meta
hindpaw, the first m etacarpal bone of the fore carpal bones have particularly dense walls.
paw is usually present, although it is by far the These walls become thinner toward the extrem
shortest and most slender of the metacarpal ities, so that the articular cartilages lie on thin
bones. It bears the first digit, which does not cortical bases. During development, the main
quite reach the level of the second metacarpo metacarpal bones have only distal epiphyses.
phalangeal joint. Metacarpal I articulates prox According to Schaeffer (1934), metacarpal I has
imally with the first carpal, and laterally with only a proximal epiphysis. On the proximal third
the second metacarpal. Distally, its laterally en of the caudal surface of each of the four main
larged head articulates with the proximal metacarpal bones there is a nutrient foramen.
B on es o f t h e T h o r a c ic L im b 75
P ro xim a l )
c a rp a l bones
D ista l
c a r p a l b o n e s \^
Meta c a r p a l [
I nt e r me t a c a r p a l
bones '
s p a c e IV
Sesamoid
fossa
_ Do r s a l
se s a mo i d
A rticular face f o r Radiu s -Heod
i A r t i c u i a r f ace f or
Ulnar C a r p a l O Q
F ig . 1-73. Left metacarpal and sesamoid bones,
Radial c a rp a l- disarticulated, dorsal aspect.
S e sa mo i d f a c e -
Base
F ig . 1-71. Left carpus, dorsal aspect. Radial carpal disarticulated.
frochlea
P ro xim a l!
phalanpes
- Base
Sesamoid--m^ 'Extensor
phalanjes i t ub e r c l e
I meiacarpal-
P ha la nx turned Unc j uai Cr e s t
to s h o w
l a t e r a l surface Ungual pro cess
U ncjuis -
F ig . 1-72. Left carpus, articulated, medial aspect. F ig . 1-74. Phalanges, disarticulated, dorsal aspect.
76
B on es of th e T h o r a c ic L im b 77
UI n a - Base
Ra d i u s - _
R a dia l carpal
Accessory canpaU
1 At \ \ carp a! Sagittal
i, i * $ /J | bones crest
Pa! m a r s e s a m o i d bo ne s
1-77. Left metacarpal and sesamoid bones, disarticu
F ig .
lated, palmar aspect.
Sulcus f o r tendon o f
Accessory carp al Flexor c a r p i r a d i a l i s
Radial carpal
m m Sesamoi d
.f-fo c s
Q- Sesamoi d
bone
- Base
Ul nar c a r p a l
P r o x i ma l ’■fll- - B o d u
p h a langes u. i'.AV J
\-H ead
M iddle
pholonpes
Un g u a l crest
F ig .1-76. Left carpal and metacarpal bones, palmar D i s t a l
aspect. Radial and accessory carpals disarticulated. p h a I a npGS ( jj
P ha la nx turned
to s h o w
i| ijjj m e d i a l s u r f a c e
Unguis
U n gual process
parts of the tendons of the four main digits at the The bony pelvis (Fig. 1-79) is formed by the
metacarpophalangeal joints, (Fig. 1-73), whereas ossa coxarum, the sacrum, and the first coccyg
cartilaginous nodules are found at both the dor eal vertebra.
sal and palmar sides of the distal interphalangeal
joints. Os Coxae
BONES OF THE PELVIC LIM B
The os coxae, or hip bone (Figs. 1-80, 1-81),
Each pelvic limb (membrum pelvinum) con is composed of three distinct bones develop-
sists of its half of the pelvic girdle (cingulum mentally. These are the ilium, ischium, and
membri pelvini), composed of the ilium, ischi pubis. They fuse during the twelfth postnatal
um, pubis, and acetabular bone fused as the week, forming the socket which receives the
hip bone (os coxae); the thigh, represented by head of the femur in creation of the hip joint.
the fem ur; the stifle or knee joint, with its me This socket is a deep, cotyloid cavity, called the
nisci, fabellae, and patella; the crus or leg, con acetabulum. The acetabulum in a medium-sized
sisting of the tibia and fibu la; and the hindpaw, dog is 1 cm. deep and 2 cm. in diameter. The
or pes. The pes includes the ankle or tarsus, lunate surface (facies lunata) is the smooth artic
with its digits, consisting of m etatarsals, pha ular circumference which is deficient over the
langes, and the sesam oid bones associated with medial portion of the acetabulum. The cranial
the phalanges. part of the lunate surface is widest as it extends
--Iliac crest
-Ilia c fossa
- -Ilia c fuberosify
A rficu la n surface
N utrient foram en
Body of iliu m
- I I I o p e c t in e a l em i n e n c e
- Lunate s u r f a c e o f ace tabu lu m
-A c e t a b u l a r f o s s a
- Pec t e n o f p u b i c bone
' P ub ic fu b e rcle
S ym ph ysis pelvis
M ed ia l a n y le of is c h ia t ic t u b e r o s i t y
Ihac tuberosity
G re o fe r i s c h i a t i c notch
lliopecfineal line
C a u d a l y l u t e a l l i ne Lunote- s u r f a c e
Body of i l l ur n -
11 i o p e c t i n e a l e m m e n c e -
P e d e n o f pubic bone
Pubic t u b e r c l e - -
O b tu naton f o r a m e n
from the acetabular margin three-fourths of the about as long as its floor, but is offset to the ex
distance to the depth of the acetabulum. The tent that a transverse plane touching the caudal
lunate surface is narrowest mid-laterally, being part of the sacrum also touches the cranial border
about one-half its maximum width. The cranial of the pubis. The lateral osseous wall of the
portion ends medially in a rounded border. pelvic canal is formed largely by the body of
Medially the acetabulum is indented by a notch, the ilium and caudally, to a small extent, by the
the incisura acetabuli. The caudal part of the bodies of the ischium and pubis as these fuse to
acetabular margin or lip which forms the caudal form the acetabulum. The floor of the bony
boundary of the notch is indented by a fissure 2 pelvis is formed by the sacropelvic surfaces of
to 4 mm. deep. The quadrangular, non-articular, the rami of the pubes and ischii. Between these
thin, depressed area which extends laterally rami and the body of the ischium is the large,
from the acetabular notch is the acetabular fossa oval to triangular obturator foram en (foramen
(fossa acetabuli). During the seventh postnatal obturatum). The symphysis pelvis is the median
week, a small osseous element, the acetabular synostosis formed by the right and left pubic and
bone (os acetabuli), located in the floor of the ischial bones. It is, therefore, composed of the
acetabulum between the ilium and ischium, be symphysis pubis cranially and the symphysis
comes incorporated with these larger bones ischii caudally. Occasionally, in young speci
(Fig. 1-82). The pelvic cavity is of considerable mens, there is in the caudal part of the symphy
obstetrical importance since, for survival of the sis a separate triangular bone which is widest
species in nature, it must be large enough to and thickest caudally.
allow for the passage of the young during partu The ilium (os ilium) is the largest and most
rition. The cranial pelvic aperture (apertura cranial of the bones which compose the os
pelvis cranialis), or pelvic inlet, is formed by the coxae. It is basically divided into a cranial, nearly
terminal line (linea terminalis) of the sacrum sagittal, laterally concave part, the wing (ala
dorsally, the cranial border o f the pubis (pecten ossis ilii) and a narrow, more irregular caudal
ossis pubis) ventrally, and the iliopectineal crest part, the body (corpus ossis ilii). The body, at its
bilaterally. The iliopectineal crest (crista iliopec- expanded caudal end, forms the cranial two-
tinea) extends from the iliopectineal eminence fifths of the acetabulum. In this cavity it fuses
(eminentia iliopectinea), which is located on the with the ischium caudally and the pubis medi
terminal line cranial to the acetabulum, to the ally. The ilium is divided into two surfaces, three
promontory of the sacrum. borders, and three angles. The cranial border is
The following conventional measurements of more commonly known as the iliac crest (crista
the pelvis are useful in obstetrics: the transverse iliaca). It forms a cranially protruding arc which
diam eter (diameter transversa) is the greatest is thin in its ventral half; the dorsal half gradu
transverse measurement of the bony pelvic cav ally increases in thickness until it reaches a width
ity. According to Roberts (1956), only in the of nearly 1 cm. dorsally in the large working
achondroplastic types of dogs like the Sealyham breeds. The iliac crest, in heavily muscled
and Pekingese are the transverse diameters breeds, presents a slight lateral eversion. The
greater than the conjugate or sacropubic di dorsal border is thicker in its cranial half than in
ameters. The conjugate (conjugata) measure its caudal half. The caudal half of the dorsal
ment is the distance from the sacrovertebral border is gently concave, forming the greater
angle or the sacral promontory to the cranial ischiatic notch (incisura ischiadica major). The
border of the symphysis pubis. The oblique di dorsal border of the ilium is continuous with
am eter (diameter obliqua) is measured from the the dorsal border of the ischium as a slight con
sacroiliac articulation of one side to the iliopec vexity dorsal to the acetabulum. This is the
tineal eminence of the other. The pelvic axis ischiatic spine (spina ischiadica). The eminence
(axis pelvis) is an imaginary, slightly curved line located dorsal to the iliosacral joint between the
drawn through the middle of the pelvic cavity thin and thick parts of this border is the caudal
from the pelvic inlet to the pelvic outlet. The dorsal iliac spine (spina iliaca dorsalis caudalis).
caudal pelvic aperture (apertura pelvis caudalis), The obtuse angle located between the cranial
or pelvic outlet, is bounded dorsally by the first and dorsal borders is the cranial dorsal iliac spine
coccygeal vertebra, bilaterally by the sacrotu- (spina iliaca dorsalis cranialis). These two spines
berous ligament, and ventrally by the caudo- and the intermediate border constitute what is
lateral border of the tuber ischiadicum on each known as the tuber sacrale in the dog and in
side and the ischiatic arch located between them. the large herbivores, in which it is more sali
The sacral part of the roof of the pelvic canal is ent than it is in the dog. The ventral border
B on es of the P e l v ic L im b 81
\*\- - C a n f i / a j i n o u s area
(margo ventralis) begins at the cranioventral tends from the auricular surface to the iliopectin
angle of the ilium or the cranial ventral iliac eal eminence. It divides the sacropelvic surface
spine (spina iliaca ventralis cranialis). About of the body of the ilium into a medial two-thirds
1 cm. caudal to this spine is a small eminence on and a ventromedial one-third. The caudally di
the thin ventral border which is known in man rected nutrient foramen is located near the mid
as the caudal ventral iliac spine (spina iliaca dle of this surface adjacent to the ventral border.
ventralis caudalis). These two spines and the The mm. sartorius and tensor fasciae latae arise
connecting border constitute what is called the from the tuber coxae. The iliac portion of the m.
tuber coxae. Grooving the ventral border just iliopsoas, the m. sacrospinalis, and portions of
caudal to the tuber coxae and extending on the the mm. coccygeus and levator ani attach to the
lateral surface of the ilium in old specimens is sacropelvic surface (Figs. 3-40 and 3-71). The
the vascular groove for the iliolumbar artery and mm. gluteus medius, gluteus profundus, and
vein. After running caudad parallel to a plane capsularis coxae arise from the gluteal surface of
through the lateral surface of the ilium, the the ilium. The mm. psoas minor, rectus ab
prominent caudal half of the ventral border ends dominis, rectus femoris, and pectineus attach
in the low, but prominent, iliopubic eminence to the iliopectineal eminence and the terminal
(eminentia iliopubica). line adjacent to the eminence.
The gluteal surface (facies glutea) of the ilium The ischium (os ischii) consists of a body,
faces laterally and slightly upward. It embodies ramus, and tuberosity. It forms the caudal third
the whole external surface of the bone. An inter of the os coxae and enters into the formation of
mediate fossa which parallels the axis of the the acetabulum, obturator foramen, and sym
bone divides the surface into a strong ridge dor physis pelvis. The ischiatic tuberosity (tuber
sally and a triangular, moderately rough area ischiadicum) is the caudolateral part of the bone.
ventrally. The medial or sacropelvic surface It gradually thickens, from the medial to the lat
(facies sacropelvina) articulates with the wing of eral side, where it ends in a pronounced rough
the sacrum by a synchondrosis which forms the hemispherical eminence. The caudal end of the
auricular surface (facies auricularis). The iliac sacrotuberous ligament attaches to the dorsal
tuberosity (tuberositas iliaca) is the rough, surface of this eminence.
slightly protruding eminence of the sacropelvic The body o f the ischium (corpus ossis ischii)
surface located dorsal to the auricular surface. is that part of the bone which lies lateral to the
The iliac surface (facies iliaca) is a nearly obturator foramen and at its cranial end forms
square, flat area cranial to the auricular surface. about two-fifths of the acetabulum. Its thick
Between the ventral and dorsal margins of the dorsal border continues with the dorsal border
body of the ilium lies the lateral part of the ter of the ilium in a slight convexity, forming the
minal line, or the iliopectineal line, which ex ischiatic spine (spina ischiadica). Caudal to the
82 Chapter 1. T h e S k e l e t a l S y ste m
spine the dorsal border is flattened and creased of the symphysis pelvis. Caudally it fuses with
by about five shallow grooves, in which lie the the ischium without demarcation at a plane
multiple tendons of the m. obturatorius internus. through the acetabular fossa. The ventral sur
In life the lesser ischiatic notch (incisura ischi face of the pubis gives origin to the mm. gracilis,
adica minor) is converted into a large open adductor, and obturatorius externus. The dorsal
ing, the lesser ischiatic foramen, by the sacrotu- or pelvic surface gives rise to the m. levator ani
berous ligament. The ramus o f the ischium and a part of the m. obturatorius internus. The
(ramus ossis ischii) joins the body at a right an pubic tubercle (tuberculum pubicum) is located
gle. The ramus and the body are twisted at their on the ventral surface of the pubic symphysis.
junction in such a way that the ramus faces dor The cranial border of the pubis, stretching from
sally, forming the ischiatic table (tabula ossis the iliopectineal eminence to the symphysis pu
ischii), and the body faces medially, forming the bis, is also called the pecten ossis pubis, or the
caudal part of the lateral boundary of the pelvic medial part of the terminal line. It serves for
cavity. The m. obturatorius internus arises from the attachment of the prepubic tendon,
the shallow fossa of the ischiatic table which lies whereby all of the abdominal muscles, except
cranial to the ischiatic tuberosity, as well as from for the m. transversus abdominis, attach wholly
the medial and cranial edges of the obturator or in part. The m. pectineus also arises here.
foramen and the adjacent pelvic surface of the
os coxae. The cranial border of the ramus forms Femur
the caudal boundary of the obturator foramen
and the caudal border meets its fellow in form The femur (Figs. 1-83 to 1-85) is the heaviest
ing the deep ischial arch (arcus ischiadicus), bone in the skeleton. In well proportioned
which is formed by the joining of the caudome- breeds it is slightly shorter than the tibia and
dial parts of the adjacent bones. The ventral sur ulna, but is about one-fifth longer than the
face of the ischiatic tuberosity gives rise to the humerus. It articulates with the os coxae prox
most powerful muscles of the thigh, the ham imally, forming a flexor angle of 110 degrees
string muscles: mm. biceps femoris, semitendi- cranially. Distally, it articulates with the tibia,
nosus, and semimembranosus. The adjacent forming a flexor angle of 110 degrees caudally.
ventral Surface of the ramus gives rise to the m. Right and left femurs lie in parallel sagittal
quadratus femoris, and a zone next to the caudal planes when the animal is standing. In fact, all
and medial borders of the obturator foramen the main bones of the pelvic limb are in about
gives rise to the m. obturatorius externus. The the same sagittal plane as those of the ipsilateral
m. adductor arises from the symphysis and the pectoral limb, but the flexor angles of the first
ventral surface of the ischium adjacent to it. two joints of each limb face in opposite direc
The mm. gemelli arise from the lateral surface tions.
of the ischium ventral to the lesser ischiatic The proximal end of the femur presents a
notch. The root of the penis, with its m. ischio- smooth, nearly hemispherical head (caput fem
cavernosus, attaches to the medial angle of the oris), supported by a neck on its proximolateral
ischiatic tuberosity. side and three processes, or trochanters. The
The pubis (os pubis) is a dorsoventrally com head caps the dorsocaudal and medial parts of
pressed, curved bar of bone which extends from the neck. The fo v e a capitis fem oris is a small,
the ilium and ischium laterally to the symphysis rather indistinct, circular pit on the medial part
pubis medially. Its caudal border bounds the of the head. Occasionally a depressed, moder
cranial part of the obturator foramen, which is ately rough, nonarticular strip extends from the
particularly smooth and partly grooved by the fovea to the nearest caudoventral nonarticular
n. obturatorius and the ascending obturator ves margin. The fovea serves for the attachment of
sels. It is divided into a body and a ramus. The the round ligament of the hip joint. The neck
body (corpus ossis pubis) is thick and triangular (collum femoris) unites the head with the rest
in cross section, and enters into the formation of of the proximal extremity. It is about as long as
the acetabulum. The iliopectin eal eminence the diameter of the head, slightly compressed
(eminentia iliopectinea), its largest process, is craniocaudally, and it is reinforced by a ridge
located on the cranial border of the bone as it of bone which extends from the head to the
joins the ilium. At its narrowest part the body large, laterally located greater trochanter.
gives way to the ramus (ramus ossis pubis), the The greater trochanter (trochanter major),
flattened, triangular part of the pubis. Medially the largest tuber of the proximal extremity of
it fuses with its fellow, forming the pubic part the bone, is located directly lateral to the head
Greater G reater
Head
H e a d in - \ V trochanter trochanter
acef abu I u m O b tura tor
N eck - - - Transver se
foramen
Neck- line
f i L i n e of
L e s s e r - -li t
*f j - Vast us l a t
trochanter
- L i n e of
Th i r d V a s t u s med.
trochanter-
Greater trochanter
i
i Trochanteric fo s s a
ir -Fo vea
L a t e r a l lip
Fabellae ♦Jjt £
In te rc o n d y lo id fossa
Lateral condtjle- Medial condole
83
84 Chapter 1. T h e S k e l e t a l S y s te m
and neck. Its free, pyramid-shaped apex usu litea). The relatively flat surface proximally,
ally extends nearly to a frontal plane lying on which is flanked by the diverging femoral lips, is
the head. Between the femoral neck and the the trochanteric surface (facies trochanterica) of
greater trochanter, caudal to the ridge of bone Nickel, Schummer, and Seiferle (1954). The
connecting the two, is the deep trochanteric largest nutrient foramen to enter the femur is
fossa (fossa trochanterica). The mm. gluteus found on the caudal surface at approximately
medius, gluteus profundus, and piriformis insert the junction of the proximal and middle thirds
on the greater trochanter. The mm. gemelli, of the bone. The m. adductor longus inserts on
obturatorius internus, and obturatorius ex the lateral lip distal to the third trochanter,
ternus insert in the trochanteric fossa. The whereas the m. adductor magnus et brevis in
lesser trochanter (trochanter minor) is a dis serts on the whole lateral lip from the third tro
tinct, pyramid-shaped eminence which pro chanter to the popliteal surface. The m. pectin-
jects from the caudomedial surface of the eus inserts on the popliteal surface and the m.
proximal extremity near its junction with the semimembranosus inserts on the adjacent distal
shaft. It is connected with the greater trochanter end of the medial lip.
by a low but wide arciform crest, the intertro The distal end of the femur is quadrangular
chanteric crest (crista intertrochanterica). The and protrudes caudally. It contains three main
m. quadratus femoris inserts distal to the inter articular areas. Two of these are on the medial
trochanteric crest adjacent to the lesser tro and lateral condyles, and the third is an articular
chanter. The most craniolateral eminence of the groove on the cranial surface. The lateral con
greater trochanter is called the cervical tubercle. dyle (condylus lateralis) is convex in both the
On the line which arches distocaudally from this sagittal and the transverse plane. The medial
tubercle is the third trochanter (trochanter ter- con dyle (condylus medialis) is smaller and less
tius), which is about as large as the cervical tu convex in both the transverse and the sagittal
bercle. The m. gluteus superficialis inserts on the plane. Each condyle articulates directly with the
third trochanter. This lateral eminence is about tibia, but most extensively with the menisci of
2 cm. distal to the apex of the greater trochanter. the tibia. They are separated by the intercondy
The transverse line (linea transversa) is dorsally lar fossa (fossa intercondylaris), which is
arched and runs from the femoral head across slightly oblique in direction as the caudal part of
the cranial surface of the intertrochanteric crest the intercondyloid fossa is located farther later
to the greater trochanter. ally than is the cranial part. The articular sur
The shaft, or body (corpus femoris), is nearly faces of the condyles are continuous caudally
cylindrical, and is straight proximally and crani with small facets on the adjacent epicondyles.
ally arched distally. Its cranial, lateral, and The facet on the lateral epicondyle is larger than
medial surfaces are not demarcated from each that on the medial one. These articulate with
other, but the caudal surface is flatter than the sesamoid bones in the tendons of origin of the m.
others. A small proximal nutrient foramen gastrocnemius. The fem oral trochlea or patellar
pierces the cranial surface of the cortex in a surface (trochlea femoris s. facies patellaris) is
distal direction. Covering all but the caudal sur the smooth, wide articular groove on the cranial
face of the femur is the large m. quadriceps surface of the distal extremity which is continu
femoris. All except the rectus femoris division of ous with the articular surfaces of the condyles.
this muscle arise from the proximal part of the Proximally, the limiting ridges diverge slightly.
body of the femur, where occasionally indistinct The medial ridge is somewhat thicker than the
lines indicate the most proximal attachments for lateral one. The patella, or knee cap, articulates
the m. vastus lateralis and the m. vastus medialis. with the patellar surface of the femur.
The caudal surface is marked by a finely rough Proximal and cranial to the medial and lateral
ened surface, the fa cie s aspera, which is narrow condyles are the m edial and lateral epicondyles
in the middle and wider at both ends. This (epicondylus medialis et lateralis). These serve
slightly roughened face is bounded by the for the proximal attachments of the medial and
m edial an d lateral lips (labium mediale et lateral collateral ligaments of the stifle joint. The
laterale), which diverge proximally, running into extensor fossa (fossa extensoria) is a small pit
the lesser and greater trochanters, and, distally, located at the junction of the lateral ridge of the
becoming obscured in the medial and lateral patellar surface and the lateral epicondyle. From
epicondyles, respectively. The sagittally con it arises the m. extensor digitorum longus. The
cave, transversely flat area enclosed distally by m. popliteus arises under the lateral collateral
these lips is the popliteal surface (facies pop- ligament from the lateral condyle of the femur.
B on es of th e P e l v ic L im b 85
On the caudal proximal surfaces of the medial tendon of the m. popliteus (Fig. 1-88), articu
and lateral condyles are facets for the articula lates with the lateral tibial condyle. The fabella
tion of the medial and lateral fabellae, the sesa located in the lateral head of origin of the m.
moid bones located in the tendons of origin of gastrocnemius is the largest fabella and re
the heads of the m. gastrocnemius. Proximal to sembles a small accessory carpal bone. It is
these facets, at the proximal edge of the pop globular in shape, except for a truncated end,
liteal surface, are located tubercles which are which faces distally and has a nearly flat articu
known as the medial and lateral supracondylar lar surface for the facet on the caudal part of the
tuberosities (tuberositas supracondylaris medi lateral femoral condyle. The fabella in the
alis et lateralis). The m. gastrocnemius arises medial head of origin of the m. gastrocnemius is
from both tuberosities. The m. flexor digitorum smaller than the lateral one, and is angular in
superficialis also arises from the lateral supra form. It may not leave a distinct facet on the
condylar tuberosity. medial condyle separate from the articular area
of the condyle. The smallest sesamoid bone of
Sesamoid Bones of the Stifle Joint the stifle region is the fabella located in the
tendon of origin of the m. popliteus, adjacent to
The patella (Fig. 1-84), or knee cap, is the its muscle fibers. It has many sides, one of which
largest sesamoid bone in the body. It is ovate in articulates with the lateral condyle of the femur
shape and curved so as to articulate with the proximal to the lateral margins of the lateral
patellar surface of the femur. The base (basis meniscus.
patellae) is blunt and faces proximally. It may
extend beyond the adjacent articular surface. Tibia
The distally located apex (apex patellae) is
slightly more pointed than the base and does not The tibia (Figs. 1-86 to 1-88) is a long, strong
extend beyond the articular surface. The articu bone which lies in the medial part of the crus, or
lar surface (facies articularis) is smooth, convex true leg. It articulates proximally with the femur,
in all directions, and in some specimens shows distally with the tarsus, and on its lateral side
longitudinal striations. Several nutrient foramina both proximally and distally with the companion
enter the bone from the medial side. The patella bone of the crus, the fibula. Its proximal half is
is an ossification in the tendon of insertion of the triangular in cross section and more massive
great extensor of the stifle, the m. quadriceps than its distal half, which is nearly cylindrical.
femoris. That part of the tendon between its in The proximal end of the tibia is relatively flat
sertion on the tibial tuberosity and the patella is and triangular, with its apex cranial. Extending
also known as the straight patellar ligament. The from the margin of the base on each side of a
patella alters the direction of pull of the tendon central elevation are articular areas which form
of the quadriceps; it protects the tendon and it the proxim al articular surface (facies articularis
provides a greater bearing surface for the tendon proximalis). The divided proximal articular sur
to play on the trochlea of the femur than would face lies on the lateral and m edial condyles
be possible without it. (condylus lateralis et medialis). The articular
The cranial articular area of the stifle is areas of the condyles are separated by a sagittal,
greatly increased by the presence of two or non-articular strip, and two eminences. Although
three parapatellar fibrocartilages (cartilagines the surface area of the two is approximately the
parapatellares). These are grooved cartilages, same, the medial condyle is oval and the lateral
one on each side of the patella, which articulate condyle is nearly circular. Both are convex in the
with the ridges of the patellar surface of the sagittal plane, and concave transversely. In the
femur. Proximally, the two cartilages may ex fresh state they are covered by articular cartilage
tend far enough above the patella to curve and have only a small area of contact with the
toward each other and meet, or a third cartilage articular cartilage of the femoral condyles.
may be located at this site. For a more complete Functionally the medial and lateral tibial con
description of these cartilages refer to Chapter dyles are separated from the medial and lateral
2, on Arthrology. femoral condyles by the m edial and lateral
The fabellae are three sesamoid bones, each m enisci (meniscus medialis et lateralis). These
less than 1 cm. long. Two of these are located in fibrocartilages are biconcave, incomplete discs,
the heads of the m. gastrocnemius caudal to the which are open toward the axis of the bone. The
stifle joint on the medial and lateral condyles central edges of these C-shaped cartilages are
(Fig. 1-85), and the third, intercalated in the thin and concave, and their peripheral margins
C rania l intercondylar / ln f e r c a n d y l a r eminence Cran, i n t e r c o n d y l a r , .Intercondylar
area ' * e mi ne n c e
areai / i i
MediaI condyle, ,M u sc u la r c/roo^e M u s c u l a r Cjroove Lateral condyle
j- L a te r a l condyle T ibi ah
T ib ia l'
tuberosity tuberosity -H e a d
--He ad
T i b i a l cre- st 11 b i o I c r e s t
-FIBULA
■Nutrient f o r a m e n
TIBIA
-TIBIA
Caudal in te rc o n d y la r a r e a
Interosseous
L a t condyle | Med, c o n d y le border
FIBULA
M-p°piiteu^ M M h v 0 W - P°p''!eal
not ch
- Nu t n i e n t -
f o r ame n
Medial
mal leol us ^Lateral malleolus
-TIBIA
_G r o o v e of the
FIB U LA Lat er al m a l l e o l u s
F ig . 1-86. Left tibia and fibula articulated, cranial aspect
F ig . 1-87. Left tibia and fibula articulated, lateral aspec'
Fibular
surface
Distal a rtic u la r
surface
Lateral malleolus-
-Medial m a l l e o l u s
Groove o f Lat m a l l e o l u s - _
F ig . 1-88. Left tibia and fibula disarticulated, caudal aspect
86
B o n es of th e P e l v ic L im b 87
are thick and convex. The intercondylar em i lique line. The mm. flexor hallucis longus, tibi
nence (eminentia intercondylaris) is a low but alis posterior, and flexor digitorum longus arise
stout divided eminence between the medial and from the proximal half of the caudal surface in
lateral tibial condyles. The two spurs which are lateral to medial sequence. Running obliquely
articular on their abaxial sides are known as the distolaterally across the lower part of the caudal
medial and lateral intercondylar tubercles surface may be a vascular groove which extends
(tuberculum intercondylare mediale et laterale). to the distal end of the bone adjacent to the
The oval, depressed area cranial to the inter- lateral malleolus.
condyloid eminence is the area intercondylaris The m edial surface (facies medialis) of the
cranialis; the smaller, depressed area caudal to tibia is wide and nearly flat proximally, as it is
it is the area intercondylaris caudalis. The menis- partly formed by the tibial crest. Near the tibial
cial ligaments attach to these areas. The con crest in large specimens is a low, but wide, mus
dyles are more expansive than the articular areas cular line for the insertions of the mm. semi
located on their proximal surfaces. Between the tendinosus, gracilis, and sartorius. The medial
condyles caudally is the large popliteal notch surface of the tibia is relatively smooth through
(incisura poplitea). The muscular groove (sulcus out, as it is largely subcutaneous in life.
muscularis) is a smaller notch which cuts into The lateral surface (facies lateralis) of the
the lateral condyle as far as the articular area. tibia is smooth, wide, and concave proximally,
On the caudolateral surface of the lateral con flat in the middle, and narrow and convex dis
dyle is the fa cies articularis fibularis proximalis, tally. Part of the m. biceps femoris inserts on the
an obliquely placed facet for articulation with medial surface of the tibial crest, and just caudal
the head of the fibula. The m. semimembranosus to this attachment the m. tibialis cranialis arises.
inserts on the caudal part of the medial condyle, This muscle intimately covers the lateral surface
and the proximal part of the origin of the m. of the tibia. The m. flexor hallucis longus arises
tibialis cranialis arises from the lateral condyle. from the proximal three-fourths of the lateral
The tibial tuberosity (tuberositas tibiae) is the border of the tibia. The m. fibularis brevis arises
large, quadrangular, proximocranial process from the lateral surfaces of the distal two-thirds
which provides insertion for the powerful m. of the fibula and tibia.
quadriceps femoris and parts of the mm. biceps The distal end of the tibia is quadrilateral and
femoris and sartorius. Extending distally from slightly more massive than the adjacent part of
the tibial tuberosity is the tibial crest (crista the body. The distal articular surface is in the
tibiae), which has a slight medial inclination. To form of two nearly sagittal, arciform grooves,
it insert the mm. gracilis, semitendinosus, and the cochlea tibiae, which receive the ridges of
parts of the mm. sartorius and biceps femoris. the proximal trochlea of the tibial tarsal bone.
The body (corpus tibiae) is three-sided The grooves are separated by an intermediate
throughout its proximal half, whereas the distal ridge. A transversely located synovial fossa ex
half is essentially quadrilateral or cylindrical. tends from one groove to the other across the
Three surfaces and three borders are recognized intermediate ridge. The whole medial part of
in the proximal half of the tibia. These are the the distal extremity of the tibia is the m edial
caudal, medial, and lateral surfaces, and the malleolus (malleolus medialis). Its cranial partis
medial, interosseous, and cranial borders. The formed by a stout pyramid-shaped process.
interosseous border (margo interosseus) is re Caudal to this is a semilunar notch. The small,
placed in the distal half of the tibia by a narrow, but distinct, sulcus for the tendon of the m. flexor
flat surface apposed to the adjacent, closely lying digitorum longus grooves the lip of the medial
fibula. This is the fa cie s fibularis of the tibia. malleolus at the center of the semilunar notch.
The caudal surface (facies caudalis) presents On the caudal side of the distal extremity is a
an oblique line which courses from the proximal much wider sulcus for the tendon of the m.
part of the interosseous border to the middle of flexor hallucis longus. The lateral surface of the
the medial border. At the junction of the proxi distal extremity of the tibia is in an oblique plane
mal and middle thirds of the interosseous border as it slopes caudolaterally. It is slightly flattened
is the distally directed nutrient foramen of the by the fibula. At the distal end of the fibular sur
bone. The m. popliteus inserts on the proximal face is a small facet, the fa c ies articularis m alle
medial part of the caudal surface, the proximal oli, for articulation with the distal end of the
part of the medial border, and the adjacent fibula. No muscles attach to the distal half of the
medial surface of the tibia proximal to the ob tibia.
88 Chapter 1. T h e S k e l e t a l S y ste m
a head, neck, and body. The body (corpus tali) the comparable surface of the tibial tarsal. The
forms the proximal half of the bone. The most most distal and the smallest articular surface on
prominent feature of the body is the proximal the dorsal part of the bone is the fa c ie s articu
trochlea (trochlea tali proximalis), the surface laris talaris distalis. This surface is confluent
which articulates with the sagittal grooves and with a small articular facet for the central tarsal
the intermediate ridge of the distal articular on the distal surface. Between the middle and
surface of the tibia. The sides of the trochlea ar distal articular surfaces is the calcanean sulcus
ticulate with the medial and lateral malleoli and (sulcus calcanei). This sulcus concurs with a sim
are known as the fa cies malleolaris medialis and ilar one of the tibial tarsal to form the tarsal
facies malleolaris lateralis, respectively. The tib sinus (sinus tarsi). On the distal end of the fibu
ial tarsal articulates with the fibular tarsal by lar tarsal is a large flat fa c ie s articularis cuboidea,
three distinct and separate facets. The large, for articulation mainly with the central tarsal
concave proximal articular surface of the tibial and by a small facet with the tibial tarsal.
tarsal bone (facies articularis calcanea proxi The central tarsal bone (os tarsi centrale s. os
malis) is plantarolaterally located. The lateral naviculare) lies in the medial part of the tarsus
part of this facet is located on a large right- between the proximal and distal rows. It articu
angled process which is articular on three sides. lates with all of the other tarsal bones. Proxi
It is the lateral process o f the talus (processus mally it articulates with the tibial tarsal by a
lateralis tali). The oval middle articular surface large, concave, roughly oval area. On the proxi
(facies articularis calcanea media) is separated mal surface of the plantar process of the bone,
from the distal part of the dorsal articular surface tuberositas plantaris, is a small facet for articu
by the deep but narrow sulcus tali. The smallest lation with the fibular tarsal. The central tarsal
articular surface for the fibular tarsal bone is articulates distally with the first, second, and
located on the extreme distolateral part of the third tarsals, and laterally with the proximal half
tibial tarsal. It is the distal articular surface of the fourth tarsal.
(facies articularis calcanea distalis). The head The first tarsal bone (os tarsale primum s. os
(caput tali) of the tibial tarsal bone is the trans cuneiforme mediale) varies greatly in develop
versely elongated distal extremity. The distal ment. When it does not exist as a separate bone
surface is rounded, and irregularly oval trans it is fused with the distally lying first metatarsal
versely, and contacts only the central tarsal to bone. It is always compressed transversely.
form the articular surface fo r the central tarsal When it is fused with the first metatarsal it
(facies articularis centralis). The neck (collum forms a rough, bent plate. The first tarsal bone
tali) unites the large, proximally located body normally articulates with the central tarsal, the
with the head. It is smooth and convex medially, second tarsal, and the first metatarsal. Occasion
and lies directly under the skin. ally the first tarsal bone articulates with the sec
The fibular tarsal bone (os tarsi fibulare s. ond metatarsal. Other possible variations are
calcaneus) is the largest and longest bone of the described in the discussion of the first digit of
tarsus. The distal half of the bone is wide trans the hindpaw, under Phalanges.
versely and possesses three facets and two proc The second tarsal bone (os tarsale secundum
esses whereby it is mortised with the tibial tarsal s. os cuneiforme intermedium) is the smallest of
bone, to form a very stable joint. The tuber cal the tarsal bones. It is a wedge of bone which ex
canei, or proximal half of the bone, is a sturdy tends toward the plantar side only a short dis
traction process which serves for the insertion tance. It articulates with the central tarsal prox
of the calcanean tendon. Its slightly bulbous imally, the third tarsal laterally, the first tarsal
free end contains the m edial and lateral proc medially, and the second metatarsal distally.
esses (processus medialis et lateralis), which are The joint with the second metatarsal is at a
separated by a wide groove. A jutting shelf, the higher level than the similar joints lateral to it.
sustentaculum tali, leaves the medial side of the The third tarsal bone (os tarsale tertium s. os
bone. On the plantar side of this process is a cuneiforme laterale) is nearly three times larger
wide shallow groove over which the tendon of and two times longer than the second tarsal
the m. flexor hallucis longus glides. On the dorso- bone. It articulates proximally with the central
medial side is a concave, oval facet, the facies tarsal, laterally with the fourth tarsal, distally
articularis talaris media, for articulation with with the third metatarsal, and medially with the
the middle articular surface of the tibial tarsal. second tarsal and metatarsal. On the plantar side
The dorsal articular surface, fa c ie s articularis it ends in a rounded plantar tuberosity which is
talaris dorsalis, is convex as it articulates with embedded in the joint capsule.
90 Chapter 1. T h e S k e l e t a l S y s te m
Fibula -
L a t e r a l •*- m e d i a l p r o c e s s e s of c a l c a n e a l i u b e r
f .Sustentaculum t a l i
Calcaneal tu b e r-
F ib u ta r tarsal -
bone
tarsal '
bone -T ib ia l tarsal bone
S u l c u s --h
f or f l exors
T i b i a l t ar sa l ^ IV ta rs a l bone - f ^ \
bon e - P l a n t a r process
Pl ant ar process
Sulcus f or tendon of Ilf1'!-Cen t r a l t ar s al bone
Central tarsal bone
M. f i b u l a r i s l ongus
IV ta r s a l bone- | - P l a n t a r process
n-i
P l a n t ar p r o c e s s ' y - m - P i a n t a r p rocess
II m etatarsal bone
III
F ig . 1-89. Left tarsus, articulated, medial aspect.
F ig . 1-90. Tarsal bones disarticulated, plantar aspect.
_^Sulcus f o r p e r f or a t i n g
m e t a t a r s a l a.
M Sustentaculum t a l i
T i b i a l tarsal -Body
, P l a n t ar process
-Central tarsal
" S a g it ta l c rest
F ig . 1-91. Left tarsus, articulated, plantar aspect. F i g . 1-92. Left metatarsal and sesamoid bones, disarticulated,
plantar aspect.
B o n es o f t h e P e l v ic L im b 91
Calcaneol tuber
Fibular ■C a l c a n e a l tuber
tarsal -
b one Trochleoi
Tibial ta rsa l
bone (body) -
■■iii /J'Hlfi
Ti bi al t a r s a l _ F . J™ g !j fi i, ' " A r t i c u l a r f aces f o r
T ib ia l tarsal
j a ] u s bone 1/
N e c k - tK lf - F i b u l a r t a r s a l bone
Central t a r s a l 1 (-
bone Head- - f
- Medial process
C entral ta rsa l--^
£>one
Fig. 1-93. Left tarsus, articulated, lateral aspect. ■S u l c u s f o r t e n d o n o f
M. f i b u l a r i s lo n g u s
Sulcus f o r
P e r f o r a t i n g m e t a t a r s a l a.
Calcaneal t u b e r _
_B a s e
Tibial tarsal
bone
Fibular
tarsal
Trochl ea Body
bone
Head
Central M edial
t a r s a l bone process - S e s a m o i d fossa
.H e a d
u/~
g, ®--Dorsal sesamoid
F ig . 1-95. Left tarsus, articulated, dorsal aspect. F ig . 1-9 Left metatarsal and sesamoid bones, disarticulated, dorsal aspect.
92 Chapter 1. T h e S k e l e t a l S y s te m
The fourth tarsal bone (os tarsale quartum s. of metatarsals II and III form a space through
os cuboideum) is as long as the combined dimen which passes the perforating metatarsal artery
sions of the central and third tarsals, with which from the dorsal to the plantar side of the paw.
it articulates medially. The joint between the The distal end of each main metatarsal bone,
fourth and central tarsals slopes upward and out like each corresponding metacarpal bone, has a
ward, whereas the joint with the third tarsal ball-shaped h ead (caput), which is separated
slopes downward and inward. Proximally, the from the body dorsally by a deep transverse
fourth tarsal articulates mainly with the fibular sesam oid fo ssa (fossa sesamoidalis). On the
tarsal and slightly with the tibial tarsal on its plantar part of the head the articular surface is
dorsomedial edge. Medially, the fourth tarsal divided in such a way that the area nearer the
articulates with the central and third tarsals and axis of the paw is slightly narrower and less ob
distally with metatarsals IV and V. The distal lique transversely than the one on the abaxial
half of the lateral surface is widely grooved for side. The four mm. interossei arise from the
the tendon of the m. fibularis longus, forming plantar side of the bases of the main metatarsal
the sulcus tendinis m. fibularis longi. Proximal bones and intimately cover most of their plantar
to the sulcus is the salient tuberosity o f the surfaces. The m. tibialis cranialis inserts on the
fou rth tarsal b on e (tuberositas ossis tarsalis medial side of the base of metatarsal II and the
quarti). Distally there are two indistinct rectan m. fibularis brevis inserts on the lateral side of
gular areas, sometimes partly separated by a syn the base of metatarsal V.
ovial fossa, for articulation with metatarsals IV
and V. All tarsal bones of the distal row possess Phalanges
prominent plantar processes for the attachment
of the heavy plantar portion of the joint capsule. The phalanges and sesamoid bones of the
hindpaw are so similar to those of the forepaw
Metatarsus that no separate description is necessary, except
for the bones of digit 1.
The term metatarsus refers to the region of The term “dew claw ” is applied to the vari
the pes, or hindpaw, located between the tarsus ably developed first digit of the hindpaw of the
and the phalanges. The metatarsal bones (ossa dog. It should not be applied to the first digit of
metatarsalia I-V) resemble the corresponding the forepaw because that appendage, although
metacarpal bones in general form. They are, rudimentary, is always present. Some breeds are
however, longer. The shortest main metatarsal recognized by the American Kennel Club (1956)
bone, metatarsal II, is about as long as the long as normally possessing fully developed first digits
est metacarpal bone. The metatarsus is com on their hindpaws. Many individuals of the
pressed transversely, so that the dimensions of larger breeds of dogs possess “dewclaws” in
the bases of the individual bones are consider various degrees of development (Kadletz 1932).
ably greater sagittally than they are transversely. In the most rudimentary condition an osseous
Furthermore, as a result of this lateral crowding element bearing a claw is attached only by skin
the areas of contact between adjacent bones is to the medial surface of the tarsus. The proximal
greater and the intermetatarsal spaces are phalanx may be absent, and metatarsal I, much
smaller. The whole skeleton of the hindpaw is reduced in size, may or may not be fused with
longer and narrower than that of the forepaw. the first tarsal. Occasionally two claws of equal
The first metatarsal bone (os metatarsale I ) is size are present on the medial side of the hind
usually atypical and will be described with the paw. These supernumerary digits probably have
phalanges of the first digit. no phylogenetic significance. Complete duplica
Metatarsal bones II, III, IV, and V (ossa met tion of the phalanges and metatarsal I is some
atarsalia II-V) are similar. A typical metatar times encountered. The first metatarsal may also
sal bone consists of a proximal base (basis), which be divided into a proximal and a distal portion.
is transversely compressed and irregular and The distal metatarsal element is never fused to
a shaft or body (corpus), which in general is tri the proximal phalanx. It may be united to its
angular proximally, quadrangular at mid-shaft, proximal part by fibrous tissue, or a true joint
and oval distally. Each body possesses one large may exist. Although the dewclaw may be lack
and several small nutrient foramina which enter ing, a rudiment of the first metatarsal is occa
the proximal halves of the bones from either the sionally seen as a small flattened osseous plate
contact or the plantar surface. Oblique grooves which lies in the fibrous tissue on the medial side
on the opposed surfaces of the proximal fourths of the tarsus.
Os P e n is 93
H ET ER O T O PIC SK ELET O N
Sectiont
94 Chapter 1. T h e S k e l e t a l S y s te m
A RTH RO LO G Y *
95
96 Chapter 2. Arth rolo gy
droses. Unions of this type may be formed by thetized dogs, concluded that “the lymphatic
hyaline cartilage, by fibrocartilage, or by a com system is the essential apparatus for the removal
bination of the two, and they are subject to of protein from joints.”
change with increasing age. The nerve supply of synovial joints is derived
Hyaline cartilage joints, or primary joints, are from peripheral or muscular branches in the
usually temporary and represent persistent parts vicinity of the joint. Included in these articular
of the fetal skeleton or secondary cartilage of nerves are proprioceptive fibers, pain receptor
growing bones. The epiphysis of an immature fibers, and sympathetic fibers related to vaso
long bone is united with the diaphysis by a motor or vasosensory functions. Some areas of
cartilaginous epiphyseal plate. When adult stat the joint capsule are more richly innervated than
ure is reached, osseous fusion occurs and a joint others. Gardner (1950) reviewed the mor
no longer exists, although a slight epiphyseal line phology and physiology of joints, including their
may mark the union. This osseous union, in some innervation, and cites over 500 references. An-
anatomical works, is called a synostosis. Similar sulayotin (1960) studied the nerves which supply
transitory hyaline cartilage joints are typical of the appendicular joints in the dog.
the union of the shafts with the femoral tro
chanters or the humeral tubercles, and of the Structure of Synovial Joints
spheno-occipital synchondrosis. Some hyaline
cartilage joints, such as the costochondral junc The joint capsule is composed of an inner
tions, remain throughout life. synovial membrane and an outer fibrous mem
Fibrocartilaginous joints, or secondary joints, brane. The synovial m embrane (membrana syn-
are sometimes referred to as amphiarthroses. ovialis) is a vascular connective tissue which
The best examples of such joints are those of the lines the inner surface of the capsule and is re
pelvic symphysis, mandibular symphysis, sterne sponsible for the production of synovial fluid.
brae, and vertebral bodies. The fibrocartilage The synovial membrane does not cover the ar
uniting these bones may have an intervening ticular cartilage but blends with the periosteum
plate of hyaline cartilage at each end. Occasion as it reflects onto the bone. Joint capsules may
ally these joints may ossify, as do hyaline carti arise postnatally if the need exists, and thus false
lage joints. joints often form following unreduced fractures.
Synovial membrane covers all structures within
S y n o v ia l J o in t s a synovial joint except the articular cartilage and
the contact surfaces of fibrocartilaginous plates.
The true joints of the extremities permit the Synovial membrane also forms sleeves around
greatest degree of movement and are most com intra-articular ligaments and covers muscles,
monly involved in dislocations. All synovial joints tendons, nerves, and vessels if these cross the
are characterized by a joint cavity (cavum ar- joint closely. Adipose tissue often fills the
ticulare), joint capsule (capsula articularis), irregularities between articulating bones, and
synovial flu id (synovia), and articular cartilage in some instances it is aspirated into or squeezed
(cartilago articularis). A few of the synovial joints out of the joint as the surfaces of the artic
have modifications peculiar to the functions they ulating bones part or come together during
perform and may possess intra-articular liga movement. Fat in such locations is covered by
ments, menisci, fat pads, or synovial projections. synovial membrane. A synovial fo ld (plica syn-
The blood supply of synovial joints is provided ovialis) is an extension of the synovial membrane;
by an arterial and venous network from parent such folds usually contain fat. Around the pe
trunks in the vicinity of the joint. The vessels riphery of some synovial joints the synovial mem
supply the capsule and also the epiphyses brane is in the form of numerous processes, or
bordering the joint. Around the articular mar synovial villi (villi synoviales). These are soft and
gins the blood vessels of the synovial membrane velvety. The synovial membrane may extend be
form anastomosing loops, referred to collectively yond the fibrous layer and act as a bursa under a
as the circuius articuli vasculosus, according to tendon or ligament, or even be in communica
Barnett, Davies, and MacConaill (1961), who tion with a synovial sheath.
quote William Hunter’s description of 1743. The fibrous m em brane (membrana fibrosa) of
Lymphatic vessels are also present in synovial a joint capsule is composed mainly of white fi
membrane and account for the rapid removal of brous tissue containing yellow elastic fibers. It is
some substances from the joint cavity. Bauer, also known as the capsular ligament. In some
Short, and Bennett (1933), working with anes synovial joints the true ligaments are quite sep
G en era l 97
arate from the fibrous capsular ligament such as and Maddock 1932). It receives its nutrition
the patellar ligaments of the stifle joint, but in from the synovia. Because of its mucin content,
most joints the ligaments are thickenings of the the synovia forms a viscous capillary film on the
fibrous portion of the joint capsule. In those articular cartilage. The articular cartilage varies
joints where great movement occurs in a single in thickness in different joints and in different
plane the fibrous membrane is usually thin and parts of the same joint. It is thickest in young
loose on the flexor and extensor surfaces, and healthy joints and in joints which bear consider
thick on the sides of the bone which move the able weight. Its thickness in any particular joint
least. Such thickenings of the fibrous layer are is in direct proportion to the weight borne by
known as collateral ligaments (ligg. collateralia) the joint, and it may atrophy from disuse.
and are present to a greater or lesser degree in Healthy articular cartilage is translucent, with a
all hinge joints. The fibrous membrane attaches bluish sheen. Elasticity and compressibility are
at the margin of the articular cartilage, or at necessary physical properties which it possesses.
most 3 cm. from it, where it blends with the This resiliency guards against fracture of bone
periosteum. by absorbing shock.
The synovial fluid (synovia) serves chiefly to A meniscus (meniscus articularis), or disc (dis
lubricate the contact surfaces of synovial joints. cus articularis), is a complete or partial fibro
In all cases these surfaces are hyaline cartilage or cartilaginous plate which divides a joint cavity
fibrocartilage. Fibrocartilage contains few blood into two parts. The temporomandibular joint
vessels and nerves, and hyaline cartilage has contains a thin, but complete, meniscus, and,
neither. Therefore, the synovial fluid serves the because the capsular ligament attaches to the
additional function of transporting nutrient ma entire periphery of the meniscus, the joint cav
terial to the hyaline cartilage and removing the ity is completely divided into two parts. Two
waste metabolites from it. Synovia also enables menisci are found in the stifle joint, and neither
the wandering leukocytes to circulate in the is complete, thus allowing all parts of the joint
joint cavity and phagocytize the products of the cavity to intercommunicate. Menisci have a
wear and tear of the articular cartilage. In many small blood and nerve supply, and are capable of
joints of man there is little, if any, free synovia. regeneration (Dieterich 1931). Their principal
In 29 normal human knee (stifle) joints, Cogges- function, according to MacConaill (1932), is “. . .
hall et al. (1940) collected on an average only to bring about the formation of wedge-shaped
0.45 ml. of synovia. Davies (1944), on the other films of synovia in relation to the weight-trans
hand, collected 10 ml. of synovia from both the mitting parts of joints in movement.” An obvi
hip and the stifle joints of normal adult cattle. ous function is the prevention of concussion.
The amount and composition vary from joint The stifle and temporomandibular joints are the
to joint. The average volume in the stifle joint of only synovial joints in the dog which possess
adult dogs of various sizes varied from 0.2 ml. to discs, or menisci.
2 ml. The general health and condition of the A ligament (ligamentum) is a band or a cord
dog has a marked influence on the amount of of nearly pure collagenous tissue which unites
synovia present in the joints. No glands are two or more bones. The term also designates
found in the synovial membrane. Synovia is remnants of fetal structures and relatively avas
thought to be a dialysate, although mucin is cular narrow serous membrane connections.
probably produced by the fibroblasts of the syn Ligaments, as the term is used in this Chapter,
ovial membrane (Davies 1944). The chemical unite bone with bone. Tendons unite muscle
composition of synovia closely resembles that of with bone. Ligaments may be intracapsular (stifle
tissue fluid. In addition to mucin, it contains and hip joints) or extracapsular where they are
salts, albumin, fat droplets, and cellular debris. developed within or in relation to the capsular
The quantitative composition of synovia is ligament. They are heaviest on the sides of joints
largely dependent on the type of tissue underly where the margins of the bones do not separate
ing the surface fibroblasts and the degree of but glide on each other. Hinge joints with the
vascularity of this tissue. greatest radii of movement have the longest
The articular cartilage (cartilago articularis) ligaments. The ligaments often widen at their
is usually hyaline cartilage. It covers the artic attached ends, where they blend with the peri
ular surfaces of bones where its deepest part may osteum. Histologically, ligaments are largely
be calcified. It contains no nerves or blood ves composed of long parallel or spiral collagenous
sels, although it is capable of some regeneration fibers, but all possess some yellow elastic fibers
after injury or partial removal (Bennett, Bauer, also. The integrity of most joints is ensured by
98 Chapter 2. A rth rolo gy
the ligaments, but in some (shoulder and hip) the surfaces of the radiocarpal articulation form an
heavy muscles which traverse the joints play a ellipsoidal joint.
more important part than do the ligaments. Such A hinge joint (articulatio angularis s. gingly-
muscles and their tendons are sometimes spoken mus) permits flexion and extension with a limited
of as active ligaments. In hinge joints ligaments degree of rotation. The most movable surface of
limit lateral mobility, and some (cruciate liga a hinge joint is usually concave. An example
ments of the stifle joint) limit folding and open would be the elbow joint.
ing of the joint as well. In certain ball-and- A condylar joint (articulatio condylaris) re
socket synovial joints the sockets are deepened sembles a hinge joint in its movement but differs
by ridges of dense fibrocartilage, known as gle in structure. The surfaces of such a joint include
noid lips (labia glenoidalia). rounded prominences or condyles which fit into
reciprocal depressions or condyles on the adja
Pathology cent bone, resulting in two articular surfaces
usually included in one articular capsule. Ex
Articular separations are spoken of as luxa amples of condylar joints include the temporo
tions or dislocations. Although most luxations mandibular joint and the knee joint. The knee,
are due to injury or degenerative changes, there or stifle, joint is best classified as a complex-
are also predisposing genetic factors (often condylar joint since it possesses an intra-articu
breed-specific) which play an important role. lar fibrocartilage which partially subdivides the
Leonard (1960) has described the etiology, diag intra-articular cavity.
nosis, and treatment of luxations in the dog. A trochoid (articulatio trochoidea), or pivot
joint, is one in which the chief movement is
around a longitudinal axis through the bones
Classification of Synovial Joints
forming the joint. The median atlanto-axial joint
and the proximal radio-ulnar joint are examples
Synovial joints may be classified according to
of trochoid joints.
(1) the number of articulating surfaces involved,
(2) the shape or form of the articular surfaces, or A saddle joint (articulatio sellaris) is character
(3) the function of the joint (Barnett, Davies, ized by opposed surfaces each of which is con
vex in one direction and concave in the other,
and MacConaill 1961).
According to the number of articulating sur usually at right angles. When opposing joint sur
faces a joint is either simple (articulatio simplex) faces are concavoconvex, the main movements
or com pound (articulatio composita). A simple are also in planes which meet at right angles.
joint is formed by two articular surfaces within The interphalangeal joints are examples of this
an articular capsule. When more than two artic type of articulation.
ular surfaces are enclosed within the same cap
sule, the joint is compound.
The classification of synovial joints, approved Movements of Synovial Joints
by the VI International Congress of Anatomists
in 1955 and revised in 1961 (Nomina Anatomica, Joint movements that are brought about by
2nd Edition. Excerpta Medica Foundation), is the contraction of muscles which cross the joints
based on the shape or form of the articular sur are known as active movements; those joint
faces. There are seven basic types: movements caused by gravity or secondarily by
A plane joint (articulatio plana) is one in the movement of some other joint or by an ex
which the articular surfaces are essentially flat. ternal force are known as passive movements.
It permits a slight gliding movement. An ex Synovial joints are capable of diverse move
ample would be the costotransverse joint. ments. Flexion, or folding, denotes moving two
A ball-and-socket, or spheroidal joint (artic or more bones so that the angle between them
ulatio spheroidea), is formed by a convex hemi becomes less than 180 degrees. Extension, or
spherical head which fits into a shallow glenoid straightening, denotes movement by which the
cavity (shoulder joint) or into a deep cotyloid angle is increased to 180 degrees. It is readily
cavity (hip joint). seen that some joints such as the metacarpopha
An ellipsoidal joint (articulatio ellipsoidea) is langeal and metatarsophalangeal joints are in
similar to a spheroidal joint. It is characterized a state of overextension. This is called dorsal
by an elongation of one surface at a right angle flexion, since the angle of a ginglymus which can
to the other, forming an ellipse. The reciprocal be reduced the most from 180 degrees is al
convex (male) and concave (female) elongated ways regarded as the flexor side. When an animal
L ig a m e n t s and J o in t s of the Skull 99
“humps up” it flexes its vertebral column. Some the meniscus as it passes between the two bones.
parts of the vertebral column are normally in a The joint cavity is thus completely divided into
state of flexion (the joints between the first few a dorsal meniscotemporal compartment, between
coccygeal vertebrae), whereas others are in a the meniscus and temporal bone, and a ventral
state of overextension, or dorsal flexion (the m eniscom andibular compartment, between the
joints between the last few cervical vertebrae). meniscus and mandible. Laterally the fibrous
Flexion and extension occur in the sagittal plane part of the joint capsule is strengthened by fi
unless the movement is specifically stated to be brous strands to form the mandibular ligament
otherwise (right or left lateral flexion of the (lig. mandibulae).
vertebral column). Adduction is the term applied The symphysis o f the m andible (symphysis
to moving an extremity toward the median plane mandibulae) is the median synchondrosis unit
or a digit toward the axis of the limb. Abduction, ing right and left mandibular bodies. The op
or taking away, is the opposite movement. Cir posed articular surfaces are interdigitated and
cumduction occurs when an extremity follows in the fibrocartilage of the symphysis persists
the curved plane of the surface of a cone. Rota throughout life.
tion is the movement of a part around its long
axis. Joints of Auditory Ossicles
the head of the malleus to the roof of the epi- Sutura frontoparietalis
tympanic recess, and a short rostral ligament Sutura squamosa
connecting the rostral process of the malleus to Sutura sagittalis
the osseous tympanic ring. The body of the incus Sutura sphenoparietalis
is attached to the roof of the epitympanic recess
by a dorsal ligam ent, and the short crus of the Frontal Bone
incus is attached to the fossa incudis by a caudal Sutura interfrontalis
ligam ent. The base of the stapes is attached to Sutura frontoparietalis
the margin of the oval window by an annular Sutura sphenofrontalis
ligament. Sutura frontonasalis
Sutura frontomaxillaris
Hyoid Apparatus Sutura frontolacrimalis
Sutura frontopalatina
The tympanohyoid cartilage articulates with Sutura frontoethmoidalis
the mastoid process of the petrous temporal
bone forming the synchondrosis temporohy- Sphenoid Bone
oidea. This articulation is adjacent to the stylo Sutura vomerosphenoidalis
mastoid foramen. Except for the temporohyoid Sutura sphenoethmoidalis
joint there are tightly fitting synovial cavities be Sutura sphenopalatina
tween all parts of the hyoid complex, as well as a Sutura sphenofrontalis
small synovial cavity between the thyrohyoid Sutura sphenosquamosa
bone and the cranial cornu of the thyroid carti Sutura sphenoparietalis
lage. Sutura pterygosphenoidalis
A p i c a l l i g. o f d e n s -
A tl a nt o - o c c i p i t a l J o i n t c a p s u l e ------ ir r ------------------
L a t e r a l a t l a n t o - o c c i p i t a l licj. — , ^
Alar I i c j a m e n t s -------------
T r a n s v e r s e Hcj. o f a t l a s ---------------------
Atlanto-axial J o i n t c a p s u l e — ^
diverge from each other and attach to the occip terspinous ligaments send some strands to its
ital bone medial to the caudal parts of the occipi ventral surface, but the supraspinous ligament
tal condyles. The transverse atlantal ligament more than the interspinous ligaments prevents
(lig. transversum atlantis) is a strong ligament abnormal separation of the spines during flexion
which connects one side of the ventral arch of of the vertebral column.
the atlas to the other. It crosses dorsal to the The ventral longitudinal ligament (lig. longi-
dens and functions to hold this process against tudinale ventrale) (Fig. 2-5) lies on the ventral
the ventral arch of the atlas. A spacious bursa surfaces of the bodies of the vertebrae. It can be
exists between the ventral surface of the liga traced from the axis to the sacrum, but it is best
ment and the dens. developed caudal to the middle of the thorax.
The dorsal longitudinal ligament (lig. longitudi-
O t h e r S y n o v ia l J o i n t s o f th e nale dorsale) (Fig. 2-6) lies on the dorsal surfaces
V er t eb r a l C olum n of the bodies of the vertebrae. It therefore forms
a part of the floor of the vertebral canal. It is nar
The synovial joints of the vertebral column rowest at the middle of the vertebral bodies and
caudal to the axis are those which appear in pairs widest over the intervertebral fibrocartilages.
between the articular processes of contiguous The dorsal longitudinal ligament attaches to the
vertebrae and the joints between the ribs and rough ridges on the dorsum of the vertebral bod
the vertebrae. The articular capsules are most ies and to the intervertebral fibrocartilages. It
voluminous in the cervical region and at the base extends from the dens of the axis to the end of
of the tail, where the greatest degrees of move the vertebral canal in the coccygeal region. The
ment occur. In the lumbar region there is essen dorsal longitudinal ligament is heavier than the
tially a sagittal interlocking of the cranial and ventral longitudinal ligament.
caudal articular processes. At the tenth thoracic
vertebra the direction of the articular processes Sh ort L ig a m e n t s o f t h e Vertebral
changes. The caudal articular processes of this C olum n
segment face laterally, and the cranial articular
processes face dorsally. The articular processes The intervertebral discs (disci interverte-
of all vertebrae cranial to the tenth thoracic are brales) are interposed in every intervertebral
in nearly a frontal plane so that the cranial artic space, uniting the bodies of the adjacent verte
ular processes face dorsally and the caudal ar brae, except the first two. In the sacrum of
ticular processes face ventrally. young specimens, transverse lines indicate the
planes of fusion of the discs with the adjoining
L ong L ig a m e n t s o f t h e Vertebral vertebral bodies. The thickness of the discs is
C olum n greatest in the cervical and lumbar regions, the
thickest ones being between the last few cervical
The nuchal ligament (lig. nuchae) (Fig. 2-4) vertebrae. The thinnest discs are in the coccyg
is composed of longitudinal yellow elastic fibers eal region, those between the last few segments
which attach cranially to the caudal part of the being smaller in every way than any of the
heavy spinous process of the axis. It extends cau others. Each intervertebral disc consists of an
dally to the tip of the spinous process of the first outer laminated fibrous ring, and a central,
thoracic vertebra. It is a laterally compressed, amorphous, gelatinous center, the pulpy nucleus.
paired band which lies between the medial sur The fibrous ring (annulus fibrosus) consists of
faces of the mm. semispinales capiti. The yellow bands of parallel fibers which run obliquely from
nature of the nuchal ligament can be traced in one vertebra] body to the next. They provide a
the supraspinous ligament to the tenth thoracic means for the transmission of stresses and strains
spinous process (Baum and Zietzschmann 1936). which are required by all lateral and upward
The supraspinous ligament (lig. supraspinale) movements. These bands of fibers cross each
(see Fig. 2-7) extends from the spinous process other in a lattice-like pattern and are over eight
of the first thoracic vertebra to the third coccyg layers thick ventrally. Near the nucleus pulpo-
eal vertebra. It is a heavy band especially in the sus the annulus fibrosus loses its distinctive struc
thoracic region, where it attaches to the apices ture and form and becomes more cartilaginous
of the spines as it passes from one to another. and less fibrous.
Bilaterally the dense collagenous lumbodorsal The pulpy nucleus (nucleus pulposus) is a ge
fascia imperceptibly blends with it throughout latinous remnant of the notochord. Its position
the thoracic and lumbar regions. The feeble in- and shape are indicated on each end of the verte-
104 Chapter 2. Arthrology
L ig a m e n t s and J o in t s of the Vertebra l C olum n
V e n t r a l l o n g i t u d i n a l lig.
_ _ _ Li g. o f n e c k
Lig. o f h e a d
Infervertebral di sc
Conjugal l i g a m e n t
Yellow l i g a m e n t
■Dorsal c o s t o t r a n s v e r s e I ig.
I
F ig . 2-6. Ligaments of vertebral column and ribs, dorsal aspect.
106 Chapter 2. Arth rolo gy
bral body as a depressed area surrounded by a passes from the head of the rib to the lateral part
line. Since its consistency is semifluid, it bulges of the disc. The last three or four ribs are dis
when the retaining fibrous ring ruptures or de placed caudally at their vertebral articulations;
generates. Resultant pressure upon the spinal the ligament also shifts caudally and attaches to
cord may cause posterior paralysis. the body of the vertebra adjacent to the disc.
The interspinous ligaments (ligg. interspi- The conjugal ligament (lig. conjugate cos-
nalia) (Fig. 2-7) connect adjacent vertebral tarum) (see Figs. 2-5, 2-6) might well be classi
spines. They consist of laterally compressed fied as a ligament of the head, probably being
bands of tissue interspersed with muscle bundles homologous with the intra-articular ligament of
of the mm. interspinalis. The bands run from the man. Because of its unique position and func
bases and borders of adjacent spines and decus tion it merits a separate description. It is a col
sate as they insert on the opposed caudal and lagenous cord which runs from the head of one
cranial borders of adjacent spines near their dor rib over the dorsal part of the disc, but under
sal ends. The stronger fibers of the interspinous the dorsal longitudinal ligament, to the head of
ligaments lie almost vertically. Some of their the opposite rib. It grooves the dorsal part of the
fibers blend dorsally with the supraspinous liga disc. A synovial membrane between the lig
ment. Great variation exists, and there seems to ament and the disc joins the joint capsules of the
be no correlation with body type. opposite rib heads. The conjugal ligament is at
The intertransverse ligaments (ligg. inter- tached both cranially and caudally to the disc by
transversaria) consist of bundles of fibers which a delicate membrane, and is attached dorsally to
unite the craniolaterally directed transverse the dorsal longitudinal ligament and dura by
processes of the lumbar vertebrae. They are not areolar tissue. The ligament functions to hold
distinct in any of the other regions of the spine. the heads of opposite ribs tightly against their
The yellow ligaments (ligg. flava) (see Fig. 2 - articular sockets and to prevent excessive cranial
4), or interarcuate ligaments, are loose, thin elas and caudal movements of the ribs. There is no
tic sheets between the arches of adjacent verte conjugal ligament uniting the first pair or the last
brae. Laterally they blend with the articular two pairs of ribs, and that connecting the heads
capsules surrounding the articular processes. of the eleventh pair of ribs is smaller than the
Ventral to this ligament is the epidural space others.
which separates the ligaments and the arches of The dorsal costotransverse ligament (see Fig.
the vertebrae from the dura covering the spinal 2-6), or the ligament o f the tubercle (lig. tuber-
cord. culi costae), is the strongest single ligament unit
ing the rib to the vertebra. It attaches just distal
LIGAM ENTS AND JO IN T S OF THE R IBS to the articular capsule of the tubercle, crosses
the capsule, and blends with the periosteum of
Each typical rib articulates with the vertebral the transverse process of the vertebra corre
column by two synovial joints and with the ster sponding to the rib. The ligaments of the tuber
num by one. There is usually a slightly enlarged cles of the first five ribs lie cranial to the joints
synchondrosis between the rib and its costal car and run obliquely craniomedially from the tu
tilage. bercles to the transverse processes, the ligaments
The costovertebral joints (articulationes cos- of the next three run almost directly medially to
tovertebrales) are formed by the articulation of the transverse processes from the dorsal surfaces
the capitulum of each rib (articulatio capitis cos of the tubercles, and those of the last four incline
tae) with the costal facets of the appropriate ver increasingly caudally as they run from the rib
tebrae, and the articulation of each tuberculum tubercles to the transverse processes of the ver
(articulatio costotransversaria) with the trans tebrae. Great variation in size and position of
verse process of the corresponding vertebra. The these ligaments exists in different dogs. The lig
articular capsules of these joints are thin-walled aments of the tubercles are usually strongest on
synovial sacs which completely surround each the last four ribs.
joint and are associated with the four ligaments The ligament o f the neck (lig. colli costae) (see
of the costovertebral articulation. These are the Figs. 2-5, 2-6) consists of collagenous bundles
ligament of the head and the conjugal ligament which extend between the neck of the rib to the
both of the capitular joint, and the ligaments of ventral surface of the transverse process and the
the tubercle and neck of the tubercular joint. adjacent lateral surface of the body of the verte
The ligament o f the head (lig. capituli costae) bra. Running from the caudal surface of the
(see Fig. 2-5) is a small ligamentous band which neck of the rib to the body of the vertebra of the
L ig a m e n t s ANd j
JOINTS OF TH E
F ig . 2 - 7 t ■
• ^'gaments of
108 Chapter 2. A rth ro lo g y
same number is a ligament; passing caudally, white membranous sheets and bands of thick
these ligaments progressively decrease in size. A ened periosteum, the sternal membrane (mem
ligament also runs from the cranial surface of the brana sterni). The dorsal part is divided into two
neck of each rib to the transverse process of the or more strands, whereas the ventral part con
vertebra of the same number. sists of a single median band. The costoxiphoid
The sternocostal joints (articulationes sterno- ligaments (ligg. costoxiphoidea) are two flat
costales) (Figs. 2-7, 2-8) are synovial joints cords which originate on the eighth costal carti
formed by the first eight costal cartilages articu lages. They cross ventral to the ninth costal car
lating with the sternum. The second to seventh tilages, and converge and blend as they join the
pairs of joints are typical, but the first and last periosteum on the ventral surface of the caudal
pairs present special features. The first sternebra half of the xiphoid process.
is widened cranially by the formation of lateral The costochondral joints (articulationes costo-
shelves of bone which articulate with the trans chondrales) are the joints between the ribs and
versely compressed costal cartilages of the first the costal cartilages. Apparently, no synovial
ribs. These costal cartilages approach their ster cavities ever develop here. In puppies these
nal articulations at a more acute angle than do joints are slightly enlarged and appear as a lon
any of the other costal cartilages. The last sterno gitudinal line of beads on the ventrolateral sur
costal joints, typically, are formed by the ninth face of the thorax.
pair of cartilages joining each other and together
articulating with the ventral surface of the fibro LIGAMENTS AND JO IN T S OF THE
cartilage between the last two sternebrae, or THORACIC LIM B
with the sternebra cranial to the xiphoid process.
The ends of the right and left ninth costal carti S h o u l d e r J o in t
lages are united by an indistinct collagenous lig
ament. No synovial joint is found here, as the The shoulder joint (articulatio humeri) (Figs.
ninth costal cartilages lie closely applied to the 2-9, 2-10) is the ball-and-socket joint between
eighth costal cartilages. A typical sternocostal the glenoid cavity of the scapula and the head
joint is vertically elongated so that its length is of the humerus. It is capable of movement in
double its width. Being in a vertical plane, it al any direction, but its chief movements are flex
lows only forward and backward movements. ion and extension. The shallow, small glenoid
The joint capsule is usually thin, except dorsally cavity of the scapula is increased in size and
and ventrally, where the heavy perichondrium deepened by the glenoid lip (labrum glenoidale),
leaving the costal cartilages thickens and spreads which extends 1 or 2 mm. beyond the edge of
out as it goes to the intersternebral fibrocarti the cavity caudolaterally. The articular capsule
lages. These are the dorsal an d ventral sterno (capsula articularis) forms a loose sleeve which
costal radiate ligaments (ligg. sternocostalia attaches just peripheral to the glenoid lip prox
radiata dorsalia et ventralia). The dorsal and imally. In places the capsule attaches several
ventral surfaces of the sternum are covered by millimeters distal to the articular part of the hu-
D o r s a l s t e r n o c o s t a l r a d i a t e Ha.
-Biceps tendon
------------L a t e r a l
, M edial
g l e n o h u m e r a l / i c j a me n t
T r a n s v e r s e Humeral ligament
LA TERA L MEDIAL
F ig. 2-9. Left shoulder joint.
meral head, where it blends with the periosteum pouch which attaches distal to the supratroch
on the neck of the humerus. A part of the joint lear foramen, so that there is no intercommuni
capsule surrounds the tendon of origin of the m. cation between the extensor and flexor pouches
biceps brachii and extends distally about 2 cm. through the supratrochlear foramen. The joint
in the intertubercular groove. The tendon with capsule dips down between the radial notch of
its synovial sheath is held in the groove by the the ulna and the articular circumference of the
transverse humeral ligam ent (lig. transversum radius. Everywhere but cranially the synovial
humerale). The capsule blends with this liga membrane attaches closely to the articular carti
ment craniomedially and with the tendon of the lage. Medially it sends a distal pouch under the
m. subscapularis medially. Laterally the joint m. biceps brachii, and similar extensions occur
capsule blends with the tendons of the mm. su- laterally under the mm. extensor carpi radialis
praspinatus and infraspinatus. Elsewhere, espe and extensor digitorum communis. On the cau-
cially caudally, the articular capsule is thin and domedial side, extensions of the capsule occur
possesses a number of irregular pouches when it under the mm. flexor carpi radialis and flexor
is distended. Medially and laterally the fibrosa digitorum profundus, caput humerale.
of the capsule is irregularly thickened to form The ulnar collateral ligam ent (lig. collaterale
the m edial and lateral glenohum eral ligaments ulnare) attaches proximally to the lateral epicon
(ligg. glenohumeralia medialis et lateralis). These dyle of the humerus. Distally it divides into two
reinforcing bands protrude appreciably into the crura. The slightly larger cranial crus attaches to
joint cavity.The heavy tendons which cross the a small lateral eminence distal to the neck of the
joint may be called active ligaments. They en radius. The flatter caudal crus passes to the ulna.
sure its integrity, and do this so well that shoul At the level of the articular circumference the
der dislocation is practically unknown in domes ligament blends with the annular ligament and,
tic animals. according to Baum and Zietzschmann (1936),
often contains a sesamoid bone.
The radial collateral ligament (lig. collaterale
E lbow J o in t radiale) is weaker than the ulnar collateral liga
ment, which it resembles. It attaches proximally
The elbow joint (articulatio cubiti) (Figs. 2 - to the medial epicondyle of the humerus, crosses
11 to 2-14) is a composite joint formed by the the annular ligament distally, and divides into
humeral condyle with the head of the radius, two crura. The weaker cranial crus attaches
the humeroradial joint (articulatio humeroradi- proximal to the radial tuberosity. The stronger
alis), and with the semilunar notch of the ulna, caudal crus passes deeply into the interosseous
the humeroulnar joint (articulatio humeroul- space, where it attaches mainly on the ulna but
naris). The proximal radioulnar joint (articula also partly on the radius.
tio radioulnaris proximalis) freely communicates The annular ligament o f the radius (lig. annu
with the main part of the elbow joint, and is re lare radii) is a thin band which runs transversely
garded as a part of it. The humeroradial part of around the radius. It attaches to the lateral and
the elbow joint transmits most of the weight sup medial extremities of the radial incisure of the
ported by the limb. The humeroulnar part stabi ulna. It lies under the collateral ligaments and is
lizes and restricts the movement of the joint to slightly blended with the ulnar collateral liga
a sagittal plane, and the proximal radioulnar ment. In conjunction with the ulna, it forms a
joint allows rotation of the antebrachium. Lat ring in which the articular circumference of the
eral movements of the elbow joint are minimal radius turns when the forearm is rotated.
because of the strong collateral ligaments and The oblique ligament (lig. obliquum s. chorda
the forward protrusion of the anconeal process obliqua) attaches to the dorsal edge of the supra
of the ulna into the deep olecranon fossa of the trochlear foramen. As a small but distinct band,
humerus. Enough rotational movement occurs it crosses the flexor surface of the elbow joint
at the radioulnar and carpal joints so that the distomedially to the tendons of the mm. biceps
forepaws can be supinated about 90 degrees. brachii and brachialis. At the level of these ten
The joint capsule is common to all three artic dons, directly peripheral to the annular liga
ular parts. It is taut on the sides but expansive ment, it divides into two parts. The shorter part
in front and behind. On the cranial or flexor sur blends with the cranial crus of the radial collat
face, it attaches proximal to the supratrochlear eral ligament. The longer branch ends on the
foramen and encompasses most of the radial medial border of the radius after looping around
fossa. Caudally, or on the extensor surface, the the tendons of the mm. biceps brachii and
joint capsule forms a loose fat-covered synovial brachialis.
L ig a m e n t s and J o in t s of the T h o r a c ic L im b 111
Hum erus
Radi al c ol l a t e r a l — Brachialis
Iicjament -B iceps
Caudal v c r a n ia l Obligue li cj.
crura-
B ic e p s t
brachialis tendon
Ulna
R adius
O b l i q u e licj.
Ul n a n
Biceps-- - c o l l a t e r a l l i e]■
— A n n u l a r licj■
Brachialis-
A nnular hcj.-
U l n a r c o l l a t e r a l l i g.
C audal v cranial crura
Interosaeous membrane
In te ro s se o u s lig
Olecranon
I icjament
J O IN T CAPSULE
F ig . 2-14. Left elbow joint, caudal aspect.
L ig a m e n t s and J o in t s of th e T h o r a c ic L im b 113
of the radius and on the palmar projections of These ligaments diverge as they run distally,
the radial and first carpals. The transverse pal thus allowing a free opening on the cranial sur
mar carpal ligament is divided into two parts. face of the antebrachiocarpal joint during flex
One lies superficial and the other lies between ion. The ulna is securely anchored to the palmar
the tendons of the superficial and deep digital side of the radial carpal by an obliquely running
flexors. The carpal canal (canalis carpi) on the ligament located just proximal to the accessory
palmar side of the carpus is formed superficially carpal bone. From the palmar surface of the
by the superficial part of the transverse palmar radius, near its distal articular cartilage, a liga
carpal ligament and deeply by the palmar part ment runs to the palmar surface of the radial
of the joint capsule. It is bounded laterally by carpal. A short leaf of this ligament runs from
the accessory carpal bone. It contains the ten the midpalmar surface of the radius to the radial
dons and synovial sheaths of the mm. flexor digi carpal. A flat band nearly 1 cm. wide runs from
torum superficialis and flexor digitorum profun the palmarolateral surface of the radius from
dus, as well as the radial, ulnar, and palmar within the distal part of the interosseous space to
interosseous arteries and veins and the ulnar and the lateral surface of the hidial carpal adjacent
median nerves. to the ulnar carpal. The accessory carpal bone is
The palmar carpal fibrocartilage (fibrocarti- secured distally by two ligaments which origi
lago carpometacarpeum palmare) (Fig. 2-15) is nate near its enlarged, rounded, free end. Dis
quite thick and sharply defined proximally. As it tally one attaches to metacarpal V and the other
crosses the palmar surfaces of the carpal bones, to metacarpal IV. Many short ligaments unite
it attaches to all except the accessory carpal bone. the carpal bones transversely, holding them as
The thickest attachment on the accessory carpal units in the two rows.
bone is to its dorsomedial border, just caudal to
the articulation of the radial carpal with the
ulnar carpal. The palmar carpal fibrocartilage is Metacarpal Joints
very heavy distally. This attaches to the palmar
surfaces of the distal row of carpal bones and the The intermetacarpal joints (articulationes
adjacent surfaces of the proximal parts of met intermetacarpeae) are close fitting joints be
acarpals III, IV, and V. The palmar carpal tween the proximal ends of adjacent metacarpal
fibrocartilage serves as the origin for most of the bones. The synovial membrane from the adja
special muscles of digits 2 and 5, as well as cent carpometacarpal joint extends a few milli
furnishing part of the origin for the interosseous meters between the metacarpal bones. Distal to
muscles. It flattens the palmar irregularities at the synovial part, the bones are united for vari
the carpometacarpal joints and furnishes a able distances by fibrous tissue, the interosseous
smooth deep surface for the carpal canal. m etacarpal ligaments (ligg. metacaipea interos-
The special ligaments o f the carpus will be sea). Distal to these ligaments are the interosse
treated briefly. Some of the smaller ones will not ous spaces of the metacarpus (spatia interossea
be described, but all are illustrated in Figures 2 - metacarpi).
17 to 2-21. The metacarpophalangeal joints (articula
The short radial collateral ligam ent (lig. col tiones metacarpophalangeae) are the five joints
laterale radiale breve) consists of a straight and formed by the distal ends of the metacarpal
an oblique part. The straight part runs from a bones and the proximal ends of the proximal
tubercle above the styloid process to the most phalanges. To these are added in each of the
medial part of the radial carpal. The oblique four main joints the two palmar sesamoid bones.
part, after leaving the styloid process, runs ob Each joint has a joint capsule that runs between
liquely to the palmaromedial surface of the the four bones which form the joint and the two
radial carpal. The tendon of the m. abductor collateral ligaments (ligg. collateralia) which
pollicis longus lies between the two parts as it unite the osseous parts. Each pair of palmar
crosses the medial surface of the carpus. sesamoid bones of the four main joints are joined
The short ulnar collateral ligament (lig. collat together by the intersesamoidean ligament (lig.
erale ulnare breve) extends from the styloid intersesamoideum). This short, cartilaginous lig
process of the ulna to the ulnar carpal. In addi ament consists of transverse fibers which unite
tion to the short collateral ligaments the cranial the paired sesamoid bones and cover their pal
distal lip of the radius is attached to the cranial mar surfaces. The lateral an d m edial sesa-
surface of the ulnar carpal by a strong ligament. m oidean ligaments (ligg. sesamoideum laterale
(Text continued on page 118.)
L ig a m e n t s and J o in t s of the T h o r a c ic L im b 115
P a lm a r c a r p a l fibrocartilacje
M. ext. d i g i f o r u m c o m m u n i s - M. e x t . p o l l i c i s l oncj . t i n d . p r o p r i u s
D i s t a l Hcjcj- ° f acip. c a r p a l M. i n f e r o s s e u s I I
M. f I e x o r d i c j i f o r u m brevis- M. f l e x o r d i g i t o r u m p r o f . t o d t c j i t I
— Dorsal r a d i o c a r p o l lig.
-Short u l n a r c o l l a t er a l lig.
CU
- C o l l at e r al ligg. of p r o x i m a l
metacarpophalangeal j o i n t
Dorsal elastic
- Coll ateral ligg. of distal
interphal angeal j o i n t
S h o r t r a d i a l c o l l a t e r a l ligg.
m a r r a d i o c a r p a l lig.
m a r u l n o c a r p a l lig. Epi phi jses
S h o r t radi al
col l at eral lig— - A r t i c u l a r di sc
Tendon of
Abductor \
p o l l i c i s longus
-Tendon of e x t
di git orum l at
intersesamoidean lig.
Medi al sesamoi dean ligg. F ig . 2-20. Ligaments of forepaw, lateral aspect.
CA = accessory carpal. V = metacarpal V.
Cruciate ligg. of sesamoi d bones
Rad i o u l na r I ig
F ig . 2-19.
Deep ligaments of left forepaw, palmar aspect.
CA = accessory carpal. I to V = metacarpals. Dorsal r ad i oc a r p a l l ig,
Col l at eral l i g of
metacarpophalangeal j o i n t - ■%=-Later al
sesamoidean ligg-
Col l ateral lig. of proximal
interphalanqeal j o i n t -
D o r s a l e l a s t i c ligg.
Collateral lig. of di st al - -t
interphalangeal j o i n t
et mediale) are short, flat bands on each side of cartilage is located on the palmar side of the
the metacarpophalangeal joint. The first part joint capsule. The joint capsule is thickened to
attaches the corresponding lateral and medial form the collateral ligaments, which attach
surfaces of the sesamoid bones to the distal sur proximally to the shallow fossae on each side of
faces of the metacarpal bone caudal to the proxi the head of the middle phalanx and extend ob
mal attachments of the collateral ligaments. The liquely caudodistally to attach to the sides of the
second part goes to the medial and lateral tuber ungual crest of the third, or distal, phalanx. The
cles of the proximal phalanx. From the distal dorsal ligam ents (ligg. dorsalia) are two elastic
ends of each pair of sesamoid bones there is a cords which extend across the dorsal part of the
thin, flat band which attaches to the palmar side distal interphalangeal joint some distance from
of the proximal phalanx. It is called the distal its surface. They attach proximally to the dorsal
sesam oidean ligament (lig. sesamoideum distale). surface of the proximal part of the middle pha
The cruciate ligaments o f the sesam oid bones lanx, where they are about 2 mm. apart. Distally
(ligg. sesamoidea cruciata) extend from the they attach close together on the dorsal part of
bases of the sesamoid bones to the diagonally the ungual crest. They passively keep the claws
opposite tubercles on the proximal end of the retracted, so that the claws do not touch the sub
proximal phalanges. In the first digit there is stratum except when their tension is overcome
usually only one sesamoid bone, and therefore by the m. flexor digitorum profundus.
only one ligament.
The dorsal sesamoid bones of the metacarpo Interdigital Ligaments
phalangeal joints are secured by delicate fibers
from the tendons of the m. extensor digitorum The interdigital ligaments (ligg. interdigitalia)
communis and the mm. interossei proximally, form a continuous superficial, V-shaped liga
and by a ligament to the dorsal surface of the mentous structure which not only holds the dig
middle phalanx distally. its together but also acts as a fastening mecha
nism for the large heart-shaped metacarpal pad.
Phalangeal Joints They originate bilaterally as small fibrous strands
from the abaxial borders of the second and fifth
The proximal interphalangeal joints (articula tendons of the m. flexor digitorum superficialis.
tiones interphalangeae proximales) are formed From their origin proximal to the metacarpo
by the heads of the proximal phalanges articulat phalangeal joints they extend distally to the
ing with the fossae of the middle phalanges in proximal digital annular ligaments which cross
each of the main digits, II to V. These are the flexor tendons at these joints of digits II and
saddle-type joints. The joint capsules have dor V. The interdigital ligaments attach to the proxi
sal walls which are thickened by a bead of carti mal digital annular ligaments of the second and
lage. Here the capsules are intimately united fifth digits and, augmented in size, run disto-
with the extensor tendons so that the sesamoid axially to the proximal digital annular ligaments
cartilages appear to be intercalated in them. On of the third and fourth digits. They attach to
the palmar side the joint capsules are intimately these annular ligaments and again increase in
fused with the flexor tendons. The collateral size, reaching a maximum width of 4 mm. in
ligaments are stout collagenous bands which do large dogs. Continuing distally they unite in a
not parallel the axis through the digit but extend single broad band located dorsal to the meta
in vertical planes as the dog stands. They attach carpal pad. The conjoined interdigital ligaments
proximally to the fossae on the sides of the distal continue to the integument of the pad and cover
ends of the first phalanges and distally to the col the flexor tendons opposite the proximal inter
lateral tubercles on the proximal ends of the phalangeal joints. This is the main supportive
middle phalanges. In the first digit, which has structure of the pad, but there are in addition
only two phalanges, the collateral ligaments several fibro-elastic strands which pass radially
attach distally to the proximal end of the distal into the substance of the pad from the interdig
phalanx. ital ligaments as they cross and are fused to the
The distal interphalangeal joints (articula annular ligaments. Proximal to the interdigital
tiones interphalangeae distales), in the second to ligaments is a feeble collagenous strand which
fifth digits, are formed by the heads of the mid runs from the palmar surface of metacarpal II to
dle phalanges articulating with the saddle a like place on metacarpal V. It is not present in
shaped fossae on the proximal ends of the distal the hindpaw, according to Baum and Zietzsch
phalanges. A single, small, spheroidal, sesamoid mann (1936).
L ig a m e n t s and J o in t s of the P e l v ic L im b 119
—S u p r a s p i n o u s Iicj.
A r t i c u l a r c a p s u l e ---------------- - Y e l l o w lig.
capitis femoris), formerly called the round liga layers of the joint capsule are separated by a
ment, is a rather heavy flattened cord which quantity of fat, the infrapatellar fa t body (corpus
extends from the fovea in the head of the fe adiposum infrapatellare), which increases in
mur to the acetabular fossa. Since it is largely thickness distally. The femorotibial sacs are less
intracapsular and not weight-bearing it is roomy than the femoropatellar. Both femoro
covered by synovial membrane. In large dogs tibial sacs are partly divided by the menisci into
it is about 1.5 cm. long, and 5 mm. wide at its femoromeniscal and tibiomeniscal parts. The
femoral attachment. Its acetabular attachment is fibrosa of the capsule is firmly attached to the
wide as it blends with the periosteum of the discs so that the two parts communicate only
acetabular fossa, and the transverse acetabular around their concave, sharp-edged axial borders,
ligament. The pelvic attachment of the ligament where the tibial and femoral condyles contact
of the femoral head is over 1 cm. wide in large each other. A free transverse communication
dogs. In and peripheral to the rectangular ace also exists between the lateral and medial condy
tabular fossa there is usually a small quantity of loid parts of the joint. Both the lateral and medial
fat. condyloid parts extend between the caudal,
proximal parts of the femoral condyles and the
St i f l e J o in t fabellae that articulate with them. The lateral
femorotibial joint capsule has three other sub
The stifle joint, or knee (articulatio genus) pouches, in addition to the extension between
(Figs. 2-24 to 2-30), is a complex condylar the lateral fabella and femur. One of these ex
synovial joint. The main spheroidal part is tends laterally between the head of the fibula
formed by the thick roller-like condyles of the and the lateral condyle of the tibia to form the
femur articulating with the flattened condyles of proximal tibiofibular joint capsule. This sub
the tibia to form the femorotibial or condyloid pouch is tight-walled since little movement is
part of the joint (articulatio femorotibialis). necessary here. Cranial to this articulation in the
Freely connected with this is the femoropatellar sulcus muscularis of the tibia another pouch ex
joint (articulatio femoropatellaris), located be tends distally about 2 cm. The tendon of the m.
tween the patella and the trochlea of the femur. extensor digitorum longus at its origin from the
The two joints are interdependent in that the extensor fossa of the femur (in the proximal part
patella is held to the tibia firmly by ligamentous of the sulcus muscularis) is ensheathed by syno
tissue so that any movement between the femur vial membrane. The tendon of origin of the m.
and tibia also occurs between the patella and popliteus on the lateral epicondyle of the femur
femur. The incongruence which exists between is never completely surrounded by synovial
the tibia and femur is occupied, in life, by two membrane, but it does possess, on its deep sur
fibrocartilages, or menisci, one located between face, a well defined synovial pouch which acts as
the adjacent medial condyles (meniscus medialis) a bursa.
and the other (meniscus lateralis) between the The lateral and m edial menisci (meniscus
adjacent lateral condyles of the femur and tibia. lateralis et medialis) are semilunar fibrocartilag
The proximal tibiofibular joint is also a com inous discs with sharp, deeply concave axial, and
ponent of the stifle joint. thick convex abaxial borders. The lateral menis
The joint capsule of the stifle joint is the larg cus is slightly thicker and forms a slightly greater
est in the body. It forms three sacs, all of which arc than the medial one. In large dogs the periph
freely intercommunicate. Two of these are be eral border of the lateral meniscus measures
tween the femoral and tibial condyles (saccus about 8 mm. The lateral meniscus does not reach
medialis et lateralis), and the third is beneath the the border of the tibia caudolaterally, but allows
patella. The patellar part of the joint capsule is the tendon of origin of the m. popliteus to pass
very capacious. It attaches to the edges of the over the tibial condyle.
parapatellar fibrocartilages and extends beyond
these in all directions. Proximally a sac protrudes Ligaments of Stifle Joint
1.5 cm. under the tendon of the m. quadriceps
femoris. Laterally and medially the patellar part The meniscal ligaments attach the menisci to
of the joint capsule extends about 2 cm. from the the tibia and femur. Four of these, two from
crests of the trochlear ridges toward the femoral each meniscus, go to the tibia, and one from the
epicondyles. Distally, the patellar and femoro lateral meniscus goes to the femur. The individ
tibial parts join without sharp demarcations. ual meniscal ligaments are as follows:
Distal to the patella the synovial and fibrous The cranial tibial ligament o f the medial
L ig a m e n t s and J o in t s of th e P e l v ic L im b 123
meniscus (lig. tibiale craniale menisci medialis) The cranial or lateral cruciate ligam ent (lig.
goes from the cranial, axial angle of the medial cruciatum craniale) runs between the caudo-
meniscus to the cranial intercondyloid area of medial part of the lateral condyle of the femur
the tibia. This attachment is immediately cranial somewhat diagonally across the intercondyloid
to the intermeniscal ligament, the cranial tibial fossa to the cranial intercondyloid area of the
attachment of the lateral meniscus, and the tibial tibia.
attachment of the cranial cruciate ligament. The caudal or m edial cruciate ligam ent (lig.
The caudal tibial ligam ent o f the medial me cruciatum caudale) runs from the lateral surface
niscus (lig. tibiale caudale menisci medialis) of the medial femoral condyle caudodistally to
goes from the caudal axial angle of the medial the lateral edge of the popliteal notch of the tibia.
meniscus to the caudal intercondyloid area of The caudal cruciate ligament is slightly heavier
the tibia. This attachment is just cranial to the and definitely longer than the cranial one. As
tibial attachment of the caudal cruciate liga their name implies, the cruciate ligaments de
ment. cussate, or cross each other. The caudal cruciate
The cranial tibial ligam ent o f the lateral me ligament lies medial to the cranial one. Being
niscus (lig. tibiale craniale menisci lateralis) intra-articular, they are covered by synovial
goes to the cranial intercondyloid area of the membrane, which, in fact, forms an imperfect
tibia where it attaches caudal to the cranial sagittal septum in the joint. However, this is in
tibial attachment of the medial meniscus. complete, allowing right and left parts to com
The caudal tibial ligam ent o f the lateral me municate.
niscus (lig. tibiale caudalis menisci lateralis) goes Although the patella is a large sesamoid bone
from the caudal axial angle of the lateral menis intercalated in the tendon of insertion of the m.
cus to the popliteal notch of the tibia just caudal quadriceps femoris, it is acceptable to regard the
to the caudal intercondyloid area of the tibia. tendon from the patella to the tibial tuberosity
The fem oral ligament o f the lateral meniscus as the patella ligam ent (lig. patellae). The
(lig. femorale menisci lateralis) is the only fem patellar ligament is separated from the joint
oral attachment of the menisci. It passes from capsule by a large quantity of fat, which is par
the caudal axial angle of the lateral meniscus ticularly thick distally. Between the distal part
dorsally to that part of the medial femoral con of the patellar ligament and the tibial tuberosity,
dyle which faces the intercondyloid fossa. just proximal to its attachment, there is fre
The transverse or intermeniscal ligament (lig. quently located a small synovial bursa. The
transversum genus) is a small transverse fibrous patella is held in the trochlea of the femur
band which leaves the caudal side of the cranial mainly by the heavy lateral femoral fascia, or
tibial ligament of the medial meniscus and goes fascia lata, and the lighter medial femoral fascia.
to the cranial part of the cranial tibial ligament of Aiding in this function are the delicate medial
the lateral meniscus. and lateral fem oropatellar ligaments (lig. fem-
The femorotibial ligaments are the collateral oropatellare mediale et laterale). They are nar
and the cruciate ligaments. The cruciate liga row bands of loose fibers which partially blend
ments o f the stifle (ligg. cruciata genus) are lo with the overlying femoral fasciae. The lateral
cated within the joint cavity. band can usually be traced from the lateral side
The tibial (medial) collateral ligament (lig. of the patella to the fabella in the lateral head of
collaterale tibiale) is a strong ligament which ex the m. gastrocnemius. The medial ligament,
tends between the medial epicondyle of the fe weaker than the lateral, usually blends with the
mur and the medial border of the tibia about 2 periosteum of the medial epicondyle of the fe
cm. distal to the medial tibial condyle. As it mur. The sides of the patella are continued into
passes over this condyle a bursa is interposed be the femoral fascia by the m edial and lateral
tween the ligament and the bone. The total parapatellar fibrocartilages (cartilago parapatel-
length of the ligament is over 4 cm. in medium laris medialis et lateralis). These usually meet
sized dogs. It fuses with the joint capsule and the dorsally. Baum and Zietzschmann (1936) men
medial meniscus. tion a suprapatellar fibrocartilage being present
The fibular (lateral) collateral ligament (lig. in older dogs in the tendon of the m. rectus fem
collaterale fibulare) is similar to its fellow in size oris. The lateral and medial cartilages ride on
and length. As it crosses the joint cavity it passes the crests of the femoral trochlea and tend to
over the tendon of origin of the m. popliteus. It prevent dislocation of the patella.
ends distally on the head of the fibula, with a few Paatsama (1952) has shown that a ruptured
fibers going to the adjacent lateral condyle of the cruciate ligament may be reconstituted by fascia
tibia. lata, which will provide a new functional liga
ment. Free forward movement of the tibia, with
(Text continued on page 127.)
Chapter 2. A rth ro lo g y
Lateral m en iscu s
C a ud al f i b u l a r lig.
P a t e l l a r lig. -
CAUDAL CRANIAL
F ig . 2-24. Ligaments of left stifle joint.
Tendon of g u a d r i c e p s
Patel Ia
Fabellae
Fi bu I a r col I a t e r a l licj
Tibial c o llateral lig
Tendon of popliteus
Cranial fibular lig.
Tendon of long d igita l ext.
— P atellar ligam ent
LATERAL MEDIAL
. v m m \I - - ~ E
onp ip
c uht ys 3
u0r faalc el i ne
F e m o r a l licj. of lat. m e n i s c u s -
n m f
Lateral m en iscu s — ------- L a t e r a l c r u c i a t e licj.
C a u d a l fib. I i g of lat. m e n i s c u s - -j
M edial c ru c ia te lig
--M edial m e n i s c u s (cut)
F ig . 2-28. Cruciate and meniscal ligaments of left stifle joint, medial aspect.
P a f e I l a r l iq . ------
M. r e c t u s f e m o r i s M s a r f o r i us
Mm. vastus l a t e r a l i s t
in te rm e d iu s (fused) — ~M.vastus medialis
-------- P a r a pa t el l ar
fi brocarti lages
--------- P a t e l l a
F a t -------
Patellar lig.-
F ig . 2-30. Parapatellar fibrocartilages, caudal aspect.
L ig a m e n t s and J o in t s of the P e l v ic L im b 127
the joint in extension, is positive evidence of a joint, the distal joint is hardly more than a syno
rupture of the cranial cruciate ligament (Schroe- vial pocket between the lateral malleolus and
der and Schnelle 1941). The cranial cruciate lig the distal lateral surface of the tibia. Besides the
ament is the one most often torn or severed, as a strong connection to the fibular tarsal, fourth
result, usually, of trauma. Hyperflexion is more tarsal, and metatarsal V, by means of the fibular
likely to cause it than other movements. In ex collateral ligament, the lateral malleolus of the
treme instances the tibial collateral ligaments fibula has two ligaments which are proper to the
also may be ruptured. Schreiber (1947) has writ distal tibiofibular joint. The cranial tibiofibular
ten an anatomical treatise on the canine stifle ligament (lig. tibiofibulare craniale) runs a short
joint. distance transversely from the cranial edge of
It has long been known that the removal of all the lateral malleolus to the adjacent lateral sur
or part of a meniscus results in a regenesis of the face of the tibia. The caudal tibiofibular liga
excised part. According to Smillie (1943) and ment (lig. tibiofibulare caudale) runs caudally
Nilsson (1949), the regenerated portion consists from the distal lateral surface of the lateral mal
entirely of connective tissue. By close attention leolus to the lateral surface of the fibular tarsal
to the radii of patellar movement the femoral bone.
trochlea, the femoropatellar ligaments can be
successfully reinforced to prevent chronic luxa T a r s a l , M e t a t a r s a l , a n d P h a la n g ea l
tion of the patella. J o in t s (A r t ic u l a t io n e s P e d is )
Tarsal Joints
T ib io f ib u l a r J o in ts
The tarsal joints (articulationes tarsi) (Figs. 2 -
The fibula articulates with the tibia at each 31 to 2-33), like the carpal joints, are composite
end by small synovial cavities and, in addition, articulations. The talocrural joint (articulatio
possesses an extensive tibiofibular syndesmosis. talocruralis), or ankle joint, permits the greatest
Barnett and Napier (1953) studied the rotatory degree of movement. The trochlea of the tibial
mobility of the fibula in eutherian mammals and tarsal formed largely of two articular ridges fits
concluded that in the dog no rotation could be into reciprocal grooves which form the cochlea
demonstrated on passive movements of the foot. of the tibia. The grooves and ridges are not quite
The proximal tibiofibular joint (articulatio in sagittal planes but deviate laterally about 25
tibiofibularis proximalis) is small and tightly fit degrees so that the open angle faces cranially.
ting; its synovial lining is a distal extension of the This allows the hindpaws to be thrust past the
lining for the lateral femorotibial part of the forepaws on the outside when the dog gallops.
stifle joint capsule. The fibrous layer is not well Due to the presence of the central tarsal bone
developed, although a recognizable spray of in the medial half of the tarsus, the intertarsal
short fibers goes from the head of the fibula prox- joint (articulatio intertarsea) is divided into
imocaudally under the fibular collateral ligament the proximal intertarsal joint (articulatio in
to the adjacent lateral condyle of the tibia as the tertarsea proximalis) and the distal intertarsal
ligament o f the fibular head (lig. capitis fibulare). joint (articulatio intertarsea distalis). The middle
The interosseous m em brane o f the crus (mem tarsal joint of the lateral side and the proximal
brana interossea cruris) extends from the proxi middle tarsal joint of the medial side are coex
mal to the distal tibiofibular articular capsule. tensive; they are formed between the tibial tar
The fibula has many muscles attaching to it, and sal and the fibular tarsal proximally, and be
many of these extend beyond their fibular at tween the central and the fourth tarsal distally.
tachment to the interosseous membrane, which, Some side movement as well as flexion and ex
more than anything else, fastens the fibula to the tension are possible here as the slightly convex
tibia. The fibers which compose this fibrous distal ends of the tibial and fibular tarsals fit into
sheet decussate, forming a lattice-like flat liga glenoid cavities of the central and fourth tarsals.
ment. Proximally an opening exists in the liga The four distal tarsal bones, the first to fourth,
ment for the passage of the large cranial tibial articulate with metatarsals I to V, forming the
artery and its small satellite vein. tarsometatarsal joints (articulationes tarsometa-
The distal tibiofibular joint (articulatio tibio tarseae). The vertical joints between the indi
fibularis distalis) receives an extension of the vidual bones of the tarsus are the intratarsal
synovial membrane from the lateral side of the joints (articulationes intratarseae), all of which
talocrural joint. Like the proximal tibiofibular are exceedingly rigid.
128 Chapter 2. A rth ro lo g y
Tendon of Flexor-
d i g i t o r u m superf.
Cal c a n e a n tendon
T ib ia
F ib u I a Tendon of Flexor
h a l l u c i s longus
-- P r o x i m a l t r a n s v e r s e I ig.
- - T i b i o f i b u l a r lig.
Tendon of Fl exor
-----F i b u l a r c o l l a t e r a l l i g . --------- d i g i t o r u m longus
S u s t e n t a c u t u m t a l i ------ Tendon of _ J|
Fibul ari s longus
— Di st al transverse lig.
Tendon of
Ti bi al i s cran-
Tl -
Tarsal f i b r o c a r t i l a g e -
DORSAL VENTRA l
P r o x i ma l t r a n s v e r s e lig.
F i b u l a r c o l l a t e r o l lig —
short p a r t - -
T i b i al col l at eral l i g . - l an g p a r t -
ihort part
Distal transverse l i g . - -
l on g p a r t
MEDIAL LATERAL
The fibrous part of the tarsal joint capsule taches primarily to the first tarsal and metatarsal
extends from the periosteum proximal to the dis bones.
tal articular cartilage of the tibia and fibula to The fibular collateral ligament (lig. collaterale
the proximal ends of the metatarsal bones. As fibulare), like the tibial collateral ligament, is di
the fibrous layer covers the individual tarsal vided into a long and a short part. The long part
bones and their ligaments it fuses to the free sur passes from the lateral (fibular) malleolus to the
faces of the bones and ligaments. Like the com base of metatarsal V, attaching along its course
parable carpal joint capsule, thickenings are to the fibular tarsal and fourth tarsal. The short
found on both the dorsal and plantar surfaces. part lies under the long part proximally. From
On the plantar surface the fibrous part is deeply the lateral malleolus one band extends to the tu
concave transversely and thickened distally to ber calcanei of the fibular tarsal; a secondhand
form the deep wall of the tarsal canal (canalis goes to the more dorsally located tibial tarsal
tarsea). The tarsal canal contains the tendon bone. Both bands run at nearly right angles to
and sheath of the m. flexor hallucis longus, the the long part of the fibular collateral ligament.
plantar branches of the saphenous artery and On the dorsal surface of the tarsus there are
vein, and the medial and lateral plantar nerves. various short dorsal ligaments. One prominent
The synovial layer of the joint capsule extends ligament unites the tibial tarsal with the third
to the edges of the articular cartilages. There and fourth tarsals. It blends proximally with the
are three lateral and four medial joint sacs. Prox proxim al transverse ligam ent o f the tarsus (lig.
imally the largest sac lines the most freely mov tarsi transversi proximalis), which holds the ten
able joint of the tarsus, the talocrural joint. Dis dons of the mm. extensor digitorum longus, ex
tal to this, in the medial part of the tarsus, are tensor hallucis longus, and tibialis cranialis to the
the proximal and distal middle tarsal sacs. Lat tibia. A small band connects the second and
erally, only a single middle tarsal sac exists be third tarsals. Oblique bands exist between the
tween the fibular and fourth tarsals. Between central and second tarsals, as well as between
the tarsus and metatarsus is the tarsometatarsal the central and third tarsals. The distal row of
sac enclosing the joint which extends between tarsal bones are joined to the proximal ends of
the distal row of tarsal bones, the first to fourth, the metatarsal bones by small vertical ligaments
and the bases of metatarsals I to V. According to on the dorsal surface. A ligamentous loop which
Baum and Zietzschmann (1936), the talocrural attaches to the fibular tarsal surrounds the ten
and the proximal middle intertarsal sacs commu don of the m. extensor digitorum longus. This is
nicate with each other and these communicate the so-called distal transverse ligament o f the
with the synovial sheath surrounding the tendon tarsus (lig. tarsi transversi distalis). On the plan
of the m. flexor hallucis longus. These authors tar surface of the tarsus the special plantar liga
further state that the distal middle tarsal sac ments are heavier than those on the dorsal side.
communicates with the tarsometatarsal sac, but Most of these fuse distally with the thickened
that the two intercommunicating proximal sacs part of the joint capsule at the tarsometatarsal
do not communicate with the two intercommu junction. Several of these ligaments are distinct.
nicating distal sacs. A ligament from the body of the fibular tarsal
The tibial collateral ligam ent (lig. collaterale passes over and is attached to the fourth tarsal
tibiale) (Fig. 2-33) is divided into a long and a on its way to the bases of metatarsals IV and V.
short part. The long, more superficial part is a Another ligament leaves the plantar surface of
rather strong band which runs from the medial the sustentaculum tali, and attaches to and
(tibial) malleolus to attach firmly to the first tar passes over the central tarsal to end in the thick
sal and feebly to metatarsals I and II. As it ened tarsometatarsal joint capsule. Laterally, a
crosses the tarsus it attaches weakly to the free conspicuous band leaves the caudolateral sur
surface of the tibial tarsal and strongly to the face of the fibular tarsal and blends with the long
free surface of the central tarsal. The short part, fibular collateral ligament attached to the base of
attaching craniodistal to the long part, divides as metatarsal V.
it passes under it. One part extends caudally to
attach on the tibial tarsal bone. The other part
is longer and parallels the long part of the tibial M etatarsal and Phalangeal Joints
collateral ligament, caudal to which it lies, after
passing under it. A few fibers may end distally The joints and ligaments of the metatarsus
on the sustentaculum tali of the fibular tarsal and digits are similar to the comparable joints
bone by a fascial connection. However, it at and ligaments of the forepaw.
130 Chapter 2. A rth ro lo g y
M YOLOGY
The muscular system, composed of contractile of adjacent myofibrils are in register with each
units of varied morphology, energized by volun other. Each m uscle fib er is composed of several
tary or involuntary nerve impulses, and nour hundred or several thousand parallel myofibrils,
ished by the blood, constitutes the motive power which also exhibit cross striations. The myofibril
for circulation, respiration, digestion, secretion, is in turn composed of several hundred thick and
excretion, and locomotion, as well as performing thin myofilaments, which consist of the proteins
a host of minor functions. The functional cell myosin (thick) and actin (thin). These myo
unit is known as a m uscle fiber, and it is cus filaments alternate and interdigitate along the
tomary to classify muscle fibers as smooth, length of the myofibril and thus produce the
cardiac, or skeletal. characteristic alternation of the isotropic (I), or
Smooth muscle fibers are spindle-shaped in light, bands and the anisotropic (A), or dark,
form, with a central nucleus. Like other muscle bands. The use of the electron microscope has
cells they possess myofibrils, but they are ho added greatly to knowledge of the significance
mogeneous and not striated. They are found in of the cross bands in contraction and relaxation.
the walls of hollow organs, and in blood vessels, An analysis of myofibril structure is in the realm
as well as in association with glands, and with of protein chemistry. The control of skeletal
the spleen, the eyeball, and hair follicles of the muscles is largely voluntary, although they are
skin. The shortest fibers are about 20 microns capable of involuntary contraction and reflex
long, whereas the longest (in the wall of the movements.
gravid uterus) may be 500 microns in length. For a consideration of structural detail the
Smooth muscle is innervated by the autonomic reader is referred to the texts of Ham and Lee-
nervous system, and in many cases is also under son (1961), or Bloom and Fawcett (1962).
hormonal influence. Other names that have been Bourne (1960) has brought together in three
used for smooth muscle are: unstriated, plain, volumes a considerable wealth of information on
involuntary, white, or visceral muscle. the structure, biochemistry, physiology, pharma
Cardiac muscle fibers form the bulk of the cology, and disease of muscle. The research re
heart. The fibers are arranged in a network of ferred to in these volumes is evidence of the
individual cellular units with intercalated discs wide interest in muscles and the variety of the
between the cell extremities. They exhibit cross unanswered problems.
striations, as do skeletal muscle fibers, and have
centrally placed nuclei like smooth muscle fibers. SKELETAL M U SC LES
Cardiac muscle is capable of rhythmic contrac
tions and is under autonomic control. Specialized The present chapter is concerned primarily
cardiac muscle fibers (Purkinje fibers) serve as a with the named axial and appendicular muscles
conducting system for impulses within the heart. of the body. (Parts of this chapter have been
Skeletal muscle fibers are long, cylindrical, adapted from the Baum-Zietzschmann Anato
multinucleated cells organized into distinct mie des Hundes, by permission of the Publisher,
bundles within connective tissue envelopes. Paul Parey, of Berlin and Hamburg.) In mam
Other names applied to skeletal muscle include malian species the skeletal muscles comprise
striated, voluntary, or somatic muscle. The cells approximately one-third to one-half of the total
appear striated because the light and dark bands body weight. They range in size from the minute
131
132 Chapter 3. M yology
stapedius muscle of the middle ear to the large neuroses but attach directly to the periosteum of
gluteus medius muscle of the rump. bones. Such origins or insertions are spoken of as
Each muscle fiber is surrounded by a thin fleshy attachments. The more fixed point of mus
sarcolem m a and a delicate connective tissue cle attachment is spoken of as the origin; the
sheath known as the endomysium. When several more movable point of attachment is called the
fibers are grouped into a fasciculus they are en insertion. In the limb the insertion of a muscle is
closed by perimysium. The definitive muscle is always considered to be distal to its origin, al
composed of several fasciculi wrapped by an though functionally it may be the most fixed
epimysium, which delimits one muscle from an point at some phase of the stride. Certain mus
other or occasionally fuses with the intervening cles have equally fixed or mobile attachments,
fascia. The size of an individual muscle fiber de and the naming of an origin and an insertion is
pends on the species as well as on the physical rather arbitrary.
condition of the animal, since individual muscle The expanded fleshy portion of a muscle is
fibers are capable of hypertrophy as well as its belly, the origin is a head, and minor inser
atrophy. tions are called slips. A muscle may have more
Conflicting statements have appeared in re than one belly (digastric) or more than one head
gard to the length of a muscle fiber within a (triceps), and frequently has several insertions.
muscle. Lockhart and Brandt (1938) found mus
cle fibers running the entire length of the sar
torius muscle (5 cm.) in a human fetus, and were Function
able to isolate fibers 34 cm. long in a 52 cm. sar
torius muscle of an adult. Huber (1916) and Van Muscles which attach to long bones (the
Harreveld (1947), working with rabbit thigh levers) and span one or more joints usually work
muscles, found that many fibers do not extend at a mechanical disadvantage. When a muscle
from end to end. They concluded that, although fiber contracts it does so at its maximum power
the longer fasciculi have longer fibers, many and it is capable of contracting to about half of
fibers end intrafascicularly. its stretched length. The contraction is initiated
Muscles take diverse shapes and are usually by a nerve impulse traveling over a motor nerve
named according to some structural or func fiber (axon) to the muscle fibers, or cells. Each
tional feature, although other criteria have also axon supplies several muscle fibers. These neuro
been used. The variations encountered in the muscular units are known as motor units, and
muscular system within a species are numerous the number of motor units functioning at any
and may constitute a breed-specific feature. one time determines the activity of the muscle.
Huntington (1903) considered problems of gross If a muscle has many motor units, each of which
myological research and the significance and includes only a few muscle fibers, then the pre
classification of muscular variations. The most cision of movement is great (as in the extrinsic
complete account of the muscles in the dog is by muscles of the eyeball).
Baum and Zietzschmann (1936). Other sources It is important to study and experiment with
include Miller (1958), Ellenberger and Baum muscles in the living body to appreciate the full
(1943), Bradley and Grahame (1959), Nickel, significance of precise muscular movement and
Schummer, and Seiferle (1954), and Sisson and the value of such movement in a neurological
Grossman (1953). The muscles of the cat were examination for the determination of intact or
described in detail by Straus-Durckheim (1845) defective nerve supply. Electromyography, as
and less completely by Reighard and Jennings shown by Basmajian (1962), is an excellent tech
(1935). nique for studying living muscles.
Of two muscles of equal size and shape, the
Origin and Insertion muscle which contains the greater number of
fibers is the stronger. Although a muscle fiber
Most skeletal muscles are attached by con can shorten to about half of its length, never do
nective tissue to a bone or cartilage. Some are whole muscles contract to this degree. Straplike
attached to an organ (eye, tongue), to another and sheetlike muscles contract to a greater de
muscle, or to the skin; others lie free beneath gree than do those of the extremities. Muscles
the skin and act as sphincters. The connective possessing tendons throughout all or part of their
tissue attachment may be in the form of a cord length are known as pennate muscles. A muscle
like tendon or a flat sheetlike aponeurosis. Some with a tendon running along one side is called
muscles have no demonstrable tendons or apo unipennate; if there is a tendon on each side of
Sk eleta l M u sc les 133
the muscle, it is bipennate; when a muscle is in oris muscle. Sesamoid bones serve three im
vaded by tendons in several places it is multipen- portant functions: (1) they protect tendons
nate. Pennate muscles are stronger (exert more which pass over bony prominences; (2) they in
force) than straplike or sheetlike types because crease the surface area for attachment of ten
they are composed of many short oblique fibers dons over certain joints; and (3) they serve to
which have an additive effect upon the insertion. redirect the pull of tendons so that greater effec
Because of their elasticity, tendons protect mus tive force can be applied to the part being
cles from sudden strains. Fleshy, rectangular moved.
muscles usually attach farther from the fulcrum Bursae are simple connective tissue sacs con
than do pennate types; they also move their in taining a viscous fluid and serving to reduce fric
sertions farther, but with less speed. tion. They are usually located between a tendon,
Muscles which straighten bone alignment, or ligament, or muscle, and a bony prominence.
open a joint, are called extensors; those which Occasionally they are located between tendons
angulate the bones, or bend the joint, are known or between a bony prominence and the skin. In
as flexors. Flexion and extension are the primary constant bursae may develop at various sites in
movements necessary for locomotion. Accom response to undue friction, and, conversely, cel
panying movements include adduction, or the lular proliferation due to infection or trauma
movement of an extremity toward the median may eliminate them.
plane; abduction, or movement away from the Synovial tendon sheaths are double-layered,
median plane (in the case of the digits the refer elongated sacs containing synovia which wrap
ence point is the axis of the limb); circumduc tendons as they pass through osseous or fibrous
tion, or moving an extremity in a plane describ grooves. The inner layer of the sheath, which is
ing the surface of a cone; and rotation, or mov fused to the tendon, attaches to the wall of the
ing a part around its long axis. The pattern of passageway and is known as the mesotendon.
movement resulting from muscle contractions, Blood vessels and nerves enter the tendon via
even for apparently simple movements, is the mesotendon. The tendon sheath with its con
brought about by the complex interactions of tained synovia serves for reducing friction dur
many muscles. The characteristic movement of ing movement.
a joint is produced by a muscle or muscles, Fascia is connective tissue which remains
called prime movers, or agonists. The muscles after the recognizable mesodermal structures
responsible for the opposite action are known as have been differentiated in the fetus. It serves
antagonists, although they actually aid the many important functions, and has considerable
prime mover by relaxing in a controlled manner clinical significance. For descriptive purposes it
so that the movement will be smooth and pre is convenient to distinguish many fascial entities
cise. For the elbow joint, the prime mover in which envelop, separate, or connect muscles,
flexion is the biceps brachii; the antagonist is the vessels, and nerves. Fascial sheets provide routes
triceps. Conversely, in bringing about extension, for the passage of blood vessels, lymphatics, and
the prime mover is the triceps. Fixation and ar nerves, as well as serving for the storage of fat.
ticular muscles are those which stabilize joints The superficial fascia beneath the skin is closely
while the prime movers are acting. Synergists associated with the dermis, and often includes
are fixation muscles which stabilize intermediate cutaneous muscle fibers. The deep fascia which
or proximal joints and enable the force of the covers and passes between the muscles is partic
prime mover to be exerted on a more distal joint. ularly tough and distinct in the limbs. It func
tions as a sleeve within which the muscles can
Accessory Structures operate, and often serves as an aponeurosis of
origin or insertion. In certain locations fascia
Associated with muscles are accessory struc blends with the periosteum of bone, forming
tures of great physiological and clinical impor interosseous membranes or annular bands which
tance, such as sesamoid bones, bursae, synovial confine tendons or redirect their force. Most
tendon sheaths, and fascia. commonly, distinct fascial septa separate groups
Sesamoid bones are located in certain tendons of muscles from one another and result in fascial
or joint capsules as small rounded nodules. Oc planes along which infection may spread or
casionally they develop in response to friction, fluids drain.
but usually they form prenatally. The patella, or
knee cap, is an example of a large sesamoid bone Connective Tissue
in the tendon of insertion of the quadriceps fem The amount of connective tissue present is
134 Chapter 3. M yology
by many fiber bundles with the underlying Innervation: Rami buccalis dorsalis et ven
sphincter colli profundus. The ventral border tralis, n. facialis.
has a distinct boundary. Only rarely does the The m. maxillonasolabialis (Figs. 3-2, 3-3, 3 -
platysma have defects. 4) has developed from the upper lip portion of
Action: To draw the commissure of the lips the orbicularis oris. It corresponds to the m. leva
posteriorly. tor labii superioris alaeque nasi of the primates.
Innervation: Rami buccalis dorsalis et ven In the dog it is distinctly separated into nasal
tralis, and ramus auricularis posterior, n. and labial portions, which are both covered by
facialis. the levator nasolabialis.
The m. sphincter colli profundus includes The stronger pars nasalis is a flat muscle, only
muscles of the cheek, lip, and external ear. It is 2 mm. thick in large dogs, which lies beneath the
divided into a pars oralis, pars palpebralis, pars apical end of the levator nasolabialis on the max
intermedia, and pars auricularis. illa and incisive bone. The nasal part arises cau-
Pars oralis. The pars oralis of the sphincter doventral to the infraorbital foramen deep to the
colli profundus includes the mm. orbicularis oris, palpebral part of the sphincter colli profundus.
incisivus dorsalis et ventralis, maxillonasolabialis, The fibers of insertion spread out as they enter
buccinatorius, and mentalis. the nasal ala and the upper lip.
The m. orbicularis oris (Figs. 3 -1 , 3-2), the The weaker pars labialis is immediately ven
principal component of the lips, extends from tral to the nasal portion and separates from it, in
the commissural region into the lips near their that it extends over the labial end of the levator
free borders. In the median segment of both lips, nasolabialis into the upper lip.
the muscle is interrupted. It lies between the skin Action: To increase the diameter of the exter
and mucosa. The other muscles of the lips and nal naris, and lift the apical portion of the
the muscles of the cheeks (platysma, and mm. upper lip.
buccinatorius, zygomaticus, and levator nasola- Innervation: Ramus buccalis dorsalis, n. faci
bialis) enter the m. orbicularis oris so that here alis.
these muscles blend with each other; the m. The m. buccinatorius (Figs. 3-2, 3-3) has de
incisivus is also attached to it. The portion of veloped from the deep part of the orbicularis
the orbicularis oris lying in the upper lip is the oris. It is a strong, flat, wide muscle which forms
stronger; separate fibers extend from it to the ex the foundation of the true cheek. It is composed
ternal naris. of two portions, which extend caudally from the
Action: The muscle closes the mouth opening labial commissure. Only a superficial portion
and is a pressor of the labial glands. Of the proceeds in an arch from one lip into the other,
bundles extending to the lateral nasal car like the orbicularis. The upper portion has incor
tilage on either side, the medial ones act to rectly been called the m. buccalis, the lower the
pull the entire nose downward (in sniffing), m. malaris.
and the lateral bundles act to increase the The pars dorsalis, or the m. buccalis of de
diameter of the external nares. In strong scriptive nomenclature, is the somewhat stronger
contractions both the medial and lateral fi portion. It arises from the upper lip and second
ber bundles function to dilate the external arily from the region caudal to the infraorbital
nares. foramen on the alveolar border of the maxilla.
Innervation: Rami buccalis dorsalis et ven It is superficial as far as the raphe. It proceeds
tralis, n. facialis. posteroventrally and, except for a narrow pos
The mm. incisivus dorsalis et ventralis lie terior portion, crosses through the ventral por
deep to the orbicularis oris. These are two thin tion. After running deeply, it ends in three or
muscles not clearly defined from the orbicularis four distinct portions in such a way that the oral
and buccinatorius; they arise on the alveolar bor portion remains superficial and proceeds sphinc
ders of the incisive bone and mandible as far as ter-like into the lower lip. This portion runs be
the corner incisor teeth, and are situated imme neath and parallel to the orbicularis oris, and can
diately beneath the mucosa of the lips. They ex be isolated in carefully prepared specimens. Be
tend to the orbicularis oris. According to Huber tween the two, the anterior end of the m. zygo
(1922), the lower one cannot be isolated as a sep maticus enters. The other portions of the pars
arate muscle. dorsalis of the buccinatorius pierce the ventral
Action: The m. incisivus dorsalis raises the up part and cross beneath the latter in a postero
per lip. The m. incisivus ventralis pulls the ventral direction to end more or less independ
lower lip downward. ently on the mandible. The posterior branch
(Text continued on page 139.)
136 Chapter 3. M y o lo g y
Trac/oh el i c i n u s ,
HeI i c is
S pine of h e lix
S c u ti form c a rtila g e
Orbicularis oris u
S p h i n c t e r c o l I i prof. —p a r s p a l p e b r a l i s
PI a t t-jsma 1
Zijcjoma t i cu s 1
S p h in c te r c o lli p rof. - pa rs i n t e r m e d i a
V\
F r o n t a l is \ A poneurosis
Maxi l i o n a so l a b i a l i s , I tCervicaauri c u l a r i s
- p a r s n a s a I is v \ ' I prof. m in or
- p a r s i a b i a l i s NN ^ v Te mp o r a l i s
Ment a I is
Orbicularis o r is 1
Buccinator - o r a l p o rtio n
Sphincter c o l l i prof. - pa rs pa I p e b r a l is
P a r o tid duct
M a n d ib u la r lymph n o d e s 1
Pa r o t i d g l a n d 1
M a n d I b u la r g l a n d '
Ext. j u g u l a r v.'
F ig . 3-2. Superficial muscles of the head, lateral aspect. (Platysma and sphincter colli superficialis removed.)
138 Chapter 3. M y o lo g y
/ I n f e r io r p a r t of cervicoauricuiaris superficialis (O c c i p i t a l i s
Interscufu Ia ri s Cervi c o s c u t u I a r i s
i
Scufu l o a u r i cu i a r i s super f . a c c e s s o r i u s ( j 11 n t e r p a r i e f o s c u t u l a r i s
S c u f u l o a u r i c u l a r i s superf. m e d i u s ] J J I Int erpari et oaur i cul ari s
t I i
S c u t u l o a u r i c u l a r i s superf. d or s al is ^ i| i I i j i Cervi coauri cuiari s
Frontalis ' I 1 i I prof. maj or
■us v 1 1I I1
1 I iCervicoauricuiaris
Superci H a ris \ , .
1 I i p r o f mi n o r
' ' \ \ 1 1I I
O rbicularis oculi, ' \ \ \ t i 1 1
' \ x ..........I,__i_i i 1I I /Cervicoauric.
L e v a t o r n a s o l a b i a l is, ' \ \
I ( j I / superficialis
M axillonasolabialis. \ I 1 / / Tempor al i s
- p a r s nasalis \ '
-pa r s l a b i a l i s . \
Mental is /
0 r b i cu I a ri s oris
B u c c i n a t o r - o r a l portion
B u c c i n a t o r —d o r s a l p o r t i o n { Ext . j ucj ul ar v.
M a n d i b u l a r buccal g l an d Depressor a u r i c u l a e
Masseter S p h i n c t e r c o l l i p r o f . - pars i n f e r m e d i a
F ig . 3-3. Deep muscles of the head and ear, lateral aspect.
M u sc les of the H ea d 139
runs almost horizontally and passes beneath the tral mid line to the lower lid. At the level of the
masseter. Furthermore, as already mentioned, dorsal border of the platysma, it is divided into
a little muscle bundle separates from the un two portions by the pressure of the platysma: (1)
crossed part of the posterior border; this arches a ventral portion, covered by the platysma, and
posteriorly and in some animals extends super (2) the free dorsal segment. The muscle is found
ficially over the anterior border of the masseter. between the pars oralis and pars intermedia of
The pars ventralis is the weaker portion of the the sphincter colli profundus, and covers the an
buccinatorius. It arises from the lower lip and, terior end of the m. zygomaticus and the m. buc
secondarily, in the region of the three lower mo cinatorius. In the lower eyelid it extends upon
lars, from the alveolar border of the mandible. It the orbicularis oculi.
proceeds beyond the caudal edge of the dorsal Action: To depress the lower lid.
portion. The anterior portions end on the maxilla Innervation: Ramus buccalis ventralis, n. faci
although most of the fibers pass beneath the mas alis.
seter. Pars intermedia. The pars intermedia of the
The fibers of the dorsal and ventral portions sphincter colli profundus (Fig. 3-2) is the contin
of the buccinatorius, disappearing beneath the uation of the palpebral portion and consists of
edge of the masseter, fuse into a loose delicate loose muscular strands which arise in the inter-
muscle lying directly upon the mucous mem mandibular region behind those of the palpebral
brane of the cheek. Closely attached to the buc portion. There the two portions extend over to
cal mucosa, they also attach on the maxilla and the other side and cross each other. Toward the
mandible peripheral to the edge of the mucosal anteroventral border of the scutulum on which
covering. Functionally, this is the origin of the they end, the strands are closer together. The
muscle. This attachment may be circumscribed superficial strands occasionally arise from the
by the following lines: the alveolar border of the superficial surface of the platysma at its dorsal
mandible in the region of the last two cheek border. From the anterior margin of the pars
teeth, the anterior edge of the coronoid process intermedia the long, broad muscular band which
to half its length, and the alveolar border of the proceeds toward the angle of the mouth is the m.
maxilla in the region of the last two cheek teeth. zygomaticus (auriculolabialis).
The “free” portions of the buccinatorius lying The m. zygomaticus (Figs. 3 -1 to 3-4), a de
anterior to the masseter also surround the ven rivative of the pars intermedia of the sphincter
tral buccal glands with fibrous masses medially. colli profundus, is usually intimately fused with
At the level of the third maxillary cheek tooth, the intermediate part at the scutulum; only ex
the parotid duct perforates the buccinatorius ceptionally can both be completely isolated at
near its dorsal border. this point. The straplike, long muscle extends
Action: To return food from the vestibule to from the anterior angle of the scutulum to the
the masticatory surface of the teeth. edge of the mouth, where it sinks into the or
Innervation: Rami buccalis dorsalis et ven bicularis oris after crossing beneath the palpe
tralis, n. facialis. bral portion of the sphincter colli profundus. Its
The m. mentalis is an incomplete division of apical portion is deep and bears no relationship
the ventral portion of the m. buccinatorius. It to the platysma. Proximally it is distinctly sepa
arises from the alveolar border and body of the rated from the m. frontalis, which arises from a
mandible near the corner teeth. The fibers unite fascia at the deep surface of the scutulum; this
with those of the opposite side and radiate into portion of the m. zygomaticus is covered by the
the lower lip, forming a strong, fat-infiltrated skin.
muscle. Action: To fix the angle of the mouth and draw
Action: To stiffen the lower lip in the apical re it back, or to fix and draw the scutulum for
gion. ward.
Innervation: Ramus buccalis ventralis, n. faci Innervation: Ramus auriculopalpebralis, n.
alis. facialis.
Pars palpebralis. The pars palpebralis of the
sphincter colli profundus has incorrectly been Muscles of the Forehead and the Dorsum of
identified with the m. malaris of man. It is found the Nose
adjacent to the caudal portions of the pars oralis
and is a thin muscle consisting of separated, deli Dorsal to the pars intermedia of the sphincter
cate muscular strands extending in front of the colli profundus is a large homogeneous muscle
masseter (beneath the platysma) from the ven mass, the orbitofrontoauricular muscle leaf. This
140 Chapter 3. M yology
extends from the concha over the scutulum to The m. superciliaris is a small, strong muscle
the forehead and the eyelids; it sends further de strand which arises from the median line on the
rivatives between the orbits to the nose and up frontal bone from the nasofrontal fascia. It ex
per lip. tends over the orbicularis oculi into the medial
The preauriculur muscles, the m. frontalis, the half of the upper lid. It passes through portions
muscles of the lids, and the m. levator nasolabi of the upper lid which bear the hairs (pili supra-
alis belong to the orbitofrontoauricular muscle orbitales) designated as the eyebrow.
complex (Figs. 3-1 to 3-4). Action: To lift the upper lid, especially its
The m. frontalis is a thin muscle which lies on nasal portion, and erect the hairs of the eye
the temporalis. It arises in front of the anterior brow.
border of the scutulum, beneath the pars inter Innervation: Ramus auriculopalpebralis, n.
media of the sphincter colli profundus, by means facialis.
of a fascial leaf, and extends to the forehead and The m. levator nasolabialis is the most ante
toward the upper eyelid. The frontal portion, rior extension of the original muscle sheet (pars
unseparated posteriorly from the m. interscutu- intermedia of the sphincter colli profundus) that
laris, unites with that of the opposite side and passes from the ear past the eye to the muzzle.
anteriorly joins the nasofrontal fascia by which It is a very flat, thin, and broad muscle (even in
it attaches to the zygomatic process. The palpe large dogs), lying immediately beneath the skin
bral portion passes beneath the orbicularis oculi on the lateral surface of the nasal and maxillary
and attaches to the orbital ligament. From the bones. It arises in the frontal region between the
concha a considerable number of muscle strands orbits from the nasofrontal fascia, the nasal pal
of the m. scutuloauricularis superficialis dorsalis pebral ligament, and the maxillary bone; occa
extend over the scutulum into the frontalis. It is sionally a few additional fibers come from the
seen from this relationship that these two muscle lacrimal bone. Spreading out, it proceeds to the
groups belong together. The m. frontalis has also nose and upper lip to push beneath the orbicu
been called the m. frontoscutularis. In the dog laris oris. The posterior portion inserts on the
it has become attached to the frontal bone and buccinatorius; the apical, larger portion passes
the orbital ligament. It has been completely beneath the orbicularis partly between both por
separated from the mm. retractor anguli oculi, tions of the m. maxillonasolabialis, to end near
superciliaris, and levator nasolabialis. the edge of the lip. The most dorsal and anterior
Action: To fix and pull the scutulum forward. fibers, however, force their way between the fi
Innervation: Rami temporalis and auriculopal- bers of the nasal portion of the maxillonasolabi
pebralis, n. facialis. alis and so break it up into separate leaflike
The m. orbicularis oculi surrounds the palpe layers in gaining attachment to the external
bral fissure. Portions of the muscle adjacent to naris. These fibers cannot be considered a spe
the borders of the lids extend from the medial cial m. nasalis (Boas and Paulli 1908). The spe
palpebral ligament over the upper lid, around cific mm. nasales of other mammals (ungulates,
the temporal angle of the lids, and along the proboscidae, primates) are divisions of the pars
lower lid back to the ligament. Thus in the dog, oralis of the sphincter colli profundus. Further
this muscle, which originally was divided into divisions of the m. levator nasolabialis, which
dorsal and ventral portions, has become one. would serve for special nasal movements, do not
Huber (1922) states that the ventral portion exist in the dog. At most, one finds a flat, thin di
comes from the m. zygomaticus, and the dorsal lator of the naris which arises on the anterodor-
portion comes from the m. frontalis. sal portions of the nasal cartilage and extends
Action: To close the palpebral fissure. into the lateral nasal ala. It is noticeable only
Innervation: Ramus auriculopalpebralis, n. in large dogs.
facialis. Action: To increase the diameter of the naris,
The m. retractor anguli oculi, as a division of and lift the apical portion of the upper lip.
the m. frontalis, arises beside the latter from the Innervation: Ramus auriculopalpebralis, n.
temporal fascia. It extends horizontally to the facialis.
lateral palpebral angle, and, in so doing, it
crosses the orbicularis oculi before it sinks into M u sc les o f th e E xtern al E ar
the fibers of the latter.
Action: To draw the lateral palpebral angle The muscles of the ear, from comparative and
posteriorly. ontogenetic viewpoints, fall into three groups.
Innervation: Ramus auriculopalpebralis, n. The preauricular group is a derivative of the pars
facialis. intermedia of the sphincter colli profundus; the
M u sc les of the H ead 141
O c c i p i fa utularis
w %
J //
uricularis superf.
Cervicoauric ula ris prof. m a j o r N
A n t e r i o r p a r t of
c e r v i c o a u r i c . superf.
I n t e r p a r i e t o s c u t u la r i s \
\ ! 'J
\ /Scutuloauricularis
I n t e rpa r i e t o a u r i cu laris^ superf. accessorius
Scutul o au r i cu l a r i s
superf. medi us
~ S c u f i f a r m cartilage
■S c u f u l o a u r i c u i a r i s
superf. d o rsa lis
Zt j qo ma t icoauri cu l ari s
Z y g o ma t i cu s v
S p h i n c t e r c o l l i prof .
—p a r s i n t e r m e d i a
s R e t r a c t o r ancju li o cu li
vOrbicularis o c u l i
\ S u p e rciIia ri s
Levator n a s o la b ia lis
Max i 11 o n a s o l a b i a I i s
- p a r s la b i a I is '
—p a r s n a s a l i s /
Fic. 3-4. Deep muscles of the head and ear, dorsal aspect.
142 Chapter 3. M yology
ventroauricular group is represented by the pars has arisen from the fusion of bilateral portions.
auricularis of the sphincter colli profundus; and The origin is from the entire dorsomedial border
the postauricular group is a derivative of the deep of the scutulum. The posterior portion of the
portion of the platysma of the neck. muscle has a distinct border and covers the m.
occipitalis and the m. cervicoscutularis, both of
Preauricular Muscles which blend with the interscutularis. Anteriorly
it has no distinct border and encroaches upon
Because of the position of the scutulum in the the frontal portion of the m. frontalis.
dog, the m. scutuloauricularis superficialis dor Action: Fixation of the scutulum.
salis, the zygomaticoauricularis, and the intrinsic The m. scutuloauricularis profundus major,
muscles—the mm. tragohelicinus, tragotubo- or large rotator of the concha (Fig. 3-5), is com
helicinus, and conchohelicinus—have all become pletely separated from the m. frontalis to which
completely separated from the orbitofronto- it belongs. As a strong, well defined muscle, it
auricular muscle complex. The m. interscutularis lies beneath the scutulum and arises on its deep
and the m. scutuloauricularis profundus major surface to extend to the concha over the m.
are also distinct. All of these preauricular mus temporalis. The muscle has an almost sagittal
cles are innervated by the ramus temporalis, n. course as it crosses the scutuloauricularis pro
facialis. fundus minor on its deep surface.
The m. scutuloauricularis superficialis dor Action: To turn the conchal fissure backward.
salis (m. auricularis anterior superior of Huber)
(Figs. 3-1, 3-3, 3-4) is the dorsal inward rotator Ventroauricular Muscle
of the ear. It separates dorsoposteriorly from the
m. frontalis, with which it is always partly The m. depressor auriculae (parotideoauricu-
united. It is a broad, strong muscle lying free laris of Huber) (Fig. 3-2) arises posterior to the
over the anterior portion of the scutulum and laryngeal region, on or near the midline, where
coursing in a fold of skin to the medial border of it blends with the cervical fascia. As a strong,
the concha, where it attaches in the region of the well defined band, it runs obliquely toward the
spine of the helix (crus helicis distale). concha, crossing the mandibular and parotid
Action: To turn the conchal fissure forward. glands in its course. The muscle is almost com
The m. zygomaticoauricularis (Figs. 3-1, 3-2, pletely covered by the platysma and inserts on
3-4) is the external inward rotator that arises as the antitragus.
a rather broad muscle from the tendinous leaf Action: To depress the ear.
lying in front of the scutulum. It is continuous Innervation: Ramus colli, n. facialis.
anteriorly with the lower division of the frontalis.
Posteriorly, it extends ventrally to the basal por Postauricular Muscles
tion of the tragus.
Action: To turn the conchal fissure forward. The postauricular muscles are derivatives of
The mm. tragohelicinus, tragotubohelicinus, the platysma and are innervated by rami post-
and conchohelicinus lie together in one muscle auriculares of the facial nerve. The muscles
complex, which bridges the space between the forming this complex consist of the cervicoauric-
superimposed conchal edges of the conchal ular musculature, the posterior conchal muscles,
canal. This muscle aggregate passes from the and the m. mandibuloauricularis.
deep surface of the lateral crus of the helix to the The cervicoauricular musculature is a con
tragus. In certain cases all of these muscles are tinuation of the deep layer of the platysma in the
independent. See Leahy (1949) for further pic region of the neck. It is divided into three layers
torial and descriptive treatment. The m. trago which, in the dog, are not completely separated.
helicinus arises from the external surface of the The superficial layer, m. cervicoauriculo-
tragus, the m. tragotubohelicinus from the tragus occipitalis, forms a completely homogeneous
and the conchal canal, and the m. concho muscle plate which comes from the dorsum of
helicinus from the external surface of the con the neck and the occipital bone. It inserts on the
cha. caudal border of the scutulum as well as on the
Action: To narrow the entrance to the conchal m. interscutularis and nasofrontal fascia. In this
canal and thus make the concha rigid. muscle complex are contained the m. cervico-
The m. interscutularis (Fig. 3-3) is a thin auricularis superficialis, the m. occipitalis, and,
muscle extending from one scutulum to the as a connecting link between the two, the m.
other, without attaching to the cranial bones. It cervicoscutularis.
M u sc les of th e H ea d 143
The in. cervicoauricularis superficialis, or temporalis of that region. Primitively, the auric
long levator (Fig. 3-4), arises from the cervical ular end of this muscle is simple. There are cases,
mid line and the external occipital protuberance. however, in which this end is double.
As a broad muscle mass it passes to the concha Action: To turn the conchal fissure outward
and ends by two branches on the dorsum of the and backward.
ear. The anterior branch is made wider by fibers The m. interparietoscutularis (Figs. 3-3, 3 -
from the lateral border of the scutulum; these 4) is only exceptionally an independent muscle.
correspond to the m. scutuloauricularis super It is described by Huber (1923) as the m. cervi
ficialis accessorius, or short levator, of other ani coscutularis medius belonging to the middle
mals. The posterior branch covers the auricular layer. It arises from the interparietal portion of
end of the interparietoauricularis. the external sagittal crest and inserts on the
Action: To raise the concha. caudal border of the scutulum, which is com
The m. cervicoscutularis (cervicointerscutu- pletely covered by the superficial layer of this
laris of Huber) (Figs. 3-3, 3-4) is a narrow, muscle complex. Ordinarily this muscle is united
intermediate portion of the muscle complex with the m. interparietoauricularis almost as far
which is not clearly defined; it goes to the pos as the scutulum.
terior border and the posteromedial angle of the Action: With other scutular muscles, it aids in
scutulum, and is united with the deep surface of fixation of the scutulum.
the interscutularis by means of a few fibers. The m. scutuloauricularis profundus minor,
Action: To draw the scutulum downward, or or short rotator (Fig. 3-5), is considered by
fix it when the scutulum is drawn forward Huber (1922) to be a special branch of the
at the same time. scutuloauricularis medius. Owing to the presence
The m. occipitalis (Figs. 3-3, 3-4) is the third of the scutulum, the short rotator is interrupted
portion of the cervicoauriculo-occipital complex. as a continuation of the m. interparietoscutularis.
From the external sagittal crest, its fibers turn From the deep surface of the scutulum near the
anteriorly in bilaterally symmetrical arches in lateral angle it descends in a somewhat vertical
such a way that they form an unpaired, oval, direction to the concha. At the same time, this
thin membranous muscle which can be followed small short rotator crosses the large rotator which
a short distance forward beneath the posterior belongs to the preauricular muscle group. It
portion of the m. interscutularis; there, on the inserts on the external surface of the lateral crus
frontal bone, they spread out into the naso of the helix opposite the attachment of the m.
frontal fascia. mandibuloauricularis.
Action: To tense the nasofrontal fascia. Action: To turn the conchal fissure laterally.
The middle and deep layers of the cervico- The m. scutuloauricularis superficialis acces
auricular musculature in the dog are divided into sorius, or short levator (Figs. 3-3, 3-4), is the
a number of individual muscles. Of these the m. other part of Huber’s m. scutuloauricularis
cervicoauricularis profundus major, the m. inter - medius, as the second indirect continuation of
parietoscutularis, and the three parts of the m. the m. interparietoscutularis. It arises from the
scutuloauricularis belong to the middle layer, posterior portion of the lateral scutular border
while the m. interparietoauricularis and m. cer with the anterior segment of the m. cervico
vicoauricularis profundus minor belong to the auricularis superficialis. It inserts basal to the
deep layer. long levator or cervicoauricularis superficialis.
The m. cervicoauricularis profundus major, Action: With the other levators, erection of
or long outward rotator (cervicoauricularis the concha.
medius of Huber ) (Figs. 3 -4 , 3-5), is a strong, The m. scutuloauricularis superficialis medi
relatively wide muscle which, covered partly by us, or middle inward rotator (Figs. 3-3, 3-4, 3 -
the cervicoauricularis superficialis, arises on the 5), is, according to Huber (1922), a branch of
external sagittal crest, the external occipital the m. scutuloauricular medius and comes
protuberance, and the neighboring attachment from the platysma. It extends from the deep sur
of the nuchal ligament. It extends to the base of face of the caudal half of the lateral edge of the
the concha and finally ends on the root of the scutulum to the lateral crus of the helix. Anterior
lateral conchal border (antitragus), where it lies to the short levator, it lies deeply between the
next to the insertion of the depressor auriculae. scutulum and the base of the concha.
This muscle covers a portion of the origin of the Action: To turn the conchal fissure forward.
interparietoauricularis, the greater part of the The m. interparietoauricularis, or middle
cervicoauricularis profundus minor, and the m. levator (cervicoauricularis profundus anterior of
144 Chapter 3. M yology
Huber) (Fig. 3-4), is only seldom completely cha to the medial edge thereof, indicates
isolated. Indeed, it belongs to the deep layer, that the m. helicis belongs to the postauric
but usually fuses with the m. interparietoscu- ular muscle group. The same branch sup
tularis, which becomes separate only near the plies the mm. obliqui et transversi, the short
scutulum. In its entire course it is covered by the levator, and the m. mandibuloauricularis.
superficial layer of the postauricular muscula The m. mandibuloauricularis (Figs. 3-5, 3 -
ture. It arises from the interparietal segment of 11) is a muscle of the auditory canal. It is a long,
the external sagittal crest and goes directly over narrow muscle, which also bears the name m.
to the dorsum of the concha, where it attaches tragicus lateralis in descriptive nomenclature. It
under the posterior terminal branch of the m. arises tendinously in the niche between the
cervicoauricularis superficialis basal to its inser angular and condyloid processes of the mandible
tion. and extends dorsally to the lateral crus of the
Action: To raise the concha. helix, covered by the parotid salivary gland. In
The m. cervicoauricularis profundus minor, its course it passes over the root of the zygomatic
or short outward rotator (Figs. 3-3, 3-5), is a process of the temporal bone, extends along the
division of the deep layer of the postauricu anterior side of the cartilaginous auditory canal,
lar musculature. At its origin, it is rather variable and ends opposite the short rotator. This muscle
in that it can be divided into two to five clearly may undergo great reduction, and in extreme
defined muscle bundles. Of these, the posterior cases may be represented only by tendinous re
one usually comes from the external occipital mains. Often it is connected directly with the m.
protuberance, whereas the other portions are helicis, whose innervation it shares.
more or less shortened and arise aponeurotically
on the m. temporalis. Covered by the long out D eep M u sc les o f th e F ace
ward rotator, the muscle runs toward the concha
and beneath it to the extended lateral conchal The deep facial muscles are completely sep
border; in extreme cases it can descend to the arated from the superficial facial musculature
conchal canal itself. and are innervated by deep branches of the n.
Action: To turn the conchal fissure outward facialis (Huber 1923).
and backward. The m. stapedius (see Fig. 17-13) was origi
The mm. obliqui et transversi auriculae (Fig. nally associated with the primitive mandibular
3-2) are unevenly distributed muscle strands on joint. During evolution the stapes and its associ
the convex surface of the concha which, for the ated muscle have been incorporated into the
most part, proceed in a longitudinal direction. middle ear. The muscle is described along with
This layer probably belongs to the deep portion the m. tensor tympani as a muscle of the middle
of the cervicoauricular musculature. A few of its ear.
divisions appear to be distinct: thus there are The m. digastricus (biventer mandibulae)
short longitudinal fibers in the transverse groove (Fig. 3-6) runs from the processus jugularis of
between the conchal fossa and the dorsum of the occiput to the ventral border of the mandi
the remaining distal part of the pinna. There ble. Although it appears as a single-bellied mus
is a pars sulci transversi, of which a partic cle in the dog, a tendinous intersection and an
ularly long segment extends over the end of the innervation by both the n. trigeminus and the n.
long levator. Directly lateral to this is the pars facialis are evidence of its dual nature. Much has
marginalis located distal to the end of the long been written concerning this muscle in mam
outward rotator in the region of the antitragus. mals (Rouviere 1906, Bijvoet 1908, and Chaine
It extends distally. There is also the m. helicis, 1914). Only the caudal belly of the digastricus
or m. helicis retroauricularis of Huber, which belongs to the deep facial muscle group.
proceeds quite independently from the deep The digastricus lies medial to the parotid and
surface of the lateral crus of the helix. It runs mandibular glands. After crossing the ventro-
distally between the medial and lateral crura to posterior edge of the insertion of the masseter, it
the insertion of the m. scutuloauricularis super has a fleshy ending on the ventromedial border
ficialis dorsalis. of the mandible over a distance of about 2.5 cm.,
Action: Erection of the free portion of the to the level of the canine tooth. Small muscle
pinna. bundles extend far forward toward the chin.
Innervation: The innervation by a branch of Action: To open the jaws.
the ramus auricularis posterior, n. facialis, Innervation: N. facialis to posterior belly and
which can be traced from behind the con n. trigeminus to anterior belly.
M u sc les of the H ea d 145
Cervicoauri cularis p ro f m a jo r
S c u t u l o a u r i c u l a r i s prof. m i n or Ii C e r v i c o a u r i c u l a r i s prof. m i n o r
i /
Interparietoauricularis y, C e r v i c o a u r i c u l a r i s s u p e r f
Interparietoscutularis „ y C e r v i c o s cu t u la ri s
Interscutularis-
In te r p ari etoauricu laris
— Interparietoscutularis
Deep s u r f a c e o f
" " - O ccipitalis
scutiform ca rtilag e - ;
/ I I
S c u tu l o a u r i c u l a r i s p r o f m a j o r ' 1 I
S cu t u lo au r i c ul ar i s s u p e r f . m e d i u s * I
i
Lateral crus of h e l i x 1
M a n d i b u l o a u r i cu l a r i s
F ig . 3 - 5 . M uscles o f the external ear, dorsal aspect.
- - J u ^ u l o h i jo i d e u s
Stylohyoideus
F ig. 3-6. Superficial hyoid muscles and the dijjastricus, lateral aspect.
146 Chapter 3. M yology
The m. stylohyoideus (Figs. 3-6, 3-11) is a the ventral oblique muscle passes beneath the
narrow muscle bundle which proceeds from the eyeball, it gradually widens, and crosses below
tympanohyoid and proximal end of the stylo the tendon of insertion of the ventral rectus. The
hyoid obliquely across the lateral surface of the ventral oblique divides as it reaches the ventral
m. digastricus to the lateral end of the basihyoid. border of the lateral rectus. Part of its tendon
It inserts immediately posterior to the m. mylo- crosses that of the lateral rectus superficially; the
hyoideus, between the m. hyoglossus and m. deep part goes underneath the lateral rectus, and
sternohyoideus. This weak muscle is very much ends in the sclera. The superficial part ends in the
reduced, and may only be observed as a narrow sclera lateral to the insertion of the dorsal rectus.
tendinous strand. Often its origin is tendinous as Action: To rotate the eyeball around an axis
far as the dorsal border of the m. digastricus. through its poles so that the ventral part is
During its course it is covered by the mandibular moved medially and dorsally.
gland. Evans (1959) has described anomalies of Innervation: N. oculomotorius.
the stylohyoideus in mongrels and beagles which The m. obliquus dorsalis, or trochlearis (Figs.
reflect the phylogenetic history of the muscle. 3 -7 , 3-8), arises from the medial border of the
Action: To raise the basihyoid bone. optic canal. It ascends on the dorsomedial face
Innervation: N. facialis. of the periorbita to a cartilaginous pulley0 lo
The m. jugulohyoideus (jugulostyloideus of cated on the medial wall of the orbit near the
some authors) (Fig. 3-6) is a small rectangular medial canthus of the eye. The small, long,
muscle which extends from the jugular process spherical tendon passes through a groove on the
of the occiput to the cartilaginous tympanohyoid medial surface of the ventroanterior end of the
and proximal end of the stylohyoid. The muscle trochlear cartilage, where it is held in place by a
is partly covered by the end of the m. sterno- collagenous ligament. As it passes through this
mastoideus. The m. jugulohyoideus arises on the pulley, or trochlea, it bends at an angle of about
laterally projecting caudal border, whereas 45 degrees. It passes dorsolaterally and under
the m. digastricus attaches to the knobby end the tendon of the dorsal rectus, at the lateral
of the jugular process. edge of which it inserts in the sclera. It is the
Action: To move the stylohyoid bone caudally. longest and slenderest muscle of the eyeball.
Innervation: N. facialis. A ction: To rotate the eyeball around an axis
through its poles so that the dorsal part is
pulled medially and ventrally.
E x t r in s ic M u sc les of th e E yeball Innervation: N. trochlearis.
The mm. recti, or straight muscles of the eye
There are seven extrinsic muscles of the eye ball, include the mm. rectus lateralis, rectus
ball: two oblique muscles, four recti muscles, medialis, rectus dorsalis, and rectus ventralis
and the retractor bulbi. Closely associated with (Figs. 3-7, 3-8). They all arise from a poorly de
these, but inserting in the upper eyelid, is the m. fined fibrous ring which is attached around the
levator palpebrae. All of the extrinsic ocular optic canal and is continuous with the dural
muscles insert in the fibrous coat of the eyeball sheath of the optic nerve. The dorsal and medial
near its equator. The level of insertion of the recti arise farther peripherally from the optic
recti muscles is nearer the corneoscleral junction canal than do the others. As the four muscles
than is that of the four parts of the retractor. In
general, the oblique muscles insert in an inter
mediate zone between the insertions of the recti
8 The pulley, or trochlea (Figs. 3-7, A, and 3-8, A) is a disc
and retractor groups. All arise from the margin of hyaline cartilage located dorsoposteriorly to the medial
of the optic canal and orbital fissure, except the canthus of the lids on the medial wall of the orbit, less than 1
ventral oblique, which comes from the anterior cm. from the orbital margin. It is spherical to oval in outline,
part of the pterygopalatine fossa. Gilbert (1947) with its long axis parallel to that of the head. It is about 1 cm.
long by 1.5 mm. thick. It is closely related to the periorbita, in
has investigated the origin and development of
which its ligaments run. Three of these may be recognized. A
the extrinsic ocular muscles in the domestic cat. long distinct ligament runs from the anterior end of the
The m. obliquus ventralis (Fig. 3-7) arises trochlea, where the trochlear tendon bends around the carti
from the anterolateral margin of a variably sized lage, to the periosteum at the medial canthus. A short but wide
opening in the palatine bone adjacent to the thickening of the periorbita anchors the trochlea to the dorsal
wall near its margin. The third ligament is a thickening in the
suture between the palatine, lacrimal, and max periorbita which runs from the posterior pole of the trochlea
illary bones. Frequently a groove harbors the to the periosteum on the ventral surface of the supraorbital
origin of the muscle and extends posteriorly. As process.
M. r e t r a c t o r b u l b i ------
tv*.
F ig . 3 -7 . Muscles of the eye.
A. Caudolateral aspect. (The eye is displaced slightly Iaterad.)
B. The m. retractor bulbi, lateral aspect.
U
Orbital f i s s u r e
M. obi iq_uus do r s a
M. r e t r a c t o r b u l b i
M. r e c t u s m e d i a l / /
' M. r ec t u s d o r s a l i s
M. l e v a t o r p o l p e b r o
Trochlea 1. r e c t u s l a t e r a l i s
Tendon o f
M.obl i q u u s dors.
quus v e n t r a l i s
M- l e v a t o r p a l p e b r a e \ M. r e c t u s d o r s a l i s
/ O p h t h a l m i c a. v v.
M. o b i i q u u s d o r s a l i s - _ O c u l o m o t o r n.
M. r e c t u s m e d i a l i s — / T r o c h l e a r n.
O p t i c n.~ ^ A b d u c e n s n.
M. r e c t u s v e n t r o l i s ~ -O rb ita l fissure
M. r e c t u s l a t e r a l i s ' N " O r b i t a l v.
M. r e t r a c t o r b u l b i ' A n a s t o m o t i c a.
F ig . 3 -8 . Muscles of the eye.
A. Dorsolateral aspect.
B. Scheme of origin of extraocular muscles. (The m. obliquus ventralis is not shown since it originates anteriorly in
the region of the palatine-lacrimal suture.)
147
148 Chapter 3. M yology
course anteriorly from this small area of origin, the ocular muscles in reaching the upper eyelid.
they diverge and insert laterally, medially, dor- The levator inserts in the upper eyelid by means
sally, and ventrally on an imaginary line circling of a wide, flat tendon which passes between the
the eyeball, about 5 mm. from the margin of the fascicles of the m. orbicularis oculi.
cornea. The muscles are fusiform, with widened Action: To lift the upper eyelid.
peripheral ends which give rise to delicate apo Innervation: N. oculomotorius.
neuroses. The muscles diverge from each other There are also smooth muscles associated
so that wedge-like spaces are formed between with the eyeball, orbit, and lids. Several of these,
them. In the depths of these spaces the four including the ventral and dorsal palpebral mus
segments of the m. retractor bulbi lie under the cles, have been referred to in the past as muscles
fascia and fat. The recti are longer and larger of Muller. Acheson (1938) described and illus
than the parts of the retractor with which they trated the inferior and medial smooth muscles of
alternate. They therefore insert a greater dis the kitten’s eye and showed their relationship to
tance from the caudal pole than do the parts of the eyelids and nictitating membrane. In the dog
the retractor. The medial rectus is slightly larger a delicate fan of muscle fibers arises from the
than the others. trochlear cartilage and inserts in the upper lid.
Action: The medial and lateral recti rotate the These fibers are nearly continuous at their in
eyeball about a vertical axis through the sertion with the edge of the m. levator palpe
equator; the dorsal and ventral recti rotate brae superioris. Walls (1942) illustrates a muscle
the eyeball about a horizontal axis through of Muller in man as an unstriped slip of the leva
the equator. tor which inserts in the upper lid. Chauveau
Innervation: N. oculomotorius to the ventral, (1891) described a similar arrangement in the
medial, and dorsal recti; n. abducens to the horse.
lateral rectus.
The m. retractor bulbi (Figs. 3-7, 3-8) arises M u sc les of th e H y o id A ppa ra tu s
deep to the mm. recti at the apex of the orbit,
where they attach to the ventral end of the The m. sternohyoideus (Figs. 3-9, 3-46) is a
pterygoid crest and the adjacent orbital fissure. straplike muscle that arises from the deep sur
This places the initial part of the muscle lateral face of the manubrium sterni and the anterior
to the optic nerve. The four fasciculi of the m. edge of the first costal cartilage. It lies in con
retractor bulbi diverge as they run to the equa tact with its fellow, and together they extend
tor of the eye. The muscle fasciculi can be di up the neck, covering the ventral surface of the
vided into dorsal and ventral pairs. The optic trachea, to be inserted on the basihyoid bone.
nerve, as it emerges from the optic canal, passes At its origin, and throughout its caudal third,
between the dorsal and ventral portions. The in the deep surface of the m. sternohyoideus is
sertion of the several parts of the retractor on the fused to the m. sternothyroideus. The caudal
globe of the eye is about 5 mm. caudal and deep third of the m. sternohyoideus is covered by the
to the recti. This muscle is sometimes spoken of m. sternocephalicus, and, in specimens in which
as the choanid. there is a decussation of fibers between the ster-
Action: To retract the eyeball. In addition, be nocephalic muscles, the caudal two-thirds of the
cause of its essentially alternate attach muscle will be covered by these crossed fascic
ments with the recti, it aids in bringing uli. The anterior portion of the muscle which is
about oblique eye movements. not covered by the m. sternocephalicus is the
Innervation: N. abducens. most ventral muscle of that portion of the neck,
except for the platysma. A transverse fibrous in
tersection separates the caudal third from the
M u sc les of th e E y e l id s cranial two-thirds of the muscle.
Variations: Occasionally muscle slips will
The mm. orbicularis oculi, retractor anguli arise from the transverse fibrous intersection of
oculi, and superciliaris have been described on the m. sternohyoideus and pass up the neck to
page 140 with the superficial muscles of the face. be inserted half in the stylohyoideus muscle, just
The m. levator palpebrae superioris (Figs. 3 - lateral to the basihyoid bone, and half in the di
7, 3-8) is the main retractor of the upper eyelid. gastricus muscle at the angle of the jaw. The m.
It arises dorsal to the optic canal between the digastricus may be considerably smaller than
dorsal rectus and the dorsal oblique muscles. It normal and separated by a short intermediate
courses deep to the periorbita and superficial to tendon, as is the homologous muscle of man and
M u sc les of the H ea d 149
P t e r y g o id med, c u t S ty lo pharyngeus
Kena tophar yncj eus
HyopharynyeuS
! ,Chondrophari/njeus
- C l e id o c e r > vicaH s
--S f e n n o o c d p i to lis
-S iernom astoideus
- Thy r o p h o r y n c j e u s
S te rn o th y ro id e u s
~S t e m o h y o i d e u s
M y lo h y o id e u d 1
S t y lo g lo s s u s
H y o g lo ssu s
--S t e m o c e p h a /ic u s
CeniohLjoideus'
T h y ro h y o id eu S C r ic o t h y r o i d e u s
Fir.. 3-9. The hyoid muscles and muscles of the neck, lateral aspect. (Stylohyoideus and digastrieus removed.)
150 Chapter 3. M yology
horse. Leahy (1949) and Evans (1959) have de form muscle which extends from the chin, paral
scribed unilateral and bilateral anomalous slips lel to the mid-ventral line, to the basihyoid bone.
in the dog. It arises by a short tendon from the mandibular
Action: To pull the basihyoid bone and tongue symphysis and, muscularly, from the inner sur
posteriorly. face of the mandible adjacent to the symphysis.
Innervation: Nn. cervicales et n. accessorius. It passes directly caudad, at first bordered on the
The m. thyrohyoideus (Figs. 3-9, 3-15) origi lateral side by the m. genioglossus and in its fur
nates on the lamina of the thyroid cartilage. At ther course by the m. mylohyoideus, which also
the thyroid attachment it is bordered dorsally by covers much of its ventral surface. Throughout
the insertion of the m. thyropharyngeus and cau its length the muscle is in close contact with its
dally by the mm. cricothyroideus and sternothy- fellow of the opposite side. It is inserted on the
roideus. Its fibers pass obliquely forward and anterior border of the basihyoid bone.
downward, over the surface of the thyroid lam Action: To draw the hyoid apparatus ante
ina, to be inserted along most of the caudal bor riorly.
der of the thyrohyoid bone. Innervation: N. hypoglossus.
Action: To draw the hyoid apparatus poste The mm. jugulohyoideus and stylohyoideus
riorly and dorsully. are described under the deep muscles of the
Innervation: Nn. cervicales et n. accessorius. face.
The m. mylohyoideus (Figs. 3-9, 3-10) lies M u sc les of M a s t ic a t io n
most ventrally in the intermandibular space. To
gether with the muscle of the opposite side, it The m. masseter (Fig. 3-13) lies on the lateral
forms a sling for the tongue. It has a long ori surface of the ramus of the mandible ventral to
gin from the medial side of the mandible. In the zygomatic arch. It projects somewhat be
most specimens the most anteriorly arising fibers yond the ventral and posterior borders of the
are opposite the first lower premolar tooth and mandible. The muscle is covered by a strong,
the most posteriorly arising fibers are slightly glistening aponeurosis, and tendinous intermus-
posterior to the last lower molar tooth. From its cular strands are interspersed throughout its
origin the muscle fibers extend medially, form depth. One can divide the muscle into three
ing a thin plate which is largely inserted on a layers (superficial, middle, and deep), using the
median fibrous raphe with its fellow of the oppo change of fiber direction as a guide to the sep
site side. The most anterior fibers curve and in aration between the layers.
sert on the mid line farther forward than their The superficial layer, the strongest part, arises
point of origin. A few of the most posterior fi from the ventral border of the anterior half of
bers curve and pass posteriorly, to be inserted the zygomatic arch. Its fibers pass posteroven-
on the basihyoid bone. The deep surface of the trally and insert, partly, on the ventrolateral
muscle is related to the m. geniohyoideus, the surface of the mandible. Some fibers project
tongue, and the oral mucosa. around the ventral and posterior borders of the
Action: To raise the floor of the mouth and mandible and insert on its ventromedial surface,
draw the hyoid apparatus anteriorly. as well as on a tendinous raphe which passes be
Innervation: Ramus mandibularis, n. trigemi tween the masseter and the m. pterygoideus me
nus. dius. The tendinous raphe continues caudally
The m. keratohyoideus (Figs. 3-16, 3-20) is a from the angle of the jaw and attaches on the
small triangular plate of muscle, one side of bone adjacent to the tympanic bulla. In speci
which attaches to the anterior border of the thy mens with well developed masseters, this layer,
rohyoid bone. The fibers run anteroventrally at its ventral border, projects somewhat over the
from the thyrohyoid bone to the keratohyoid m. digastricus.
bone, to be attached along the dorsal border of The middle layer, the weakest part, arises
the bone. In some specimens a few fibers attach from the zygomatic arch, medial to the origin of
to the ventral end of the epihyoid bone. The the superficial layer and in part posterior to it.
deep surface of the muscle is related to the root Most of its fibers pass ventrally to be inserted
of the tongue and the oral mucosa, and the outer on the ventral margin of the masseteric fossa and
surface is related to the m. keratopharyngeus. the narrow area just ventral to the fossa. In some
A ction: To decrease the angle formed by the specimens a small bundle of fibers, which belong
thyrohyoid and keratohyoid bones. to this layer, run in a more anterior direction to
Innervation: N. glossopharyngeus. be inserted on the anteroventral margin of the
The m. geniohyoideus (Fig. 3-10) is a fusi fossa.
M u sc les of the H ead 131
M yiohLjoideuti
- Mandibular foramen
Lymph node
Ire tro p h a ry n g eal)
M a s s e te rs u p e rf portion'
SternOthijroideuS
M a s s e t e r —m i d d l e p o r t i o n '
U tcjast r ic u s / S terrohyoideus
M y I o h yo< d e u s / \ ' T h y rO p h a r L / n ^ e u s
S tylohyoideus Hyopharyngeus
Fic. 3 11. Musdes of mastication, lateral aspect.
152 Chapter 3. M yology
The deep layer is impossible to isolate at its auricular muscles. From its large origin the mus
origin because many of its fibers intermingle cle fibers curve forward and downward beneath
with those of the temporalis. Some fibers, how the zygomatic arch to invest and insert on the
ever, arise from the medial surface of the zygo coronoid process of the mandible, as far down
matic arch. The majority of its fibers are di as the ventral margin of the masseteric fossa. On
rected posteroventrally and are inserted in the the lateral side of the coronoid process the fibers
posterior part of the masseteric fossa and on the are intermingled with fibers of the deep layer of
ridge adjacent to it. A few fibers pass down the m. masseter; on the medial side the fibers lie
along the anterior margin of the temporal mus in contact with the mm. pterygoidei. A bundle
cle to be inserted on the anterior ridge of the of muscle fibers arises from the dorsal nuchal line,
masseteric fossa. near the base of the zygomatic process of the
Action: To raise the mandible in closing the temporal bone, and sweeps forward dorsal and
mouth. parallel to the zygomatic arch. It blends grad
Innervation: N. massetericus of the ramus ually into the main mass of the muscle.
mandibularis, n. trigeminus. Action: To raise the mandible in closing the
The m. temporalis (Figs. 3-11, 3-12, 3-13) is mouth.
the largest and strongest muscle of the head. It Innervation: N. temporalis of the ramus man
occupies the temporal fossa, from which it ex dibularis, n. trigeminus.
tends downward around the coronoid process of The m. pterygoideus lateralis (Fig. 3-13) is a
the mandible. During the course of its down much smaller and shorter muscle than the m.
ward extension, it is related anteriorly to the or pterygoideus medialis. It arises from the sphe
bit and orbital fat, medially to the mm. noid bone in a small fossa, which lies ventral to
pterygoidei and laterally to the m. masseter. the alar canal, round foramen, and orbital fis
Dorsolaterally it is covered by the postauricular sure. The ventral boundary of its origin is a bony
muscles, the scutulum, and the ear. It arises ridge also on the sphenoid bone. This short mus
largely from the parietal bone and to a lesser ex cle passes ventrolaterally and slightly posteriorly,
tent from the temporal, frontal, and occipital to be inserted on the medial surface of the con
bones. The margins of the muscle at its origin dyle of the mandible just ventral to its articular
are the orbital ligament and external frontal surface.
crest anteriorly, the zygomatic arch laterally, Action: To raise the mandible.
the dorsal nuchal line posteriorly, and the ex Innervation: Nn. pterygoidei of the ramus
ternal sagittal crest medially. Closely applied to mandibularis, n. trigeminus.
the muscle, within these margins, is a strong, The m. pterygoideus medialis (Fig. 3-13)
glistening fascia. In dolichocephalic dogs the arises from the lateral surface of the pterygoid,
temporal muscle meets its fellow of the oppo palatine, and sphenoid bones. It passes postero-
site side and forms a mid-dorsal sulcus. In dogs laterally to be inserted on the inner surface of
with brachycephalic heads the temporal mus the mandible, adjacent to the angle and ventral
cles usually do not meet on the mid line, and to the insertion of the mm. temporalis and ptery
the area is devoid of muscle, except for the post- goideus lateralis. Many fibers insert on a fibrous
raphe that passes between the insertion of this
muscle and the superficial layer of the masseter
muscle. When viewed from the pharyngeal side,
the medial pterygoid completely covers the lat
eral one.
The mandibular alveolar nerve passes across
the lateral face of the m. pterygoideus medialis
and the medial surface of the m. pterygoideus
lateralis, thus separating the two muscles. The
m. pterygoideus medialis extends to the poste
rior margin of the mandible and is inserted on
the posterior margin and slightly on the pos
teromedial surface.
Action: To raise the mandible.
Innervation: Nn. pterygoidei of the ramus
mandibularis, n. trigeminus.
F ig . 3-12. The m. temporalis, lateral aspect. (Zygomatic The m. digastricus is described with the deep
arch removed.) muscles of the face.
M u sc les of th e H ead 153
P t e r y g o i d e u s m e d ’ Olis
,P te rg g o id e u s la t e r a lis
i
,A re o o f in s e rt io n
o f P t e ry g o id , tat.
-A rea o f insertion of i
P t e r y g o i d e u s med. 'M a s s e t e r
'Body of m a n d ib le , cut ii
ii
V
P terygoideus med
-Temporahs
- - P t e r y g o i d , l at
M u sc les of the T ongue sal to the m. mylohyoideus. The most anterior fi
bers run upward and forward, to be inserted on
The m. styloglossus (Figs. 3-15, 3-16) ex the mid-ventral surface of the tip of the tongue.
tends from the stylohyoid bone to the tongue. It These fibers form the substance of the frenulum.
is composed of three muscle heads which insert The remaining fibers sweep upward and back
in the tongue at different levels along its long ward in a fanlike arrangement, to be inserted
axis. along the mid-ventral surface of the tongue in
The short head arises from the distal half of close contact with the fibers of the correspond
the posterior surface of the stylohyoid bone. It ing muscles of the opposite side. A distinct bun
curves downward and forward across the lateral dle of fibers runs directly posteriorly, to be
surface of the epihyoid bone. Immediately after inserted on the basihyoid and keratohyoid bones.
crossing the epihyoid bone the fibers diverge Action: To depress the tongue. The posterior
and insert on the base of the tongue among the fibers draw the tongue forward; the anterior
inserting fibers of the hyoglossal muscle. fibers curl the tip of the tongue downward.
The anterior head arises from the anterior sur Innervation: N. hypoglossus.
face of the proximal half of the stylohyoid bone. The m. propria linguae consists of many mus
These fibers curve downward and forward, pass cular bundles which are located among the fas
over part of the inserting fibers of the short head, cicles of insertion of the extrinsic muscles of the
intermingle with fibers of the m. hyoglossus, and tongue. They are arranged bilaterally in four
insert in the tongue, along its ventrolateral sur poorly delineated groups: (1) longitudinalis dor
face. salis, (2) longitudinalis ventralis, (3) transversus
The long head arises just above and lateral to linguae, and (4) verticalis linguae. The dorsal
the origin of the fibers of the short head. These longitudinal fibers lie directly under the dorsal
fibers immediately cross the stylohyoid bone, mucosa of the organ and are well developed.
then curve downward and forward along the The transverse and oblique fibers form a rather
ventral border of the anterior head. They con wide zone under the dorsal longitudinal bundles.
tinue forward along the ventral mid line of the A few long muscle strands lie ventral to the
tongue and across the lateral side of the genio- above-mentioned zone and compose the ventral
glossus muscle, to their insertion on the ventral longitudinal muscle. Thus the muscle bundles
surface of the anterior half of the tongue, near run in diverse directions.
the median plane. Action: To protrude the tongue and bring
Action: To draw the tongue backward when about complicated intrinsic, local move
all three heads act together. Each muscle ments; to prevent the tongue from being
head depresses the tongue sector to which bitten. The tongue functions in mastication
it is attached. and deglutition, as well as serving as the pri
Innervation: N. hypoglossus. mary organ of taste. Bennett and Hutchin
The m. hyoglossus (Figs. 3-14, 3-15, 3-16) is son (1946) have discussed the action of the
located in the root of the tongue. It arises from tongue in the dog.
the ventrolateral surface of the basihyoid and Innervation: N. hypoglossus.
the adjoining end of the thyrohyoid bone. It runs
forward dorsal to the m. mylohyoideus and lat
eral to the mm. geniohyoideus and genioglossus.
At the base of the tongue it crosses the medial M u sc les of th e P harynx
side of the m. styloglossus to be inserted in the
root and posterior two-thirds of the tongue. The m. hyopharyngeus (Figs. 3-9, 3-15, 3 -
Action: To retract and depress the tongue. 17) is often divided into two parts: the m. chon-
Innervation: N. hypoglossus. dropharyngeus and the m. keratopharyngeus.
The m. genioglossus (Figs. 3-14, 3-15) is a The m. chondropharyngeus, the larger part,
thin, triangular muscle which lies in the inter- arises from the lateral surface of the thyrohyoid
mandibular space, in and beneath the tongue. bone under cover of the hyoglossal muscle. The
The apex of this triangular muscle corresponds m. keratopharyngeus arises from the keratohyoid
to its origin on the medial surface of the man bone. The muscle fibers of both parts form a
dible, just posterior to the origin of the genio muscle plate, the fibers of which pass upward
hyoideus. The fibers run backward and upward over the larynx and pharynx to be inserted on
in a sagittal plane. In their course the muscle the medial dorsal raphe of the pharynx, opposite
fibers lie lateral to the m. geniohyoideus and dor the insertions of the muscles of the opposite side.
M u sc les of the H ead 155
E p i h y o i d bone.
C e n io q lo s s u s
S t y l o h y o i d --------- --
T h y ro h y o id - -
S tyloglossus, s h o r t head
Styl ogl ossus, ant. he a d , ! / J v g u l o h y o i de us
/K e ra to p haryngeus 1 Hyo-
, - C h o n d r o p h a n y n g e u s j P^ a r y r 9euS
,~ T h y r o p h a r y n g e u S
S t e r n o t h y n o i deus
— C r i c o t h y r o i deus
-----S t e r n o h y o i d e u s
Gen loqlossus Hyogl ossus
'Th y r o h y o i d e u s
• T h y r o p h aryngeus
- Cricopharyngeus
-Esophagus
- S te r n o t h y r o i d e u s
- Trochea
Near their insertions the caudal fibers are over stylopharyngeus blend with the m. palatopha
laid by inserting fibers of the m. thyropharyn- ryngeus on the dorsal wall of the pharynx. A few
geus. The m. hyopharyngeus is the most anterior fibers run forward from their palatine origin and
pharyngeal constrictor. are dispersed in the soft palate, nearly as far for
Action: To constrict the anterior part of the ward as the hamulus of the pterygoid bone.
pharynx. Action: To constrict the pharynx and draw it
Innervation: Nn. glossopharyngeus et vagus. forward and upward.
The m. thyropharyngeus (Figs. 3-15, 3-17) Innervation: Nn. glossopharyngeus et vagus.
lies on the larynx and pharynx just caudal to the The m. pterygopharyngeus (Figs. 3-18,3-19)
hyopharyngeus muscle. It arises from the arises from the hamulus of the pterygoid bone,
oblique line on the lamina of the thyroid carti passes backward lateral to the m. levator veli
lage, and goes upward and forward over the dor palatini, and continues upward over the pharynx
sal border of the thyroid lamina. The fibers to be inserted on the mid-dorsal raphe. Its fibers
spread out over the dorsal surface of the pharynx are intermixed with fibers of the m. palatophar
and insert on the median dorsal raphe of the yngeus and the m. stylopharyngeus as they radi
pharynx, just caudal to the m. hyopharyngeus. ate toward their insertions.
Some of the most anterior fibers of insertion Action: To constrict the pharynx and draw it
overlie fibers of the m. hyopharyngeus. forward.
Action: To constrict the middle part of the Innervation: Nn. glossopharyngeus et vagus.
pharynx.
Innervation: Nn. glossopharyngeus et vagus.
The m. cricopharyngeus (Figs. 3-16, 3-17) M u sc les of the So ft P alate
lies on the larynx and pharynx immediately cau
dal to the m. thyropharyngeus. It arises from the The m. tensor veli palatini (Fig. 3-19) is a
lateral surface of the cricoid cartilage and passes very small muscle that arises from the muscular
upward to be inserted on the median dorsal process at the anterior margin of the tympanic
raphe. As the muscle fibers pass over the dorsal bulla. From its origin it passes downward over
wall of the pharynx they blend, at their caudal the wall of the pharynx to the hamulus of the
margin, with muscle fibers of the esophagus. pterygoid bone. At the hamulus the muscular
Action: To constrict the caudal part of the fibers become tendinous and pass over a troch
pharynx. lear ridge on the hamulus. In their distal course
Innervation: Nn. glossopharyngeus et vagus. these tendinous fibers radiate forward and are
The m. stylopharyngeus (Figs. 3-18, 3-20) is dispersed in the palate.
a weak muscle that extends from the stylohyoid Action: To stretch the palate between the
bone to the anterodorsal wall of the pharynx. In pterygoid bones.
most specimens the fibers arise from the caudal Innervation: Ramus mandibularis, n. trigemi
border of the proximal end of the stylohyoid nus.
bone; on some specimens, however, a few fibers The m. levator veli palatini (Figs. 3-19,3-20)
arise on the epihyoid bone. From their origin the is slightly larger than the m. tensor veli palatini.
fibers run backward and inward beneath the It arises from the muscular process adjacent to
constrictor muscles on the dorsolateral wall of the tympanic bulla and passes downward and
the pharynx, where they are loosely arranged backward on the wall of the pharynx. In its distal
and intermingle with fibers of the m. palato- course it passes between the m. palatopharyn
pharyngeus. geus and the m. pterygopharyngeus and radiates
Action: To dilate, elevate, and draw the phar to its insertion on the caudal half of the soft pal
ynx forward. ate lateral to the m. palatinus.
Innervation: Nn. glossopharyngeus et vagus. Action: To raise the caudal part of the soft pal
The m. palatopharyngeus (Figs. 3-18,3-19) ate.
is a poorly developed muscle, medial to the m. The m. palatinus (m. uvulae) (Fig. 3-19) is a
tensor veli palatini, whose fibers are loosely as small, straight muscle that runs longitudinally
sociated as they encircle the pharynx. Dyce through the soft palate. It arises from the pala
(1957) divides the muscle into a dorsal and ven tine process of the palatine bone and passes with
tral portion. Most of the fibers arise from the soft its fellow to the caudal free border of the soft
palate and sweep obliquely upward and back palate.
ward over the pharynx to the mid-dorsal line. Action: To shorten the palate and curl the pos
Some fibers of the mm. pterygopharyngeus and terior border downward.
M u sc les of the H ea d
Tcnyue- O ra l p h a ry n x
S c ff p a l a t e , c u t e d y e - ■Nasal p h a n y n x
J tylcglOSSUS-
P te n y y o p h a r y n y ew
T o n s il-
S ty lop h a n y n y e u S
tiy a p h a n y n g e us-
An tic ula Uon o f thynohyoid
- on d t h y r o i d c a n f i l o y e
M ed ian naphe-
-Thy ro p h a n y n y eus
C m cophanynaeus
E S op h a y us
P a !a t in u s
S ty loylossus
__ , Epihyoid
S ty lo h y o id ~ te n y y o p h a r y n q e u s
■Ptenycjopharyn -eus
-Pala tophortjnqeuA
Pa la tinus
°alat/ne process
F i t . 3-19, Muscles ol the pharynx and palate, deep dissection ventrolateral aspect.
_ -Nasal pDanynx
J o t t p al at e
Oral p h a r y n x i (
Stylopharyngeus' Keratohyoideus
Fic. 3 -20 Muscles of the pharynx and palate, deep dissection, lateral aspect.
M u sc les of the H ea d 159
Thyroid c o rti
c ot hy roi deu s
(reflected)
F ig . 3-21. Laryngeal muscles, lateral aspect. (The thyroid cartilage is cut left of mid line and reflected.)
M. v e n t r i c u l a r i s
i C or ni c u l at e process o f a r y t e no i d c a r t i l a g e
}rarytenoid ca rtila ge
Ep i g I a t t i s ri c o i d c a r t i l a g e
C u n e i f o r m p r o ces s A r t i c u l a t i o n w i t h t h y r o i d cart.
arytenoid ca rtilag
-M. c r i c o a r y t e n o i deus lat.
V e ntricu la r I i
- Tr a c he a
M. t h y r o a r y t e n o i
Vocal lig ■ ! ' C r i c o t h y r o i d l i g.
1M. vocal is
F ig . 3-22. Laryngeal muscles, lateral aspect. (The thyroid cartilage is cut left of mid line and removed; the mm. thyroarytenoi-
deus, arytenoideus transversus, and cricoarytenoideus dorsalis have also been removed.)
-Epiglottis
L a t ventricle -
- L ar yn g ea l sa c cul e
C r a n i a l c o r n u of t h y r o i d c a r t i l a g e - -M. v e n t r i c u l a r i s
Laryngeal saccule - ~ - M. t h y r o a r y t e n o i d e u s
" M. a r y t e n o i d e u s transversus
C o r n i c u l a t e ca r t i l a g e ' ~M. c r i coa r y t e n o i d eu s dorsalis
C ricoid c a r t i l a g e ' > llll§ ^ lt§ i ' - C au d a l c or n u of t h y r o i d c a r t i l a g e
Trachea /
F ig . 3-23. Laryngeal muscles, dorsal aspect. (The right corniculate cartilage has been cut and the right laryngeal saccule is re
flected.)
160
M u sc les of the H ea d 161
from the opposite side, and blends with the more the platysma, it contains the pars palpebralis,
dorsally located m. ventricularis which spans the and pars intermedia, (parts of the m. sphincter
mid line. colli profundus) and covers the buccinator and
Action: To constrict the glottis and adduct the the large facial vessels and nerves. The parotid
vocal folds. duct is loosely surrounded as it lies behind the
The m. hyoepiglotticus (Fig. 3-14), a small, labial commissure. The fascia spreads out into
spindle-shaped muscle, arises from the medial the lips. Posteriorly it turns over the external
surface of the keratohyoid bone. It passes medi surface of the m. masseter.
ally to the mid line, then turns dorsally and The parotideomasseteric fa s c ia (fascia paroti-
passes to the ventral mid line of the epiglottis to deomasseterica) is the continuation of the above
be inserted. The fibers of fellow muscles blend described portion of the superficial fascia cover
into a common tendon of insertion, which fades ing the m. masseter and going to the external
into the ventral surface of the epiglottis. surface of the parotid gland and mandible; the
Action: To draw the epiglottis downward. branches of the facial nerve and the parotid duct
are therefore surrounded by it. Dorsally this
F a s c ia e o f t h e H ead fascia goes over the zygomatic arch into the
superficial temporal fascia and spreads out on
The superficial fascia of the head lies directly the pinna of the ear. It contains the pars inter
beneath the skin; for the most part it is easily media and pars auricularis of the m. sphincter
displaceable, but in the muzzle it fuses with the colli profundus, and the platysma with which it
skin. It contains the cutaneous muscles of the extends into the superficial cervical fascia pos
head, portions of the platysma and the m. teriorly. Ventrally it goes into the submandibular
sphincter colli profundus. It covers the entire fascia.
head like a mask and continues on the neck like The superficial subm andibular fa s c ia (fascia
a cylinder. It is divided into the pars temporalis, submandibularis superficialis) is the intermandib-
pars nasofrontalis, pars buccalis, pars parotideo- ular portion of the fascia of the head; it courses
masseterica, and pars submandibularis. In many between the bodies of the mandible on either
places the special nerves and vessels for the skin side as a continuation of the superficial buccal
pass through the superficial fascia of the head. and masseteric fasciae, and covers the m. mylo
The superficial temporal fa s c ia (fascia tem hyoideus and the body of the hyoid bone, with
poralis superficialis) conceals the muscles of the its musculature. Posteriorly it runs into the
scutular group as well as the scutulum itself. region of the larynx and into the superficial
Medially it goes into the superficial temporal cervical fascia.
fascia of the other side without attaching to the The deep fascia of the head is found on all
median system of cranial ridges. Anteriorly it parts of the head. As the deep tem poral fa scia
continues as the superficial nasofrontal fascia; (fascia temporalis profunda) it is thick as it
more laterally and ventrally, however, and also covers the temporal muscle and spreads out, en
displaceable with respect to the underlying tis closed by and attached to the external frontal
sue, it goes over the orbital ligament to the lids. crest, external sagittal crest, dorsal nuchal line,
Posteriorly it extends over the long levator of the and the zygomatic process. If a part of the pari
pinna and becomes the superficial fascia which etal bone is not covered by the temporal muscle,
contains the platysma. Laterally it passes over to as frequently occurs in brachycephalic breeds,
the pinna and, anterior to this, spreads over the then this fascia fuses with the periosteum of the
zygomatic arch into the parotideomasseteric bone. Anteriorly, the deep temporal fascia be
fascia. comes the deep nasofrontal fa s c ia (fascia naso
The superficial nasofrontal fa scia (fascia naso frontalis profunda), called the galea aponeurotica
frontalis superficialis) comes from the superficial in man, and attaches to the orbital ligament.
temporal fascia and, containing the mm. fronta Ventrally the deep temporal fascia passes over
lis and levator nasolabialis and their divisions, the zygomatic arch and the masseter as the deep
covers the nasofrontal region. It spreads out into masseteric fa scia (fascia masseterica profunda).
the upper eyelid, the nose, and the upper lip; It then spreads over the m. buccinator, extends
posteriorly, between the palpebral and labial into both lips, and passes over the mandible and
commissures, it goes into the fascia of the cheek. larynx, as the buccopharyngeal fa sc ia (fascia
The superficial bu ccal fa s c ia (fascia buccalis buccopharyngea). From the m. masseter, pos
superficialis) comes from the superficial naso teriorly, the buccopharyngeal fascia passes
frontal fascia. In addition to buccal portions of around the parotid gland, forming the fascia
162 Chapter 3. M yology
parotidea, crosses the digastricus, and goes be as the inspiratory part, lies on the dorsal surface
neath the mandibular gland and thence into the of the thorax, where its aponeurosis covers the
deep cervical fascia. Everywhere, on the head, m. splenius and its fleshy part covers the mm.
the deep fascia lies beneath the large superficial longissimus dorsi and iliocostalis from ribs 2 to
vessels. 10. The muscle arises by a broad aponeurosis
from the superficial leaf of the fascia spino-
M USCLES OF THE TRUNK transversalis and, by means of this, from the
The trunk muscles are divided topographically tendinous raphe of the neck as well as from the
into the muscles of the cervical, thoracic, and spines of the first six to eight thoracic vertebrae.
lumbar vertebrae; muscles of the lateral and This aponeurosis fuses caudally with that of the
ventral thoracic wall, including the diaphragm; mm. latissimus dorsi and serratus dorsalis cauda
muscles of the abdomen; and muscles of the tail. lis. The fleshy portion of the muscle begins at
The special muscles of the trunk are partly about the dorsal border of the m. latissimus dorsi;
covered by those passing from the trunk to the it ends immediately lateral to the m. iliocostalis,
limbs. This applies especially to those of the with distinct serrations on the cranial borders
neck and the thoracic wall. In the lumbosacral and the lateral surfaces of ribs 2 to 10. The
region, the axial muscles continue into the dorsal fibers of the muscle, as well as those of its apo
coccygeal muscles, and the muscles of the pelvic neurosis, are directed caudoventrally.
limb overlap those of the trunk. Action: To lift the ribs for inspiration.
Innervation: Nn. intercostales (branches from
the branch to the m. intercostalis externus).
M u sc les o f th e C e r v ic a l , T h o r a c ic ,
The narrower m. serratus dorsalis caudalis, or
and L um bar V ertebra e
expiratory part, consists of three rather distinctly
The muscles of the vertebrae, as far caudally isolated portions. These arise by a broad apo
as the sacrum, represent the trunk muscles in neurosis from the lumbodorsal fascia from which
the narrow sense. They are grouped, aside from the m. obliquus externus abdominis and m. ob-
the cutaneous musculature, into five layers, liquus internus abdominis also arise. After ex
which lie beside and one above another. Of these tending cranioventrally, they end on the caudal
the two superficial layers and part of the third border of the eleventh, twelfth, thirteenth, and,
layer are discussed with the muscles of the occasionally, also the tenth rib.
thoracic limb. Action: To draw the last three or four ribs
The muscles of the first layer are: mm. trape caudally for expiration.
zius and cleidocephalicus; the second layer: mm. Innervation: Branches from the nn. inter
latissimus dorsi and rhomboideus; the third costales (from the trunk or the ramus me
layer: mm. serratus ventralis, serratus dorsalis, dians of the thoracic nerves).
and splenius; the fourth layer: mm. iliocostalis, The m. splenius (Fig. 3-25) is a flat, fleshy,
longissimus thoracis, longissimus cervicis, longis- triangular muscle with the caudal end as the
simus capitis, longissimus atlantis, spinalis et apex, and the cranial end as the base of the tri
semispinalis dorsi et cervicis, and semispinalis angle. It lies on the dorsolateral portion of the
capitis; and the fifth layer: mm. multifidus, inter- neck, extending from the third thoracic vertebra
spinales, intertransversarii, and the dorsal mus to the skull. Its fibers run in a cranioventrad di
cles on the atlanto-occipital and axio-atlantal rection and cover the mm. semispinalis capitis,
joints—the mm. obliquus capitis caudalis, ob- longissimus capitis, and the terminal part of the
liquus capitis cranialis, and rectus capitis dorsalis, m. spinalis et semispinalis dorsi. It arises by
consisting of three parts. fleshy fibers from the end of the first and some
The m. serratus dorsalis (Figs. 3-24, 3-25, 3 - times of the second thoracic spine, and from
31) is a wide flat muscle partially covering the about 1 cm. of the ligamentum nuchae immedi
m. longissimus and the m. iliocostalis. Lying ately in front of the first thoracic spine. A third
under the mm. rhomboideus, serratus ventralis, point of origin is from the median dorsal raphe
and latissimus dorsi, it arises by a broad apo of the neck as far cranial as the first cervical
neurosis from the tendinous raphe of the neck vertebra. This tendinous raphe runs from the
and from the thoracic spines, and inserts on the first thoracic spine, where it fuses with the liga
proximal portions of the ribs. The muscle is com mentum nuchae, anteriorly to the occiput. The
pletely divided into two portions (see the special final origin of the m. splenius is by an aponeuro
investigations of Maximenko 1929, 1930). sis from the deep leaf of the fascia spinotrans-
The m. serratus dorsalis cranialis, also known versalis, which extends caudally to the fifth or
M u sc les of the T run k 163
1 t
E xternal in t e r c o s t a l m .III1 1V R ib
F ig . 3-24. Muscles of neck and thorax, lateral aspect.
164 Chapter 3. M y o lo g y
I liocosfalis
I
J p ie n iu s L ongissim us |
Spm alis e t se m isp in a h s
S e r r a t u s d o rs a lis
Longissimus cervicis
Fic. 3-25. Topography of the mm. splenitis and serratus dorsalis cranialis.
sixth thoracic spine. At the cranial border of the volve the development of long fascicles instead
atlas the in. splenius is enclosed in a coarse apo of short metaineric ones, the shifting of muscle
neurosis which inserts on the dorsal nuchal line insertions onto the neural spines for better lever
of the occipital bone and the mastoid part of the age, and the fusion of the caudal portions of the
temporal bone. The in. splenius may occasion m. iliocostalis with the m. longissiinus to form
ally send a strong serration to the transverse the h i . sacrospinalis or erector spinac. Slijper
process of the axis. At the lateral border of the (1946) described the functional anatomy of the
atlas the dorsal surface of the m. longissiinus epaxial spinal musculature in a wide variety of
capitis attaches firmly to the in. splenius and. by mammals.
means of a strong tendon so formed, inserts along
with the m. splenius on the mastoid part of the Iliocostalis System
temporal bone.
Action: To extend and raise the head and neck. The ni. iliocostalis (Fig. 3-26) consists of a
In unilateral action to draw the head and series of fascicles lateral to the other epaxial
neck laterally. It also functions in fixation of muscles. The caudal members of the series arise
the first thoracic vertebra. on the iliuin and constitute a lumbar portion
Innervation: Nn. cervicales. whereas the cranial fascicles extend to the
seventh cervical vertebra and constitute the
E p a x ia l S p in a l M u sc u la tu r e thoracic portion.
The m. iliocostalis lunihorum is a strong mus
The dorsal trunk musculature, associated with cle mass which arises from the pelvic surface of
the vertebral column and ribs, may be divided the wing of the ilium, the iliac crest, and from an
into three longitudinal muscle masses, each com intermuscular septum located l>etween the m.
prising many overlapping fascicles. The muscles iliocostalis and m. longissiinus. This septum is
act as extensors of the vertebral column and also attached to the ilium and the deep surface of the
produce lateral movements of the trunk when lumlx)dorsal fascia. As the fibers of the muscle
acting only on one side. run cranioventrad, strong lateral fascicles from
On the lateral aspect is the m. iliocostalis sys the ends of all the lumbar transverse processes
tem; intermediately, the h i . longissiinus system; join them. The cranial end of the lumbar portion
and deep medially, the m. trunsversospinalis sys runs toward the ribs and is distinctly separated
tem. Various fusions of these three primary seg from the in. loiigissiinus. With increasingly
mental muscle masses result in patterns which weaker fleshy serrations, the m. iliocostalis luin-
differ according to the species. The specializa boruin attaches to the thirteenth, twelfth,
tions seen in the epaxial musculature, associated eleventh, and tenth ribs, and occasionally, by a
with the leaping-gallop type of locomotion, in long delicate tendon, to the ninth rib also.
F i b e r s f r o m l on g i ss i m us t h o r a c i s et
L on gi ssi mu s t h o r o c i s et l u m b o r u m
Spi nal is thoracis lumborum]
K Lumbodorsol fo scio c o v e r in g
Sp i na l is cern'ci s IV
t r an s v er s os pi na l is m u s c u l a t u r e
M
Semi s p i n a l is c a p i t i s (Compl exus)
u sc les
of
the
T
run k
I l i oco st ah s t ho r o c is Longi Jsi mus t h o r o c i s et l u m b o r u m
l l iocast al is lumborum
The m. iliocostalis thoracis is a long, narrow and ventral surface of the ilium, and medially
muscle mass extending from the ribs, except the from the spinous processes and supraspinous lig
first and last, to the transverse process of the ament. Its fibers run craniolaterally. The apo
seventh cervical vertebra. Its origin lies medially neurosis is divided into many strong tendinous
under the cranial segments of the m. iliocostalis strands between which narrower intermediate
lumborum. It lies lateral to the m. longissimus portions extend. Cranially, it is dissipated at the
and reaches its greatest size between the fifth fifth rib. From the eleventh to the seventh rib, it
and third ribs. It is composed of individual por serves as an origin superficially for the m. spi
tions which originate on the cranial borders of nalis et semispinalis thoracis et cervicis. In the
the vertebral ends of the ribs; they extend crani- lumbar region the m. longissimus sends off seven
olaterally and, after passing over one rib, form a medially directed fascicles from the ilium and
common muscle belly. From this belly, terminal the intermuscular septum. These fascicles cover
serrations arise which, by means of long tendons the roots of the lumbar transverse processes, and
(stronger cranially), end on the costal angles of end on the accessory processes of the sixth to
the ribs and most cranially on the transverse first lumbar vertebra. The weak, most caudal
process of the seventh cervical vertebra. portion runs to a fleshy insertion on the arch of
Action: Fixation of the vertebral column or the seventh lumbar vertebrae and to the inter-
lateral movement when only one side con vertebral disc of the lumbosacral joint. There
tracts; aids in expiration by pulling the ribs are also independent, more dorsally placed, me
caudally. dial tendons going to the cranial articular proc
Innervation: Dorsal branches of the nn. tho- esses of the seventh, sixth, and fifth lumbar ver
racales and lumbales. tebrae.
The m. longissimus thoracis has serrations
Longissimus System which run to the caudal borders of the ribs by
means of broad tendinous leaves. Each tendi
The m. longissimus (Figs. 3-26, 3-27,3-38, A) nous leaf separates into a medial and a lateral
is the medial portion of the m. erector spinae. terminal tendon, the edges thicken, and between
Lying medial to the m. iliocostalis, its overlap them pass dorsal branches of the thoracic nerves.
ping fascicles extend from the ilium to the head. The medial tendons of these ventral serrations
The m. longissimus consists of thoracolumbar, end on the accessory processes of the thirteenth
cervical, and capital regional divisions. The m. to sixth thoracic vertebrae. Since accessory proc
sacrococcygeus lateralis can be regarded as the esses are lacking from the fifth to first thoracic
caudal continuation of the m. longissimus on the vertebrae, the medial tendons insert on the cau
tail; this muscle is discussed with the tail mus dal ends of the transverse processes. The lateral
cles. tendons of the m. longissimus thoracis insert on
The m. longissimus thoracis et lumborum is the thirteenth to sixth ribs, where they attach
the strongest muscle of the trunk in the lumbar medial to the attachment of the m. iliocostalis on
and thoracic regions. Lateral to the spinous the edge of a flat groove adjacent to the costal
processes of the lumbar and thoracic vertebrae tubercle. Cranial to the sixth rib the muscle be
(which are covered by deeper muscles), and dor comes so narrow that its tendons appear undi
sal to the lumbar transverse processes and the vided. The terminal tendons end on the costal
ribs, it runs from the iliac crest to the last cer tubercles of the fifth to first ribs immediately
vical vertebra. In the lumbar region, it is inti lateral to the costal tubercular joint. Occasion
mately fused with the m. iliocostalis lumborum ally, further divisions of the terminal tendon in
to form the m. erector spinae (m. sacrospinalis). sert on the transverse processes of the sixth and
In the thoracic region the m. spinalis et semi fifth cervical vertebrae, where they fuse with
spinalis dorsi et cervicis arises from its strong serrations of the m. longissimus cervicis.
aponeurotic covering. The thoracolumbar divi Action: Extension of the vertebral column.
sion reaches its greatest development in the cra Raising of the cranial portion of the body
nial part of the lumbar region; in the thoracic from the pelvis, sacrum, and loin; in con
region it gradually narrows, whereas the m. ilio junction with other muscles, fixation of the
costalis gets larger. vertebral column; deflection of the back by
The m. longissimus lumborum (Eisler 1912) is fixation of the cervicothoracic junction;
covered by an exceptionally dense aponeurosis sudden raising of the caudal portion of the
which is separated from the thoracolumbar fas body, which is initiated by means of the
cia by fat. It arises caudally from the iliac crest rear extremities.
M u sc les of the T runk 167
M u l t i f i d us c e r v i c i s
1
Median fib ro u s naphe Licjam entum nuchae
S p i n a l is e t s e m i s p i n a l i s t h o r a c i s et c e r v i c i s
R. s e m i s p i n a I is cap. ( B iventer)
j Loncj i s s i m u s
Re c t u s c a p , d o r s a l i s maj.
Semispin. cap. : i 11i ocost al i s
6 iventer -
Comple xu s
Obl i q_ uus c a p . -
c a u d a l is
Obl iq u us cap,
cram alis
0 motransversar. -
Intertransv. cervicis d a r s a l i s
Intertransversarius in term edius
Semispinolis c a p itis '
L ong i s s i mus ca p itis
I n l e r t r a n s v e r s a r i u s ve n t r a l is
Lonqissim us cervicis
■Scalenus
T r a n s v e r s us c a s t a r u m ’
F ig . 3-27. Muscles of neck and head, deep dissection, lateral aspect.
168 Chapter 3. M yology
In t e r t r a n s v e r s a r i i
Intertransversarii
05
ventralis cervicis Fic. 3-28. Deep axial muscles. ZD
170 Chapter 3. M yology
semispinalis thoracis et cervicis in the dog ex covered by the mm. longissimus and splenius.
tends from the spinous process of the eleventh The muscle lies rather deep as it extends from
thoracic vertebra to the spinous process of the the first five thoracic and the last cervical verte
axis. This combined muscle is clearly separated bra to the occiput. It surrounds each half of the
into lateral and medial parts. The lateral part ligamentum nuchae laterally and dorsally, meet
is the m. spinalis et semispinalis thoracis; the ing its fellow of the opposite side. The two mus
medial part is the m. spinalis cervicis. cles are separated only by the nuchal ligament
The m. spinalis et semispinalis thoracis (Fig. and the median fibrous raphe. It is divided into
3-26) is the lateral part of the compound muscle the dorsally located m. biventer cervicis and the
which arises from the fascia of the m. longissi- ventrally placed m. complexus, which can be
mus. From the spinous processes of the eleventh separated as far as their insertions, despite the
to the seventh thoracic vertebra, this muscle intimate connections between them.
with its nearly horizontal fibers is unsegmented; The m. biventer cervicis (Fig. 3-26) arises, by
from the ninth thoracic vertebra forward it three strong serrations medial to the m. longis
sends off gradually ascending bundles to the spi simus cervicis and capitis, from the transverse
nous processes of the thoracic vertebrae. The processes of the fourth, third, and second thora
muscle divides into eight separate bundles, cic vertebrae. Fascial strands also come from the
which insert on the spinous processes of the lateral surfaces of the spinous processes under
sixth thoracic to the sixth cervical vertebra by neath the m. semispinalis dorsi; other fibers are
means of superficial tendons which become added to the dorsal border from the fascia spino-
progressively more distinct cranially. The seg transversalis at the level of the shoulder. The m.
ment to the sixth cervical vertebra almost com biventer cervicis is firmly connected with the
pletely covers the segment to the seventh cervi median fibrous raphe of the neck. It appears to
cal vertebra. Each ends by well-developed be divided, by four (rarely five) very oblique
tendinous leaves on the spinous processes. Each tendinous inscriptions, into separate portions
also gives off a wide, leaflike portion to the having longitudinal fibers. It inserts on a dis
neighboring caudal, tendinous leaves of the m. tinct, oval, rough area ventrolateral to the ex
spinalis cervicis. In the cranial thoracic region ternal occipital protuberance on the caudal sur
variations may occur, in that intermediate face of the skull.
bundles with their own tendons may appear The m. complexus (Fig. 3-26) arises from the
(Stimpel 1934). caudal articular processes of the first thoracic to
The m. spinalis cervicis (Fig. 3-26) is the me the third cervical vertebra in common with the
dial, flat muscular strand bearing four tendinous m. longissimus capitis (laterally) and the m. mul-
inscriptions. It arises from the tendon of the tifidus cervicis (medially). The caudal segments
most cranial thoracic segment of the m. semispi are more fleshy; the one arising on the first
nalis thoracis and from the cranial border of the thoracic vertebra has a tendinous covering me
first thoracic spine, but it also receives a few dially which is also related to one of the por
bundles from the spinous process of the seventh tions of the m. multifidus. Fibers also arise in
cervical vertebra. Separated from the muscle of the fascia of the m. obliquus capitis caudalis
the opposite side only by the median ligamen somewhat cranial to the caudal border of the
tous septum, it runs cranially ventral to the lig- atlas. The fibers run craniomedially to end lat
amentum nuchae. It inserts on the spinous proc erally on the dorsal nuchal line by means of a
esses of the fifth to second cervical vertebrae; it tendon coming from a strong superficial fibrous
is covered in part by portions of the m. multifi- covering.
dus. Action: To raise the head and neck; in unilat
Action: To fix the thoracic vertebral column eral action to flex the head and neck later
and to raise the neck. ally.
Innervation: Medial branch of the dorsal Innervation: Dorsal branches of the nn. cervi
branches of the cervical and thoracic cales.
nerves. The m. multifidus (Figs. 3-28, 3-38, A) is
The m. semispinalis capitis (Figs. 3-26,3-27) a muscle composed of numerous individual por
is the strong continuation to the head of the m. tions which extend from the sacrum to the sec
spinalis et semispinalis thoracis et cervicis. The ond cervical vertebra; it is augumented at both
capital portion of the semispinalis strand covers joints of the head, as well as in the tail, by more
the cranial end of the m. spinalis et semispinalis or less modified muscles—at the head by the
thoracis et cervicis laterally; its broad origin is mm. obliquus capitis caudalis and cranialis, in
M u sc les of the T run k 171
the tail by the m. sacrococcygeus dorsalis medi- differentiated; these are the mm. rotatores longi
alis. The m. multifidus as a segmental muscle ex and breves. In addition, there are the mm. inter-
tends from mammillary, transverse, or articular spinales lumborum, thoracis, and cervicis be
processes of caudally lying vertebrae to spinous tween the spinous processes, and the mm. inter -
processes of cranially lying ones. As a rule, two transversarii coccygeus, lumborum, thoracis,
vertebrae are passed over by each bundle. The and cervicis, which in general run between the
m. multifidus, aside from the oblique capital transverse processes. The intertransverse mus
muscles, is divided into four portions: the pars cles of the tail are described with the coccygeal
lumborum, pars thoracis, pars cervicis, and the muscles.
pars coccygeae, which is described with the coc The mm. rotatores (Fig. 3-28) are developed
cygeal muscles as the m. sacrococcygeus dorsalis as eight long and nine short rotators; in the dog
medialis. they are confined strictly to the cranial thoracic
The m. multifidus lumborum is a strong, region, where the pairs of articular processes are
seemingly homogeneous muscle which runs tangentially placed, thus allowing rotatory
from the sacrum to the spinous process of the movements.
eighth or ninth thoracic vertebra. It is divided The mm. rotatores longi extend between the
into eleven individual, flat portions which are transverse and spinous processes of two alternate
united with each other. They originate from the vertebrae; the most caudal extends from the
three articular processes of the sacrum (includ transverse process of the tenth to the spinous
ing the mammillary process of the first coccygeal process (basal to the insertion of the correspond
vertebra) and from the mammillary processes of ing segment of the multifidus) of the eighth tho
the seventh lumbar to the twelfth thoracic ver racic vertebra, and the most cranial extends be
tebra. After the several parts pass over two seg tween corresponding points of the third and the
ments, they end laterally on the ends of the spi first thoracic vertebra. These segments are more
nous processes of the sixth lumbar to the ninth vertical than those of the m. multifidus, along
(occasionally eighth) thoracic vertebra immedi the caudal border of which they appear.
ately beneath the supraspinous ligament. The mm. rotatores breves pass between verte
The m. multifidus thoracis lies more ventrally brae. They are situated more deeply than are
on the vertebral column, and its segments are the long rotators. The most caudal belly runs be
more vertical than those of the lumbar part. It tween the transverse process of the tenth and
arises by nine distinctly isolated portions on the the spinous process of the ninth thoracic verte
mammillary and transverse processes of the bra, the most cranial belly between similar
eleventh to the third thoracic vertebra and in points on the second and the first thoracic verte
serts on the spinous processes of the eighth tho bra. Often this portion is surrounded extensively
racic to the seventh cervical vertebra. by tendinous tissue (Kruger 1929).
The m. multifidus cervicis is covered by the Action: Rotation of the greater cranial portion
m. semispinalis capitis. It appears under the ven of the thoracic vertebral column about the
trolateral border of the m. spinalis et semispinalis longitudinal axis in unilateral action; other
thoracis et cervicis, where it extends from the wise, fixation.
articular process of the second thoracic vertebra Innervation: Medial branches of the rami dor-
to the spinous process of the axis. It consists of sales of the thoracic nerves.
six incompletely separable individual portions The mm. interspinales (Fig. 3-28) are dis
which themselves are again partially divided tinctly separable into lumbar, thoracic, and cer
into lateral principal, medial accessory, and deep vical portions; the lumbar portion is covered by
accessory parts, according to Stimpel (1934); the m. multifidus. In the thoracic region, after
collectively they arise essentially from the artic removal of the mm. semispinalis and longissi-
ular processes. mus, the mm. interspinales are visible at the
Action: As a whole, the m. multifidus, along ends of the spinous processes. They run be
with the other dorsal back muscles, fixes the tween contiguous edges of spinous processes
vertebral column, especially in bilateral ac and overlap these edges somewhat. They also
tion. extend between the spinous processes of the first
Innervation: Medial branches of the rami dor- thoracic to the fifth cervical vertebra.
sales in the lumbar, thoracic, and cervical Action: Fixation of the vertebral column.
regions. Innervation: Medial branches of the rami dor-
From the medial surface of the m. multifidus sales of the spinal nerves.
certain deep muscles have become extensively The mm. intertransversarii (Fig. 3-28) are
172 Chapter 3. M yology
deep segments split off from the longissimus sys run cranially from the m. scalenus and form a
tem. They are separable into coccygeal, lumbo- homogeneous longitudinal strand. This is found
thoracic, and cervical parts, and, as delicate ventral to the m. scalenus and dorsal to the m.
muscle bundles, they pass over one or two, or, at longus colli; it extends from the ventral border
most, three vertebrae. of the winglike transverse process of the sixth
The mm. intertransversarii coccygei are dis cervical vertebra to insert by three separate ter
cussed with the muscles of the tail. minal segments on the caudal branch of the
Only weak mm. intertransversarii lumborum transverse process of the fourth, third, and sec
et thoracis are formed on the trunk. These sep ond cervical vertebrae. This strand is covered by
arate parts run between the mammillary proc the m. scalenus caudally and by the intermedi
esses of the seventh lumbar to the thirteenth or ate portion of the mm. intertransversarii crani
twelfth thoracic and the accessory processes of ally.
the fifth lumbar to the ninth thoracic vertebra, At its cranial end the cervical vertebral col
and between the transverse processes of the umn serves special functions. There is a corre
twelfth to the eighth and those of the eighth to sponding special development of the first two
the fourth thoracic vertebra. cervical vertebrae, as well as of their joints. The
The mm. intertransversarii cervicis are larger. specialized musculature dorsal and ventral to
They are divided into three separate muscle the atlas and axis is adapted to these special
strands: a dorsal one, running between articular functions. There are three portions of the m.
and transverse processes of the cervical verte rectus capitis which run between regions on the
brae, and, corresponding to the intertransverse spine of the axis, the atlas, and the occiput, and
muscle of the lumbar and thoracic regions, an which can be compared to the m. interspinalis;
intermediate strand, and a ventral strand. these are the mm. rectus capitis dorsalis major,
The mm. intertransversarii dorsalis cervicis lie intermedius, and minor. There are also two
between the lines of insertion of the mm. longis oblique muscles, the mm. obliquus capitis cau
simus cervicis, longissimus capitis, and semi dalis and cranialis, which can be considered
spinalis capitis. As a segmental muscle strand modifications of the m. multifidus or derivatives
it extends from the eminence on the cranial of the m. intertransversarius.
articular process of the first thoracic vertebra to The m. rectus capitis dorsalis major (Fig. 3-
the wing of the atlas. Its individual bundles run 27) is covered by the m. semispinalis capitis as it
craniolaterally in the form of five indistinctly runs between the spine of the axis and the
separated bellies from the first thoracic and the squama of the occiput. It arises cranial to the at
seventh to fourth cervical vertebrae to the trans tachment of the ligamentum nuchae on the cau
verse processes of the sixth to second cervical dal end of the spine of the axis, and it ends me
vertebrae. The cranial portion of the muscle ex dially on the occiput. The dorsal portion of the
tends from the eminence of the third and second m. obliquus capitis cranialis, which lies on the
cervical vertebrae to the caudal border of the border of the wing of the atlas, also inserts on
wing of the atlas. the ventrolateral part of the occiput.
The mm. intertransversarii intermedii cervicis The m. rectus capitis dorsalis intermedius is
form a strand which is composed of five or six a strong, almost triangular muscle. Covered by
distinctly separable, delicate parts which extend the major part, it arises cranially on the axis and
only between transverse processes; they lie ven with diverging fibers runs over the atlas to the
tral to the insertion of the m. serratus ventralis occiput where it inserts on the ventral nuchal
cervicis and dorsal to the m. scalenus, and are line. It is obviously only the deeper part of the
partly covered by these two muscles. The seg m. rectus capitis dorsalis major, although it fuses
ments course between the terminal tubercles of in part with the small extensor of the head.
the ends of the transverse processes from the The m. rectus capitis dorsalis minor is a short,
first thoracic to the second cervical vertebra. On flat muscle, lying between the atlas and the oc
the sixth cervical vertebra it is on the transverse ciput on the capsule of the atlanto-occipital joint
process proper, and, from the fifth cervical ver immediately next to its fellow of the opposite
tebra forward, it is on the caudal branch of the side. It arises on the cranial edge of the dorsal
transverse process and the border of the wing of arch of the atlas and inserts above the foramen
the atlas; the most cranial portion runs under magnum near the ventral nuchal line, where it
the dorsal m. intertransversarius of the axis. The fuses with the intermediate extensor of the head.
deep fibers pass from segment to segment; the Action: All three portions extend the atlanto-
superficial ones pass over one segment. occipital joint.
The mm. intertransversarii ventralis cervicis Innervation: Ramus dorsalis of n. cervicalis 1.
M u sc les of the T runk 173
The m. obliquus capitis caudalis (Fig. 3-27) is sternothyroideus arises from the first costal car
a strong, flat muscle lying under the mm. semi tilage, and passes up the neck covered by the m.
spinalis capitis and splenius; it covers the atlas sternocephalicus. Although weaker than the m.
and axis dorsally. It arises along the entire spi sternohyoideus, it covers more of the lateral sur
nous process and the caudal articular process of face of the trachea. It inserts on the lateral sur
the axis and runs obliquely craniolaterally over face of the thyroid lamina.
the capsule of the atlanto-axial joint to insert on A ction: To draw the hyoid apparatus, larynx,
the border of the wing of the atlas near the alar and tongue caudally.
notch. Innervation: Ramus ventralis of n. cervicalis 1.
Action: Unilateral: rotation of the atlas and The m. scalenus (Figs. 3-24, 3-30) bridges
thus the head on the axis; bilateral: fixation the space between the first three ribs and the
of the atlanto-axial joint. cervical vertebrae. The muscle is divided into a
Innervation: Rami dorsales of the nn. cervi superficial and a deep portion.
cales 1 and 2. The m. scalenus prim ae costae arises as three
The m. obliquus capitis cranialis (Fig. 3-27) portions on the cranial border of the first rib.
extends obliquely craniolaterally over the at The two deep segments, which are not clearly
lanto-occipital joint; it lies under the m. splenius separated, run to the transverse process of the
and is divided into two portions. The principal seventh cervical vertebra and to the wing-like
part arises on the lateroventral surface and lat process of the sixth cervical vertebra. In its
eral border of the wing of the atlas. Inclined dor- course it is covered proximally by the supracos-
somedially, it runs over the jugular process and tal part of the m. scalenus, and distally by a
inserts on the mastoid part of the temporal bone superficial portion (also arising on the first rib).
and from there upward on the dorsal nuchal The superficial segment, on the other hand, ex
line. The accessory portion is a superficial flat tends from the first rib to the transverse proc
belly extending to the atlantal end of the m. esses of the fifth, fourth, and third cervical ver
obliquus capitis caudalis; it takes its origin on the tebrae.
tip of the wing of the atlas and, provided with The m. scalenus supracostalis forms the prin
tendinous leaves, inserts between the principal cipal part of the superficial layer. In the form of
portion and the m. rectus capitis dorsalis major two or three flat, distinctly separated portions i t
on the dorsal nuchal line. arises from the outer surfaces of the ribs and in
Action: Extension of the adanto-occipital joint. serts on the cervical vertebrae. In so doing it
Innervation: Ramus dorsalis or n. cervicalis 1. covers parts of the m. scalenus primae costae,
the caudal portions of the m. intertransversarius
M u scles o f t h e V e n t r a l N e c k R e g io n intermedius, and the first three or four serrations
of the m. serratus ventralis. This part of the m.
The muscles of the ventral neck region are di scalenus attaches to the transverse processes
vided into two groups. The first group is closely (caudal branches) of the fifth and fourth (and
related to the trachea and esophagus and in also, occasionally the third) cervical vertebrae.
cludes the large superficially located mm. bra- The dorsal portion arises underneath the corre
chiocephalicus and sternocephalicus and the sponding segment of the serratus on the third
small, deeply located mm. sternohyoideus, ster- rib and the middle portion on the fourth; both
nothyroideus, and scalenus. The second group in may arise in common on the fourth rib. The ven
cludes muscles that lie on the ventral surfaces of tral portion is the longest; its origin from the
the cervical vertebrae: the mm. longus capitis, eighth or ninth rib, by means of a long tendinous
longus colli, rectus capitis ventralis, and rectus leaf, is covered by the m. obliquus abdominis ex
capitis lateralis. ternus.
The m. brachiocephalicus and m. stemoceph- Action: To draw the neck downward. In uni
alicus are described with the muscles of the tho lateral action, to bend the neck sideward.
racic limb. When the neck is fixed, the supracostal part
The m. sternohyoideus is closely allied with can act in inspiration.
the m. sternothyroideus throughout its course. Innervation: Rami ventrales of the nn. cervi
It is described with the muscles of the hyoid ap cales and thoracales.
paratus. The m. longus capitis (Figs. 3-27, 3-29) is a
The m. sternothyroideus (Figs. 3-9, 3-24, 3 - long, flat muscle which lies on the lateral and
46) lies deep to the m. sternohyoideus and has a ventral sides of the cervical vertebrae lateral to
similar tendinous intersection which divides the the m. longus colli. It arises from the caudal
muscle into cranial and caudal portions. The m. branches of the transverse processes of the sixth
174 Chapter 3. M yology
R e c t u s c a p i t is
- -v e n t r a lis
R e c t u s c a p it is
I a t e r a ! is
L a n g u s c a p i t is
p u l l e d l a t e r a lly
- L o n g u s c a lli
- T ra n s v e rs e p r o c e s s
o f 6 thc e r v ic a l
v e r te b r a
S c a le n u s
p n m a e c a s to e
to the second cervical vertebra, and extends branch of the first cervical nerve). It originates
cranially to the axis, where it receives a strong, on the ventral surface of the caudal half of the
tendinous leaf laterally. After crossing the wing of the atlas (lateral to the n. rectus capitis
atlanto-occipital joint, it inserts (tendinous later ventralis); it passes sagittally toward the cranium
ally, muscular medially) on the muscular over the atlanto-occipital joint, and inserts on
tubercle of the basioccipital, between the tym the base of the jugular process of the occiput.
panic bullae. This muscle can be considered a special portion
Action: To flex the atlanto-occipital joint and of the m. intertransversarius ventralis.
to draw the neck downward. Action: Flexion of the atlanto-occipital joint.
Innervation: Rami ventrales of the nn. cervi- Innervation: Ramus ventralis of n. cervicalis 1.
cales.
The m. longus colli (Fig. 3-29) is a long mus T h e F a s c ia e o f t h e N eck
cle composed of separate bundles; it lies adja
cent to its fellow on the bodies of the first six The superficial and deep cervical fasciae are
thoracic and all of the cervical vertebrae, and the direct continuations of the superficial and
thus is divided into thoracic and cervical por deep fasciae of the head.
tions. On the neck the bilateral muscle is en The superficial fascia of the neck (fascia colli
closed by the right and the left m. longus capitis. superficialis) is cylindrical in form as it clothes
The thoracic portion consists of three incom the whole neck. It is delicate, lies directly under
pletely separated parts which arise on the the skin, and is easily displaced. It originates from
convex ventral surfaces of the first six thoracic the superficial temporal, parotideomasseteric
vertebrae. These portions, complicated in their and intermandibular fasciae, and it continues
make-up, are provided with tendinous coverings. caudally into the superficial omobrachial fascia
Diverging cranially from those of the opposite and ventrally into the superficial trunk fascia of
side, the fibers of these three portions become the sternal region. It contains the m. sphincter
partly tendinous laterally; the medial fibers in colli superficialis and the platysma; with these it
sert immediately beside this tendon on the covers the mm. trapezius, omotransversarius,
ventral border of the wing of the sixth cervical cleidocephalicus, and sternocephalicus, and it
vertebra, as well as on the transverse process of bridges the external jugular vein which lies in
the seventh cervical vertebra. The continuation the jugular groove. The bilateral portions of the
of the cervical portion consists of four separate fascia meet dorsally and ventrally. At the dorsal
bundles. These bundles arise on the ventral mid line there is no special attachment to the
border of the transverse process of the sixth to underlying portions (median raphe) so that on
the third cervical vertebra and end on the ven the neck, just as on the back and loins, this
tral spine of the next preceding vertebra. The fascia can be lifted in a big fold with the skin. In
caudal V-shaped segment formed by the mus many places the smaller cutaneous vessels and
cles of both sides has lying in its angle the nerves pass through the superficial fascia.
cranial part of the thoracic portion. The cranial The deep fascia of the neck (fascia colli pro
segment ends on the ventral tubercle of the funda) is a strong binder which extends under
atlas. the mm. sternocephalicus, cleidomastoideus,
Action: To draw the neck downward. omotransversarius, and cleidocervicalis; it covers
Innervation: Rami ventrales of the nn. cervi- the mm. sternohyoideus and sternothyroideus
cales. superficially and surrounds the trachea, thyroid
The m. rectus capitis ventralis (Fig. 3-29) is a gland, larynx, and esophagus, but passes over
short, strong muscle which lies dorsal to the end the large cervical vessels and nerves (nn. vago-
of the m. longus capitis. It extends from the sympatheticus and recurrens, a. carotis com
ventral arch of the atlas to the basioccipital munis, and v. jugularis internus) to the superfi
bone. As it crosses the atlanto-occipital joint, it cial surface of the mm. scalenus and longus colli.
converges somewhat with its fellow of the op From these it goes to the superficial surface of
posite side. the mm. serratus ventralis and rhomboideus, to
Action: Flexion of the atlanto-occipital joint. end in the median raphe of the neck. Cranially,
Innervation: Ramus ventralis of the n. cervi- it continues as the external pharyngeal fascia
calis 1. and passes under the mandibular and parotid
The m. rectus capitis lateralis (Fig. 3-29) is a glands and finally ends on the hyoid bone and
small muscle which lies lateral to the m. rectus the base of the head. Under the salivary glands
capitis ventralis (separated from it by the ventral the deep cervical fascia is united with the deep
176 Chapter 3. M yology
fascia of the m. masseter. The dorsal part of the on the thorax, the m. transversus costarum
deep cervical fascia continues over the m. covers the superficial (ventral) ends of the first
rhomboideus to the superficial surface of the ribs; the m. transversus thoracis crosses the carti
scapula with its muscles, and thus runs into the lages of the sternal ribs and the sternum deeply.
deep fascia of the shoulder. On the surfaces of The mm. retractor costae and subcostalis are
the cervical parts of the mm. serratus ventralis special muscles of the last rib.
and scalenus, the deep cervical fascia continues The mm. intercostales externi (Fig. 3-32)
to the thoracic parts of these muscles medial to form the stronger outer layer of the intercostal
the shoulder to become the deep thoracic fascia. spaces; they are 4 or 5 mm. thick in large dogs,
Ventral to the m. scalenus it attaches to the first but become weaker in the region of the floating
rib and the manubrium sterni. The deep fascia ribs. They extend from the mm. levatores cos
sends various divisions between the layers of tarum, which are indistinctly set apart, to the
muscles of the neck. One of these is a strong costochondral junctions; they may also extend
leaf which passes beneath the cervical parts of into the spaces between the costal cartilages as
the mm. serratus ventralis, trapezius, and rhom the mm. intercartilaginei. The fibers of the ex
boideus but external to the m. splenius. This, the ternal intercostal muscle arise on the caudal
spinotransverse fascia (fascia spinotransversalis), border of each rib, and run caudoventrally to
is an important fascial leaf in muscle dynamics. the cranial border of the next rib.
One part of the deep cervical fascia ventral to Action: Inspiration. For both external and in
the vertebral column is the prevertebral fascia ternal intercostal muscles, the more proxi
(fascia prevertebralis), on which the superficial mal attachment is to be looked upon as the
surface of the mm. longi colli lies. Cranially, it is fixed point, since, for each arched rib, the
attached to the base of the head; caudally, it points farther from the tubercle are rela
continues with the mm. longi colli over the first tively easier to move than those closer to
six thoracic vertebrae into the thorax to unite the tubercle. Thus the m. intercostalis ex-
with the endothoracic fa scia (fascia endotho- temus acts to increase the transverse di
racica). ameter of the thorax.
Finally, from the deep leaf of the cervical Innervation: Muscular branches of the nn.
fascia, there detaches a delicate proper fascia o f intercostales 1 to 12.
the trachea (fascia tracheae propria), which sur The mm. intercartilaginei externi are unsepa
rounds the trachea. Much loose connective tis rated continuations of the mm. intercostales ex
sue is accumulated around both the trachea and terni into the interchondral spaces. They are
esophagus, to provide them with a high degree lacking in the first two or three spaces because
of displaceability. The carotid sheath is a special the external intercostals end proximal to or at
loose condensation of fascia in which the com the costochondral junctions. Distal to the ends
mon carotid artery, internal jugular vein, lym of the external intercostals the internal inter
phatics (tracheal duct), and the vagosympa costals make their appearance. With each suc
thetic trunk are located. It is located dorsolateral cessive segment, the external interchondral mus
to the trachea and is the fascial union of the cles extend farther distally, so that the ninth and
lateral and prevertebral parts of the deep cervi tenth interchondral spaces are completely filled,
cal fascia. although occasional defects in the muscle are
found. Although they are rather strongly de
M u sc l e s o f t h e L a t e r a l a nd veloped in the false or asternal interchondral
V e n t r a l T h o r a c ic W a l l spaces, the muscle is completely absent in the
twelfth interchondral space.
The spaces between the ribs are filled by the A ction an d Innervation: Same as for the mm.
mm. intercostales, which appear in a double intercostales externi.
layer, internal and external, and cross each The mm. levatores costarum (Fig. 3-32) are
other. Where these muscles occur between the present as twelve special formations of the ex
costal cartilages, they are specifically called mm. ternal intercostal muscles. They are flat, spindle-
intercartilaginei. Each m. intercostalis externus shaped muscles covered by the mm. longissimus
gives rise to a m. levator costarum proximally. thoracis and iliocostalis; they are fleshy at their
The fibers which make up its almost spindle- origins on the transverse processes of the first to
shaped belly do not come from the following rib, twelfth thoracic vertebrae. After running caudo
but come rather from the transverse process of ventrally to the angle of the rib next caudad,
the corresponding thoracic vertebra. Cranially, they end on the cranial borders of the second to
M u sc les of the T runk 177
L o n g is s im u s II l i o c o s t a li s
I
S p in a lis e t s e m is p in a lis S e r r a t u s dorsalis c a ud alis
I A
Se rra tus dorsalis c ro n ia l is 1 '
Serratus~ - -Obi i q u u s
ventralis, cut i n t e r n u s abd.
S ca len us- -
Obliquus
externus abd.
In te rc o s ta lis e x te rn u s
Fic. 3-31. Superficial muscles of thoracic cage, lateral aspect.
L e v a ta r c o sta e
i
- - In t e r c o s t a I is
e x te rn u s
iI n t e r c o s t a l i s i n t e r n u s
F ig . 3 -3 2 . Deep muscles of thoracic cage, lateral aspect.
178 Chapter 3. M yology
thirteenth ribs. They then pass over into the The m. retractor costae (Fig. 3-33) is a thin
mm. intercostales externi. muscle lying under the tendon of origin of the
A ction: Inspiration; the fixed point is the m. transversus abdominis. It bridges the space
transverse process of the vertebra. between the transverse processes of the first 3 or
Innervation: Delicate little trunks of the nn. 4 lumbar vertebrae and the last rib (Iwakin
intercostales 1 to 12, given off shortly be 1928). Seen from the interior, this thin muscle,
fore their division into lateral and medial the fibers of which cross those of the m. trans
branches. versus abdominis, lies directly under the peri
The mm. intercostales interni (Figs. 3-32, 3 - toneum. Over its cranial border lies the arcus
34) form the weaker internal layer of the inter lumbocostalis of the diaphragm. Farther distally
costal musculature; this layer is 2 or 3 mm. thick the caudal fiber bundles extend upon the last rib
in large dogs. The internal intercostals extend and partly encroach upon the peritoneal surface
from the vertebral column, where they leave of the pars costalis of the diaphragm. The m.
free only a small triangular space adjacent to the retractor costae belongs to the system of the m.
vertebrae, to the distal ends of the ribs; they are intercostalis internus and is innervated by the
only very slightly separated from the interchon- last thoracic nerve (Kolesnikow 1928).
dral muscles. The fibers course from the cranial The m. transversus thoracis (Fig. 3-34) is a
border of one rib to the caudal border of the rib flat, fleshy muscle, lying on the inner surfaces of
next cranial to it. In this cranioventral course the the sternum and sternal costal cartilages. It
fibers attain angles of inclination which, at the forms a continuous triangular leaf which covers
vertebral column, decrease from 78 to 71 de the second to eighth costal cartilages. Only ex
grees, and, at the sternum, from 68 to 54 degrees. ceptionally do slips from its lateral border reach
Thus, they are steeper than the mm. inter the sternum. A delicate, special bundle maybe
costales externa, which they cross. given off to the first costal cartilage. Its fibers
Action: Expiration. arise by a narrow aponeurosis, on the lateral in
Innervation: Nn. intercostales. ternal surface of the sternum, from the second
The mm. intercartilaginei interni are contin sternebra to the caudal end of the xiphoid proc
uations of the mm. intercostales interni. They ess. They end with indistinct segmentations on
fill the interchondral spaces and are 4 or 5 mm. the second to seventh costal cartilages, some
thick. After removal of the m. rectus abdominis, what ventral to the costal symphyses.
they appear at the sites where the external inter
chondral muscles are lacking. D ia ph r a g m
Action and Innervation: Same as for the mm.
intercostales interni. The diaphragm (diaphragma) (Figs. 3-33, 3 -
The m. transversus costarum (Figs. 3 -2 4 ,3 - 34) is a musculotendinous plate between the
31) is a flat, almost rectangular muscle which thoracic and abdominal cavities; it projects for
runs caudoventrally. It covers the cranial part of ward into the thoracic cavity like a dome. On
the lateral surface of the thorax from the origin the thoracic side, it is separated from the pleura
of the ventral portion of the m. scalenus supra- by the endothoracic fascia; on the abdominal
costalis on the first rib to rib 3 or 4. Here it joins side, it is separated from the peritoneum by the
the deep fascia of the trunk and radiates into the transversalis fascia. Peripherally, this wall which
m. obliquus externus abdominis. It covers the separates the body cavities attaches to the ven
tendon of origin of the m. rectus abdominis tral surfaces of the lumbar vertebrae, the ribs,
superficially. and the sternum. The fibers of the diaphragm
This muscle may correspond with the rarely arise on these skeletal parts and radiate toward
occurring m. stemalis of man. Artemenko (1929) the tendinous center.
regards it as a division of the m. obliquus ex The central tendon of the diaphragm, in the
ternus abdominis. The belief of other authors dog, is relatively small. It consists of a triangular
who relate it with the m. rectus abdominis is, body, the blunt point of which is directed ven
fallacious (Zimmermann 1927); the muscle thus trally, and two, steeply rising, slender, pointed
does not warrant the name m. rectus thoracis. columns, which are rather large. From the
The m. transversus costarum has no genetic re cranial aspect this tendinous area appears to be
lationship to the m. scalenus. displaced somewhat ventrally. The two-layered
Action: Inspiration. disposition of the tendon fibers is easily followed.
Innervation: Lateral branch of the nn. inter To the right, at the base of the body of the
costales. tendon, there is a concentric arrangement of
M u sc les of the T runk 179
-C o sta l a rc h
W i
--C e n tra l te n d o n
A o rtic h i a t u s
T r a n s v e r s u s t h o r a c is
i
R e t r a c t o r c o s ta e In te rc o sta lis
ii n t e r n u s
L e ft c ru s
-R ig h t crus
-T ronsversus
abdom in is
P o s tc a v o -
E sophogus-
A o rta
13 ih r i b -
strong fibers about the foramen venae cavae further differentiated into a lateral, intermediate,
which courses slightly cranioventrally. On the and medial portion.
columns of the tendon, fibers run in an arch The crus laterale o f the right diaphragm atic
from the crural musculature directly to those of crus originates mainly from the tendon of origin
the costal parts. Special strong reinforcements coming from the third lumbar vertebra. It ex
extend lengthwise along the borders. Fibers tends ventral to the psoas muscles in an almost
from the muscle surrounding the esophagus transverse arch—the arcus lumbocostalis. The
radiate on the body of the tendon to the sternal pleura and peritoneum encroach directly upon
and ventral parts of the costal diaphragmatic one another dorsal to the arch. After crossing the
musculature. Transverse fibers course from one lumbar musculature, the fiber bundles of the
side to the other as a reinforcing apparatus. lateral crus run toward those of the pars costalis
Peculiar whorls are formed near the bases of with which they coalesce into a narrow tendi
both columns. Into the dorsal border of the nous band, the extension of the end of the column
foramen venae cavae, muscle fibers from the of the tendinous center. In the wedge between
costal portion often radiate (Pancrazi 1928). these portions is a triangular area which is free
The muscular part of the diaphragm sur of muscle—the trigonum lumbocostale; only
rounds the central tendon on all sides, and its fascial coverings of the diaphragmatic muscula
fibers stream into the latter in a radial direction. ture radiate into it. This portion of the peripheral
It is divided into the pars lumbalis, a pars costalis diaphragmatic attachment crosses the m. retrac
on each side, and the pars sternalis, all of which, tor costae and the last rib ventrally. On each
with the exception of the lumbar portion, have a side the splanchnic nerves and the sympathetic
uniform thickness of 3 or 4 mm. in large dogs. trunk cross dorsal to the lumbocostal arch.
The pars lumbalis of the diaphragmatic mus The crus laterale o f the left diaphragmatic
culature is formed by the right and left dia crus arises in a similar way from its correspond
phragmatic crura. At the aortic hiatus (hiatus ing tendon; however, it has another special
aorticus) they enclose the aorta, the azygos and lateral division which radiates into the lumbo
hemiazygos veins, and the lumbar cistern of the costal arch from the ventral border of the psoas.
thoracic duct. Although at first glance they ap Thus on the left side the trigonum lumbocostale
pear to be symmetrical, they are not symmetrical is muscular; therefore the relationships of the
in their construction or in the strength of their left crus laterale correspond entirely to those in
fibers. The right crus is considerably larger than man. The course of the fibers into the tendinous
the left. Each crus arises sagittally by a long center is the same as on the right side.
bifurcate tendon, one part of which is longer The crus intermedium of the right diaphrag
and stronger and comes from the cranial edge of matic crus derives its fibers from the principal
the body of the fourth lumbar vertebra. The part of the tendon of origin and from the right
shorter and somewhat weaker part of the tendon column of the tendinous aortic ring. On the left
comes from the body of the third lumbar verte side, the fibers of this part come from the left
bra. Both portions of the tendon of each side column of the ring along its entire length. These
unite (the right is considerably stronger) to form fiber masses, only indistinctly separable from
an almost sagittal tendon which appears medial those of the crus mediale, radiate into the medial
to the m. psoas minor. The bilateral tendons borders of the bilateral columns of the central
press closely against the aorta, and, from their tendon.
lateral surfaces in particular, they give rise to The musculature of the crus mediale is the
more and more muscle fibers. This results in a thickest (5 or 6 mm.) and originates asymmetri
flat, fan-shaped muscle which bears a medial cally on the two sides. On the right side its fibers
tendon of origin. The muscle parallels the dorsal originate from the terminal portion of the right
thoracic wall. A tendinous strand descends on column of the aortic ring; on the left side the
each side of the aorta, immediately anterior to fibers originate from the apical portion of the ring
the celiac artery, to form the aortic ring. Seen underneath the aorta itself. With blunt edges
from the abdominal cavity, each crus of the dia facing each other, the two parts extend ventrally
phragm is a triangular muscle plate whose until they reach the dorsal border of the body of
borders give rise to the tendinous portions; as a the central tendon. The thick borders of the
whole, this plate of muscle radiates forward medial crura are fused by means of fibrous
toward the concavity of the diaphragmatic ten tissue. Distally they separate for the trans
don. The muscle fiber arrangement is somewhat mission of the esophagus with its vessels
different in the two crura, although each is and the two vagal trunks, thus forming the
M u sc les of the T run k 181
hiatus esophageus. Ventral to this, they fuse the mid-plane the diaphragm forms an arch
by partial crossing of the fiber bundles. There is bulging into the thoracic cavity; this arch extends
an evident asymmetry in the development of the freely downward from the first few lumbar
right and left diaphragmatic crura: from the vertebrae, passing cranioventrally over more
tendon of origin of the right crus come all three than half of the height of the thoracic cavity;
crura dextra and, in addition, the crus mediale near the sternum it turns in a caudoventral di
sinistrum. From the tendon of the left diaphrag rection. The summit of the diaphragm comes to
matic crus come the crus intermedium sinistrum lie at the junction of the middle and ventral
and a portion of the crus laterale sinistrum. thirds of the muscle. On expiration the dia
The generally homogeneous pars costalis on phragm protrudes farthest into the thoracic
each side consists of fibers radiating from the cavity and recedes on inspiration. The dia
costal wall to the tendinous center. This muscle phragm undergoes an excursion of at least 1J
arises by indistinct serrations from the medial thoracic segments at each respiration.
proximal part of the thirteenth rib, distal part of The muscle of the diaphragm is covered on
the twelfth rib, costochondral junction or sym the convex thoracic side by the fascia endo-
physis of the eleventh rib, as well as the whole thoracica and the pleura, on the concave ab
length of the tenth and ninth, and at the bend on dominal side by a continuation of the fascia
the eighth costal cartilage. In the caudal part of transversalis and the peritoneum. Both the fascia
the line of origin the serrations encroach distally and serosa are so thin in the dog that over the
on those of the m. transversus abdominis. In the tendinous portion they can only be seen micro
region of the tenth, ninth, and eighth costal carti scopically.
lages (often only the eighth alone) openings may The convex thoracic side of the diaphragm
be found which allow the passage of the first lies against the surfaces of the lungs, from which
three cranial serrations of the m. transversus ab it is separated by a capillary space. At about the
dominis. The serrations of the diaphragm reach mid-plane of the thorax the mediastinum de
beyond those of the m. transversus abdominis scends from the thoracic vertebrae; the two
and insert cranial and caudal to them on the cor pleural leaves on either side of the mediastinum
responding costal cartilages. Interspersed with separate on the diaphragm to become its pleural
many radial, fatty strands, the bundles of the covering. The attachment of the mediastinum is
costal part run centrally into the lateral borders median only from the dorsal portion of the dia
of the columns and body of the central tendon. phragm to the esophagus. Ventral to the esopha
The pars sternalis of the diaphragm is an un gus the mediastinum makes a strong deflection
paired medial part unseparated from the bi to the left, to return to the mid-plane just above
lateral costal portions. Its fibers arise on the base the sternum. Here the pleura connecting to the
of the xiphoid cartilage, the adjacent transver- postcava branches off in a convex arch.
salis fascia, and the eighth costal cartilages. They In the dorsal part of the mediastinum the
extend dorsally to the apex of the body of the aorta, the azygos and hemiazygos veins, and the
central tendon. thoracic duct extend to the hiatus aorticus. The
The diaphragm projects far into the thoracic esophagus passes to the hiatus esophageus with
cavity, and its costal part lies on the internal sur the dorsal and ventral vagal trunks. On the right
face of the last few ribs. A capillary space be side the esophagus is covered by pleura, which
tween the diaphragm and the ribs, the recessus comes from the mediastinum. In the ventral part
phrenicocostalis, is thus formed. This decreases of the mediastinum the left phrenic nerve lies in
on inspiration but increases in size on expiration. its own mediastinal fold, and the phrenicoperi-
During active flattening of the summit of the cardial ligament runs to the diaphragm near the
diaphragm, the inflated lung pushes into the mid line. The postcava and the right phrenic
opened space, and upon cessation of the dia nerve reach the diaphragm in the plica vena
phragmatic action it is again pushed out of the cava. The stomach and liver attach by ligaments
space. Even during the most extreme inspira to the concave peritoneal surface of the dia
tion the space is not entirely filled by the lung. phragm.
Similar relationships exist in the region dorsal to Action: Retraction of the diaphragmatic sum
the diaphragmatic crura over which (along the mit and thus inspiration.
vertebrae covered by the psoas muscles) a bi Innervation: Nn. phrenici (from the ventral
lateral recessus phrenicolum balis extends back branches of the fifth, sixth, and seventh
ward to the middle of the lumbar vertebrae. In cervical nerves).
182 Chapter 3. M yology
M u sc les o f th e A b d o m in a l W all wall, 6 to 8 cm. from the mid line in large dogs,
it forms a wide aponeurosis. This can be differ
From without inward the abdominal muscles entiated into an abdominal and a pelvic tendon,
are: the obliquus externus abdominis, the ob separated by means of the superficial (subcutane
liquus internus abdominis, the rectus abdominis, ous or external) inguinal ring.
and the transversus abdominis. The m. rectus The abdom inal aponeurosis, or tendon, is by
abdominis extends in the ventral abdominal wall far the largest part of the aponeurosis of the m.
on each side of the linea alba from the external obliquus externus abdominis; it is the part which
surface of the thorax to the pecten ossis pubis. arises from the pars costalis of the muscle. This
The mm. obliqui and the transversus are in the flat tendon extends over the m. rectus abdominis
lateral abdominal wall. In general these muscles to the linea alba, where it unites with that of the
arise from the outer surface of the ribs, the opposite side. It extends caudally also to attach
lumbar region, or the tuber coxae to pass in the to the pecten ossis pubis. The deep trunk fascia
lateral wall to the ventral abdominal wall or to closely adheres to the aponeurosis, obscuring the
the pelvis. In the ventral wall the tendons of the direction of its fibers. The aponeurosis fuses
two oblique muscles cross the rectus muscle deeply near the mid line with the aponeurosis of
superficially, while the tendon of the transverse the m. obliquus internus abdominis and with it
muscle crosses deeply. In this way the so-called forms the external leaf of the sheath of the rec
“sheath of the rectus” is formed. The abdominal tus abdominis. It lies closely upon the superficial
muscles are covered superficially by the large surface of the m. rectus abdominis, where it is
cutaneous muscle of the trunk (m. cutaneus intimately connected with the tendinous in
trunci). scriptions of the rectus. Cranial to the pecten os
The oblique muscles, the fibers of which cross sis pubis the abdominal tendon is separated from
each other at about right angles, form the ob the pelvic tendon by the superficial inguinal
lique girdle of the abdomen. The straight and ring, the medial crus of which it forms. The deli
transverse muscles, which also cross each other cate fibers cover the caudal 2 to 3.5 cm. of the
at right angles, form the straight girdle of the m. rectus abdominis. At the level of the superfi
abdomen. cial inguinal ring, the external leaf of the rectus
The m. obliquus externus abdominis (Figs. 3 - sheath is rather sharply defined by a fibrous
31, 3-3 5 , 3-37) is an expansive sheet covering band. Strong fibers come from the opposite side
the ventral half of the lateral thoracic wall and and, after crossing the mid line, stream upward
the lateral part of the abdominal wall; in large in the direction of the tendon of origin of the m.
dogs the thoracic portion is 2 to 3 mm. and the pectineus. This tough strand becomes fixed, and
abdominal portion 4 to 7 cm. thick. According with it the outer leaf of the rectus sheath, m.
to its origin, the muscle is divided into two parts. pectineus, and prepubic tendon pass to the ilio
The pars costalis arises by indistinct serrations pectineal eminence. This, in man, is called the
in a caudally rising line from the middle parts of reflected ligam ent (ligamentum reflexum). At
the fourth or fifth to the twelfth rib, and the ad the same place, the tendinous fibers of the lat
jacent deep trunk fascia which covers the exter eral crus of the superficial inguinal ring attach to
nal intercostal muscles. It is partly covered by form the strong caudal commissure.
the ventral edge of the m. latissimus dorsi at its The pelvic aponeurosis, or tendon, arises es
origin. The unserrated pars lumbalis arises from sentially from the lumbar part of the muscle;
the last rib and, in common with the pars cos however, the serration arising from the twelfth
talis of the obliquus internus abdominis, from rib also takes part. Like the abdominal tendon,
the principal lamina of the thoracolumbar fascia. it is intimately fused with the deep trunk fascia
The cranial serrations of the muscle extend and ventrally is not separated from the abdomi
between the terminal serrations of the m. ser nal tendon. It extends down into the niche be
ratus ventralis and cover the terminal tendon of tween the abdominal wall and the femur. It
the longest part of the m. scalenus. The caudal crosses the contents of the inguinal canal and
serrations are higher on the costal wall than the forms its lateral wall. The free dorsal border of
cranial ones; thus the line of origin of the lum the pelvic tendon courses along the femoral ves
bar portion meets the lateral border of the m. sels which come through the femoral ring to en
iliocostalis. ter the femoral triangle; ventrally the pelvic
The fibers of the external oblique muscle run tendon is separated from the abdominal tendon
caudoventrally, the caudal part being more hori by the sharp-edged sagittal slit, the superficial in
zontal than the cranial. In the ventral abdominal guinal ring. The lateral crus of the pelvic tendon
M u sc les of the T run k 183
meets the medial crus of the abdominal tendon arising from the lumbar region is divided accord
to form the cranial and caudal commissures of ing to its terminal insertion into a costal and an
the superficial inguinal ring. Rarely, a small abdominal part; the portion coming from the in
heterotopic bone may be present in the aponeu guinal ligament represents the inguinal part.
rosis caudal to the medial crus where the m. pec- The strong cranial part, pars costalis, proxi-
tineus arises (Baumeier 1908). mally, is often separated from the middle part
The inguinal ligament (ligamentum ingui by a distinct fissure (containing vessels of the
nale) comes from the ilium and runs over the lat abdominal wall); its fleshy ending is on the thir
eral surface of the m. iliopsoas to blend with the teenth rib and on the cartilage of the twelfth rib.
lateral part of the prepubic tendon. It serves as The middle portion, pars abdom inalis, gives
the origin for the principal part of the pars in- rise to a broad aponeurosis at the lateral border
guinalis of the m. obliquus internus abdominis. of the m. rectus abdominis. This line of transition
The prepubic tendon is a tough tendinous is often irregular; it extends from the bend of the
mass which extends from the iliopectineal emi twelfth costal cartilage to the iliopectineal emi
nence and the tendon of origin of the m. pectin- nence. This, together with the abdominal ten
eus to the same structures of the opposite side. don of the m. obliquus externus abdominis, ex
It is firmly attached to the median pubic tuber tends over the outer surface of the rectus as part
cle situated on the external surface of the sym of the superficial leaf of the rectus sheath. It
physis caudal to the free edge. The paired por ends on the linea alba. A narrow cranial lamina
tions of the tendon have a slightly caudomedial of the aponeurosis is split off from the princi
course. The m. rectus abdominis radiates into pal portion and runs over the inner surface of
the prepubic tendon. On either side, the m. ad the m. rectus abdominis to aid in forming the
ductor longus arises close by. deep leaf of the rectus sheath.
A m. obliquus abdominis externus profundus According to Kassianenko (1928), this part of
jis not rare. It is a plate of muscle consisting of the muscle becomes amplified at its cranial bor
two, exceptionally three, deep serrations be der (in 30 per cent of dogs) by one to three
neath the principal muscle which gives rise to slender muscle bundles (pars costoabdominalis),
the pelvic tendon (Baum and Zietzschmann which arise from the medial surface of the thir
1936). teenth, twelfth, and eleventh costal angles; their
Action: Along with other abdominal muscles, tendons are related to that portion of the tendon
compression of the abdominal viscera. This of the internal abdominal oblique muscle which
action, known as abdominal press, aids in helps make up the deep leaf of the rectus sheath.
such vital functions as expiration, urination, The caudal portion of the m. obliquus inter
defecation, and parturition. Flexion of the nus abdominis, pars inguinalis, is rather dis
vertebral column when fellow muscles con tinctly separated from the middle portion by a
tract. Lateral bending of the vertebral col fissure (containing vessels of the abdominal
umn. wall). This is the part of the muscle which comes
Innervation: Lateral ventral branches of the from the tuber coxae, by means of a short apo
last 8 or 9 nn. thoracales and the lateral neurosis, and from the inguinal ligament. The
branches of the nn. iliohypogastricus and m. obliquus internus abdominis extends beyond
ilioinguinalis. the caudal border of the m. obliquus externus
The m. obliquus internus abdominis (Figs. 3 - abdominis. It sends its outermost fibers in the re
31,3-35, 3-37) is a flat muscle lying medial to gion of the inguinal canal caudoventrally toward
m. obliquus externus abdominis in the lateral ab the linea alba.
dominal wall, where it is almost completely cov Action: Compression and support of the ab
ered by the external oblique. In large dogs it is dominal viscera.
4 to 6 mm. thick. Its fibers arise from the princi Innervation: Medial branches of the last few
pal lamina of the thoracolumbar fascia caudal to nn. thoracales and the nn. iliohypogastricus
the last rib, in common with the lumbar portion and ilioinguinalis.
of the m. obliquus externus abdominis. It origi The m. transversus abdominis (Figs. 3 -3 3 ,3 -
nates mainly from the tuber coxae. Some fibers 36, 3-37) is the deepest abdominal muscle and,
arise also from the fascia covering the m. ilio like the oblique muscles, it is developed into an
psoas and the inguinal ligament. Its fibers in extensive leaf which reaches a thickness of 2 to
general run cranioventrally and thereby cross 4 mm. in large dogs. It lies in the lateral and
those of the external oblique muscle at approxi ventral abdominal wall on the internal surface
mately a right angle. The portion of the muscle of the m. obliquus internus abdominis and adja-
184 Chapter 3. M yo lo g y
-R e c tu s a b d o m in is
J n te rc o s to lis e x t e r n u s -
-O b liq u u s e x te r n u s
a b d o m i n is
- U m b ih c u s
O b liq u u s i n t e r n u s
a b d o m in is
L in e a a !b o
C u t edge o f a p o n e u r o s is
o f Obliq. e xte rn u s a b d
S o r to n u s -
Ext. c r e m a s t e r -
P re p u b iC te n d o n ------- P e c f m e u s
S p e r m o t i c cc rd
-A d d u c to r
G ra c ilis
F ig. 3-35. Superficial muscles of trunk, ventral aspect. (M pectoralis profundus removed.)
M u s c jl e s of the T run k 185
- - R e c tu s a b d o m in is
Intercostal 16 e x t e r n u s -
In tercost a I is m t e r n u s -
-T e n d in o u s i n s c r ip ti o n
T r a ns v e r s u s o b d o m m i s
Cut edcje o f
Rectus abdominis
C u t edge o f-
Obliquus m t e r n u s abd.
~ S p e rm a tic co rd
~ -P r e p u b ic tendon
Adductor -
G racilis- -
cent costal cartilages; it arises from the eighth Action and Innervation: Same as for the inter
costal cartilage, last lumbar transverse process, nal abdominal oblique.
and the tuber coxae. It is divided into a lumbar The fascia transversalis covers the inner sur
and a costal part. faces of the mm. transversi abdominis. It runs
The pars lum balis arises by broad, short ten between the iliac fascia on the ventral, lateral
dons from the transverse processes of all the border of the m. iliopsoas and the ventral mid
lumbar vertebrae and the deepest division of the line of the abdomen; during its course it covers
thoracolumbar fascia. This fascia completely the pelvic part of the m. rectus abdominis, which
surrounds the m. iliocostalis. Out of the dorsal is free of the end aponeurosis of the transversus
parts of this fascial leaf arises the m. obliquus in abdominis. Farther cranially, it fuses with the
ternus abdominis which radiates into the lateral deep leaf of the rectus sheath. In the lateral ab
abdominal wall. dominal wall it runs cranially to the diaphragm
The pars costalis, not divided from the lumbar and continues on the abdominal surface of the
part, arises muscularly on the medial sides of the latter, which it completely covers. The fascia
thirteenth and twelfth ribs and the eleventh transversalis may contain much fat. The fascia
to eighth costal cartilages in such a way that its contains strong reinforcements of coarse elastic
line of origin crosses that of the diaphragm. fibers which run in anastomosing strands from
From one to three serrations have pleural behind forward, thus crossing the course of the
coverings. The entire muscle extends ventrally fibers of the m. transversus abdominis. These in
and slightly caudally from the internal surface filtrations come from the entire length of the m.
of the thorax, 3 or 4 cm. cranial to the origin iliopsoas, and are especially strong ventrally. Be
of the m. obliquus internus abdominis. The neath the point of separation of the m. cremaster
medial branches of the ventral divisions of the externus from the caudal border of the m. ob
last few thoracic and the first few lumbar nerves liquus internus abdominis, the elastic masses
run over the superficial surface of the m. trans are the thickest; toward the ribs they become
versus abdominis. The muscle is marked by these correspondingly thinner. The peritoneum and
into several (usually six) “segments” which oc the transversalis fascia evert to form the
cur in the part behind the last rib, the remainder processus vaginalis just caudal to the caudal
appearing medial to the costal arch. The muscle border of the m. rectus abdominis. The m. cre
extends on the inner surface of the m. rectus ab master externus is located on its lateral and
dominis and beyond its dorsal border before giv caudal sides.
ing rise to its end aponeurosis. This forms a lat The inguinal ligament (lig. inguinale) (Fig. 3 -
erally convex line, the summit of which lies at 36) is closely related to the fascia transversalis
the region of the umbilicus, 5 cm. from the mid and, like it, contains much elastic tissue. In com
line; toward the xiphoid cartilage it lies only 1.5 parison with that in other domestic animals it is,
cm. from the mid line. The end aponeurosis in the dog, a relatively incomplete structure and
forms most of the inner leaf of the sheath of the independent of the external oblique. It is a
rectus abdominis. It unites inseparably at the strong band extending in the iliac fascia from
linea alba with the external leaf. In the costal re the tuber coxae obliquely over the m. iliopsoas,
gion, it unites only with part of the end aponeu marking the caudal border of origin of the fascia
rosis of the m. obliquus internus abdominis. transversalis. Together with this fascia it extends
The cranial part of the muscle, by the devel ventrolaterally along the m. iliopsoas to radiate
opment of incomplete fissures, encroaches di into the transversalis fascia which covers the vag
rectly upon the m. transversus thoracis, and the inal process. The main part of the inguinal lig
end aponeurosis covers the outer surface of the ament continues distally between the internal
free end of the xiphoid cartilage. The caudal part inguinal and femoral rings to attach to the lateral
becomes aponeurotic in the region of the last border of the prepubic tendon. By taking this
two to four “segments” near the lateral edge of course it forms the caudal border of the internal
the m. rectus abdominis. This does not cross the inguinal ring. At its ilial end, it gives origin to
deep surface of the rectus abdominis muscle; in part of the m. obliquus internus abdominis. By
stead it traverses the outer surface to take part fusing with the deep trunk, iliac, pelvic, and
in the formation of the external leaf of the rectus transversalis fasciae, it acts as a binder in clos
sheath. It fuses toward the pelvis with a tendi ing the potential space which might exist be
nous strand of the rectus. On the deep surface of tween the pelvic and abdominal walls.
the pelvic end of the rectus there is no aponeu The m. rectus abdominis (Figs. 3 -35, 3-36,
rotic covering; there is only a thin continuation 3-37) is a long, rather wide, flat muscle which
of the fascia transversalis and peritoneum. extends, one on each side of the linea alba on
187
M u sc les of th e T runk
Rectus a b d o m i n i s
Fat
E x t a b d . o b liy u a
In t. a b d . o b i.
--T r a n s . a b d ■
—p g p i f o n & u m
Ext. a b d ■ obi-
--Int. a b d . o b i ■
____ T r a n sabd■
the thoracic and abdominal walls between the of the wide and long aponeuroses of the m. ob
external and internal leaves of the rectus sheath, liquus externus abdominis, most of the aponeu
and runs from the first costal cartilage to the rosis of the m. obliquus internus abdominis, and,
pecten ossis pubis. Cranially, in large dogs, it is near its caudal end, a portion of the aponeurosis
7 to 8 cm. broad; caudally, it gradually narrows of the m. transversus abdominis. The internal
to 3.5 to 4 cm. Its thickness is 5 to 7 mm., de leaf of the rectus sheath is formed by the end
creasing toward the lateral border. The fibers aponeurosis of the m. transversus abdominis, the
of the muscle course longitudinally. It arises by fascia transversalis, and, cranially, by an internal
a broad, flat tendon from the sternum and the leaf of the aponeurosis of the m. obliquus inter
first costal cartilage and rib, where it is covered nus abdominis. At its pelvic end, the m. rectus
by the terminal tendon of the m. transversus abdominis lacks an internal aponeurotic cover
costarum. It also has a fleshy origin by means ing, being covered here by only a thin continua
of a special serration from the sternal portion tion of the transversalis fascia and peritoneum.
of the ninth costal cartilage. As it passes over The inguinal canal (canalis inguinalis) (Fig. 3-
the ventral abdominal wall, it lies in a nearly 36), in both sexes, is a connective tissue-filled
horizontal position, with the medial border fac fissure between the abdominal muscles and their
ing the linea alba. Occasionally the terminal por aponeuroses. In the male the inguinal canal
tion of the muscle is wide enough to help in the serves as the passageway for the processus vagi
formation of the medial wall of the inguinal nalis with the m. cremaster externus and the
canal and to appear at the level of the superficial spermatic cord; in the female it may contain the
inguinal ring. United by the linea alba and cov vaginal process with the round ligament and
ered externally by a strong tendinous covering, much fat. It is relatively short. It begins at the
the two recti end on the pecten ossis pubis, from deep inguinal ring, which is formed by (1) the
one iliopectineal eminence to the other. At its ventral end of the inguinal ligament, (2) the cau
insertion each muscle unites with the tendon dal border of the m. obliquus internus abdomi
of origin of the m. pectineus and the prepubic nis, and (3) the lateral border of the m. rectus
tendon. A conical, paired segment of superficial abdominis. In large dogs the lower angle of this
fibers, however, continues farther and ends on ring is 3 cm. lateral to the mid line, 2 cm. cra
the tubercle on the ventral surface of the pelvic nial to the origin of the m. pectineus, and 2 to
symphysis. This crosses the thickened border of 3 cm. caudal to the caudal border of the m.
the external leaf of the rectus sheath. This long transversus abdominis. The deep inguinal ring is
muscle is divided into segments by three to six covered externally by the aponeurosis of the m.
(usually five) transverse, zigzag tendinous inter obliquus externus abdominis. The path of the
sections. Their distinctness varies. Their number canal is determined by the processus vaginalis.
does not correspond with the number of enter Since the latter pushes over the caudal border of
ing nerves. Intimately attached to the tendinous the m. obliquus internus abdominis for a short
intersections are fibers of the external leaf of the distance, the medial wall of the inguinal canal is
rectus sheath. The fibers of the internal leaf of formed by the superficial surface of this muscle.
the sheath are not as firmly attached. The first The superficial surfaces of the aponeuroses of
intersection is at the level of the seventh costal the mm. transversus abdominis and rectus ab
cartilage; the last segment is usually the longest; dominis also aid in forming the medial wall. The
all other relations vary (Strauss 1927). lateral wall is formed solely by the aponeurosis
Action: All functions which are dependent of the external oblique. The canal is open to the
upon abdominal press, such as expiration, outside because the abdominal and pelvic parts
urination, defecation, and parturition; sup of the aponeurosis of the m. obliquus externus
port of the abdominal viscera; to bring the abdominis separate and then come together in
pelvis forward; flexion of the back. the slitlike, superficial inguinal ring (annulus in
Innervation: Medial branches of the ventral guinalis superficialis). In this way the pelvic ten
branches of the nn. thoracales and medial don forms the lateral border, crus laterale, and
branches of the nn. iliohypogastricus and the abdominal tendon forms the medial border,
ilioinguinalis. crus m ediate, of the more or less sagittal slit.
The sheath of the rectus abdominis (Figs. 3 - Where the borders meet, the cranial and caudal
35, 3-37) covers both surfaces of the rectus ab angles, or commissures, are formed. The caudal
dominis muscle. It is formed primarily by the commissure is strong, as it is backed by the ten
aponeuroses of the other abdominal muscles. don of origin of the m. pectineus. The cranial
The external leaf of the rectus sheath consists commissure is much weaker, as the parallel
M u sc les of th e T a il 189
strands of collagenous tissue which form the ab its free edge, where it ends in the superficial tho
dominal and pelvic parts of the aponeurosis of racic fascia. The fibers of the ventral border
the external oblique are held together mainly coming from the flank reach each other in the
by the deep trunk fascia. Unlike in man, there mid-ventral line caudal to the sternum. The gap
is a minimum of cross-over of fibers of the apo thus existing between the muscles of the oppo
neurosis at the cranial angle. site sides is filled in by a division of the abdomi
The linea alba (Fig. 3-35) is a mid-ventral nal cutaneous muscle, right and left muscles
strip of collagenous tissue which extends from spread out to the prepuce as the m. preputialis
the xiphoid process to the symphysis pelvis. It in the male, and to the mammary glands as the
serves for the main insertion of the abdominal m. supramammaricus in the female.
transverse and external and internal oblique The m. preputialis, filling the space between
muscles. The medial borders of the right and the opposite abdominal cutaneous muscles in
left rectus muscles lie closely against its lateral the region of the xiphoid cartilage in the male, is
borders. At the level of a transverse plane an unpaired longitudinal strand. Toward the
through the last ribs, the linea alba contains a umbilicus a pair of muscular strands arise from
scar, the umbilicus, a remnant of the umbilical the m. preputialis. They radiate into the prepuce
ring and cord. The linea alba, just caudal to the in such a way that they come together archlike in
xiphoid process, is a little over 1 cm. wide and the prepuce ventral to the glans. In so doing they
less than 1 mm. thick. It gradually narrows and are firmly united with each other and with the
thickens caudally. Caudal to the umbilicus it ap external preputial leaf.
pears as a line, being less than 1 mm. wide but The m. supramammaricus of the bitch is ho
considerably thicker. It blends with the prepu- mologous with the m. preputialis of the male. In
bic tendon and attaches to the cranial edge of contrast to the muscle in the male, this muscle is
the pelvic symphysis. more delicate and narrower, and is paired from
The m. cutaneus trunci (Figs. 3-45,3-52), its beginning. From the region caudal to the xiph
according to Langworthy (1924), is a derivative oid cartilage they extend caudally in loose
of the m. pectoralis profundus. As a thin leaf it bundles, over the mammary gland complex, to
covers almost the entire dorsal, lateral, and ven the pubic region. Cranial to the paired inguinal
tral walls of the thorax and abdomen. It begins mammary glands, each blends with the ipsilat-
caudally in the gluteal region and, running for eral m. cutaneus trunci.
ward and downward, covers the dorsal and lat Action: The m. cutaneus trunci shakes the skin
eral surfaces of the abdomen and thorax. It ends to help rid the animal of external foreign
in the axilla and on the caudal border of the deep bodies. It pulls the prepuce cranially over
pectoral. It lies in the superficial trunk fascia and the unsheathed glans penis.
is not attached to the vertebral spines. It is prin Innervation: Cutaneous branches of the ven
cipally a longitudinal muscle; its origin is in the tral branches of nn. cervicalis 8 and tho-
superficial gluteal fascia. The dorsal borders of racalis 1 (Langworthy 1924).
the muscle on each side run parallel along the
spines of the lumbar and thoracic vertebrae.
Only in the region behind the scapula, where
the muscle begins to extend downward on the M U SC LES OF TH E TAIL
thorax, do the fibers arise from the mid line and
meet those of the opposite side. Since this part The coccygeal vertebrae are largely enclosed
of the muscle is also not attached to the spines of in muscles. The mm. sacrococcygeus dorsalis lat
the vertebrae, it is free over the vertebral col eralis and medialis, dorsal in location, are exten
umn to be included in raised folds of the skin. Its sors or levators of the tail. The mm. sacrococcyg
ventral border crosses, in the fold of the flank, to eus ventralis lateralis and medialis, ventral in
the lateral and ventral abdominal wall. The location, are flexors or depressors of the tail. The
course of the fibers is slightly ventrocranial. Its mm. coccygeus, levator ani, and the inter-
craniodorsal border covers the m. trapezius, a transversarius caudae, lateral in location, are
portion of the m. infraspinatus, and the m. latis- the lateral flexors of the tail. The dorsal muscles
simus dorsi, and ends by means of the muscular are direct continuations of the epaxial muscula
axillary arch in the medial brachial fascia. The ture of the trunk. The coccygeal muscles lie on
principal part of the muscle, however, with its the lumbar vertebrae, sacrum, and coccygeal
loose fiber bundles, passes to the superficial sur vertebrae, and insert on the coccygeal vertebrae,
face of the m. pectoralis profundus adjacent to exclusively. They have fleshy endings as well as
190 Chapter 3. M yology
tendinous ones of variable length. The most cau dividual muscles run between the spines of cra
dal tendons go to the last coccygeal vertebrae. nial vertebrae and the dorsolaterally located tu
Proximally the muscles, as well as the vertebral bercles, as well as on the mammillary processes
bodies, are larger. The coccygeal muscles of the on the cranial ends of more caudal coccygeal
dog resemble those of the cat (Schumacher vertebrae. Toward the tip of the tail the muscle
1910). segments become shorter, smaller, and more ho
The m. sacrococcygeus dorsalis lateralis, or mogeneous. They arise from the small processes
long levator of the tail (Fig. 3-38), is a flat, seg which are dorsolateral to the caudal edge of the
mental muscle strand becoming stronger toward rodlike coccygeal vertebrae. They pass over only
its dorsal border, 2.5 to 3 cm. high in the lumbo one segment and end on dorsolateral humps
sacral region in large dogs and 1 to 1.5 cm. thick which correspond to the mammillary processes
at its free edge. It may be regarded as a continu of the lumbar vertebrae. The superficial tendons
ation of the m. longissimus on the tail. In the end in common with the long tendons of the m.
caudal part of the lumbar region it lies between sacrococcygeus dorsalis lateralis. Muscle fibers
the m. longissimus, laterally, and the mm. mul- also accompany the tendons.
tifidus lumborum and sacrococcygeus dorsalis Action: Extension of the tail, possibly also lat
medialis, medially. It is covered by the thick eral flexion.
coccygeal fascia. Out on the tail, it extends be Innervation: Branches of the truncus coccyg-
tween the mm. intertransversarius dorsalis and eus dorsalis.
sacrococcygeus dorsalis medialis. It has a fleshy The m. sacrococcygeus ventralis lateralis, or
origin from the aponeurosis of the m. longissimus long depressor of the tail (Fig. 3-39), is strong;
and a tendinous origin from the mammillary in large dogs, at the sacrum, it is 2.25 cm. high
processes of the first to sixth lumbar vertebrae, and 0.75 cm. thick. It consists of numerous long,
the articular processes of the sacrum, and the individual parts which are arranged like those of
mammillary processes of at least the first eight the long levator and which end by means of long
coccygeal vertebrae. It is indistinctly divided into tendons from the sixth to the last segment. The
long individual parts which partly cover one an first segment comes from the ventral surface of
other. From this muscular belly which extends the body of the last lumbar vertebra and from
from the second sacral to the fourteenth coccyg the sacrum; the remainder arise from the ven
eal vertebra (when 20 coccygeal segments are tral surfaces and the roots of the transverse proc
present), there appear 16 thin, long tendons. esses of the coccygeal vertebrae. From the seg
These are arranged into a flat bundle by the ac mented bellies of the third and successive seg
cumulation of successive tendons. They lie em ments caudally, the individual long tendons
bedded in the thick, deep coccygeal fascia. The arise and are embedded in the thick, deep coc
first tendon ends on the mammillary process of cygeal fascia. The first of these is attached to the
the fifth coccygeal vertebra, the next ends on the ventrolateral tubercle (processus hemalis) of
sixth, and so on, to the last one. Cranial to their the proximal end of the sixth coccygeal verte
terminations a few take on a little tendon of the bra, the second on the corresponding elevation
underlying segment of the m. sacrococcygeus of the seventh, and so on to the last coccygeal
dorsalis medialis. vertebra. Before inserting, each of these tendons
Action: Extension or lifting of the tail, possibly acquires the little tendon of the segment of the
also to move it to one side. short depressor which has been crossed by the
Innervation: Branches of the truncus coccyg- segment of the long depressor.
eus dorsalis. Action: Flexion of the tail and, occasionally,
The m. sacrococcygeus dorsalis medialis, or lateral movement.
short levator of the tail (Fig. 3-38), is the direct Innervation: Branches of the truncus coecyg-
continuation on the tail of the m. multifidus and, eus ventralis.
like the latter, it is composed of relatively short, The m. sacrococcygeus ventralis medialis, or
individual segments. It lies next to the median short depressor of the tail (Fig. 3-39), consists of
plane on the sacrum and coccygeal vertebrae segmental, short individual parts extending from
and extends from the seventh lumbar to the last the last sacral vertebra throughout the length of
coccygeal vertebra. The individual segments can the tail. It lies against the ventral surface of the
be isolated at the root of the tail. They are com vertebrae and, with the muscle of the opposite
posed of deep, short muscle masses and a strong, side, forms a deep furrow (for the a. coccygea).
superficial, long part which possesses a little ten At the pelvic outlet the bundles are very strong
don that spans four or five vertebrae. These in and the segmentation is indistinct. Soon, how
M u sc les of the T a il 191
ever, independent segments are separated out. by a strong intermuscular septum of the coccyg
The fibers of each of these segments arise essen eal fascia.
tially from the ventral surface of one vertebra. Action and Innervation: Same as for the m.
Superficially, a small flat tendon is then formed. intertransversarius dorsalis coccygeus
This unites with the tendon of the long depres (supra).
sor which lies immediately lateral to it, and this The pelvic diaphragm (diaphragma pelvis) in
common tendon then passes over the following quadrupedal mammals is the vertical closure of
segment to end on the hemal process of the next the pelvic cavity through which the rectum
following vertebra. passes. The two muscles of the pelvic diaphragm
Action and Innervation: Same as for the m. are the m. coccygeus (m. coccygeus lateralis)
sacrococcygeus ventralis lateralis (supra). and the m. levator ani (m. coccygeus medialis).
The m. intertransversarius dorsalis coccyg- The m. coccygeus (m. coccygeus lateralis)
eus (Figs. 3-38, A; 3-40, B) lies between the sa (Figs. 3-40, 3-71, 3-86) is a strong muscle aris
crum and the middle of the tail. In general, it ing by means of a narrow tendon on the ischiatic
consists of short, individual parts, of which only spine cranial to the internal obturator muscle. It
the first is well developed. This portion arises on crosses the sacrotuberous ligament medially and,
the long, dorsal sacroiliac ligament, on the lateral spreading like a fan, extends to the lateral sur
part of the third sacral vertebra, and forms a face of the tail. There it ends, between the mm.
large, round muscle belly which ends on the intertransversarii, on the transverse processes of
transverse process of the fifth or sixth coccygeal the second to fifth coccygeal vertebrae. It is par
vertebra by means of a long tendon. In its course tially covered by the caudal branch of the m.
it receives supplementary fibers from the trans gluteus superficialis. In large dogs it is 2.5 to 3.5
verse processes of the first few coccygeal verte cm. wide and 5 to 6 mm. thick.
brae. These deep elements gradually become in Action: Bilateral: to press the tail against the
dependent muscles which extend from one anus and genital parts and, in conjunction
transverse process to that of the following verte with the depressors, to draw the tail be
bra. They lie on the dorsal surfaces of the trans tween the rear legs. Unilateral: lateral flex
verse processes or their rudiments, where they ion.
are partly covered by the long tendons of the Innervation: Ventral branches of the third
levators. These muscle segments become so sacral nerve.
small in the caudal half of the tail that they can The m. levator ani, also known as the m. coc
hardly be isolated. Superficial parts of the first cygeus medialis or the m. ilioischiopubococcyg-
large segment give rise to two or three long, flat eus (Figs. 3-40, 3-71), lies largely cranial to the
tendons which extend to the thick coccygeal coccygeus. It is a broad, flat, triangular muscle
fascia and to the rudiment of the transverse originating on the medial edge of the shaft of the
process of the sixth or seventh or even the ilium, on the inner surface of the ramus of the
eighth coccygeal vertebra. This is the m. ab pubis, and on the entire pelvic symphysis. Bi
ductor caudae dorsalis, or m. coccygeus acces laterally, the muscles spread out and radiate up
sorius. ward toward the root of the tail. In so doing,
Action: With the m. intertransversarius they surround a large median, fatty mass, as well
ventralis coccygeus, lateral flexion of the as the genitalia and the rectum. Caudally, each
tail. encroaches upon the inner surface of the m.
Innervation: Branches of the truncus coccyg obturator internus. After decreasing in size, the
eus ventralis. muscle then appears at the caudal edge of the m.
The m. intertransversarius ventralis coccyg coccygeus, passes into the coccygeal fascia, and
eus (Fig. 3-40, B), situated ventral to the trans ends on the hemal process of the seventh coc
verse processes, begins at the third coccygeal cygeal vertebra by means of a strong tendon
vertebra. It forms a round belly, composed of immediately next to the tendon of its fellow of
segments, and, at the base of the tail, is smaller the opposite side. This muscle can be divided
than the dorsal muscle; however, it has a more into an ischial part, m. ischiococcygeus, and an
constant size and is well segmented, and thus is iliopubic part, m. iliopubococcygeus, between
easily traced to the end of the tail. Ventrally the which the n. obturatorius passes. The fibers of
muscle is covered by the long tendons of the both parts enter the tendon at an angle. The
long depressor of the tail. From the third to the deep surface of the muscle is firmly covered by
fifth coccygeal vertebra the ventral and dorsal the pelvic fascia, which is also connected with
mm. intertransversarii are separated by the m. the m. sphincter ani externus. Pettit (1962) has
coccygeus lateralis; otherwise they are separated summarized many cases of perineal hernia in the
Chapter 3. M yo lo g y
- Zna l u m b a r v e r te br a
- N ul t i f i d u S
lumborum
- L o n g is s im u s
d orsi
-S acrococcygeus
d o rs a lis la t e r a lis
- 7 th l u m b a r ve rte b ra ,
ySpinOuS p ro c e s s
S a c ro c o c c y g e u s
d o r s a l is m edial is
/ n te rtro n s v e rs a riu s
d o r s a l i s coccygeus
Sacrococcygeus A K "1 S a c r o c o c c y g e u s
d o rs o /is m e d ia /is-T - d o r s a lis la te ro lis
o r ig in s
•i n s e r f i o ns
Coccygeus
Socrococcygeus
v e n t r a lis m e d ia lis
F ig u r e 3-40A
F ig u re 3-39
iT r a n s v e r s e p r o c e s s
1 o f sa c ru m
7 th caccyg.
v e rte b ra
In ie r tr o n s v e r s a r iu s
v e n tra lis coccygeus
G lu te u s m e d iu s '
C accyg eus
G lu te u s s u p e r f i c i a li s ■
- L e v a to r a n i
G lu te u s p r o f u n d u s
'S a c r o t u b e r o u s l i g a m e n t
F ig u re 3-40B
Fic. 3-39. Sacrococcygeal muscles, ventral aspect.
F ig . 3-40. Muscles of the pelvis.
A. Mm. levator ani and coccygeus, ventral aspect.
B. Coccygeal and gluteal muscles, lateral aspect.
194 Chapter 3. M yology
dog and described their surgical repair in regard sphincter encircle the anus ventrally. The re
to the muscles of the pelvic diaphragm. maining superficial ventral fibers end on the
Action: Bilateral: to press the tail against the urethral muscle and the bulbocavernosus muscle
anus and genital parts; unilateral: to bring of the male. In the female comparable fibers
the tail cranially and laterally. The mm. blend with the constrictor vulvae.
levatores ani, in combination with the The m. coccygeoanalis (Fig. 3-41) is homol
levators of the tail, cause the sharp angula ogous with the caudo-anal and caudo-caverno-
tion between the sixth and seventh coccyg sus muscles described in the cat by Straus-
eal vertebrae which is characteristic for def Durckheim (1845), and in the dog by Langley
ecation; compression of the rectum. and Anderson (1895). It arises as a band of
Innervation: Ventral branches of the third smooth muscle fibers about 3 mm. wide and less
(last) sacral and the first coccygeal nerve. than 1 mm. thick on each side of the sacrum or
The m. rectococcygeus (Fig. 3-41) is a paired the first coccygeal vertebra. At their origins
smooth muscle composed of fibers from the ex there is a considerable decussation of fibers ven
ternal longitudinal musculature of the rectum. tral to the rectococcygeus muscle. Each band
The fibers sweep caudodorsally from the sides of passes ventrocaudally across the lateral surface
the rectum and pass through the fascial arch of the rectum, to which it contributes some
formed by the attachment of the external anal fibers. It becomes wider distally as it passes be
sphincter to the fascia of the tail. Right and left hind the anal sac and into the sphincters. The
portions of the muscle fuse beneath the third bulk of its fibers appear to end near the duct of
coccygeal vertebra. The median muscle thus the anal sac, although some fibers insert in the
formed lies between the ventral sacrococcygeal external sphincter. Occasionally, a rudiment of a
muscles and passes caudally to insert on the fifth ventral anal loop may be present. A ventral por
and sixth coccygeal vertebrae. The attachment tion of the muscle band, in combination with
of the rectococcygeus muscle on the tail serves some fibers from the external sphincter, con
to anchor the rectum and provide for caudal tinues distally as the retractor penis muscle.
traction in defecation. Extension of the tail dur Superficially the m. coccygeoanalis is covered by
ing defecation aids in evacuating the rectum be the levator ani, with which there may be some
cause of the attachments of the mm. rectococ fiber interchange.
cygeus, coccygeus, and levator ani. The mm.
coccygeus and levator ani cross the rectum FASCIAE OF THE TRUNK AND TAIL
laterally and tend to compress it; the m. recto
coccygeus, by shortening the rectum, aids in On the trunk, as on other parts of the body,
evacuation of the fecal column. there is a superficial and a deep fascia, known
Action: To aid in defecation. collectively as the external fascia of the trunk. It
Innervation: Autonomic fibers from pelvic covers the muscles and bones of the thorax and
plexus. abdomen. In addition, there is an internal fascia
The m. sphincter ani internus is the caudal of the trunk, which serves a special function in
thickened portion of the circular coat of the anal the formation of the body cavities.
canal. It is composed of smooth muscle fibers The internal fascia of the trunk lies on the
and is smaller than the striated external anal deep surfaces of the muscles of the body wall
sphincter. Between the two sphincter muscles, and on the superficial surfaces of the serous
on either side, lies the anal sac. The duct from coverings of the cavities. In the thoracic cavity,
the anal sac crosses the caudal border of the it is the fa s c ia endothoracica; in the abdominal
internal sphincter muscle. cavity, the fascia transversalis. The latter covers
The m. sphincter ani externus (Fig. 3-41), the m. transversus abdominis on its deep surface
composed of striated muscle fibers, surrounds and fuses ventrally with its aponeurosis. Crani
the anus, covers the internal sphincter except ally the fascia transversalis covers the diaphragm
caudally, and is largely subcutaneous. The as a thin membrane. The internal trunk fascia is
cranial border of the external sphincter is united reinforced by yellow elastic tissue wherever it
by fascia to the caudal border of the levator ani. covers a movable or expansible structure, such
Dorsally the external sphincter attaches mainly as the diaphragm. The fa s c ia iliaca covers the
to the coccygeal fascia at the level of the third deep lumbar muscles and is connected with
coccygeal vertebra. This attachment is such that the last few lumbar vertebral bodies and with
a cranially directed concave fascial arch is the ilium. The fascia pelvina clothes the pelvic
formed, through which the rectococcygeus mus cavity; it lies deeply on the bones and gluteal
cle passes. About half of the fibers of the external muscles concerned and it continues on the pelvic
F a s c ia e of the T runk and T a il 195
M. c o c c y g e u s
M. l e v a t o r a n i M. r e c to c o c c y g e u s
M. co ccyge oa na lis
M. s p h i n c t e r a n i externus
- (re fle c te d )
_ - M . s p h in c t e r a n i i n t e r n u s
---- A n a l sac
R e c tu m
- M. r e t r a c t o r pen is
M. I e v o t o r a n i "
( c u t p* r e f l e c t e d ) ■ - M. b u lb o c a v e r n o s u s
E p a x ia l m u s c u la tu re II Lumbar vertebrc
Left Right
Aorta Postcava
F ig . 3-42. Schematic plan of cross section through lumbar region, showing areas of fat deposition.
196 Chapter 3. M yo lo g y
surface of the muscles of the pelvic diaphragm. after completely surrounding the lumbar part of
In obese dogs it contains much fat. the laterally projecting m. iliocostalis, it termi
The superficial external fascia of the trunk nates on the free ends of the lumbar transverse
(fascia trunci superficialis) is relatively thick; it processes where the pars lumbalis of the m.
covers the thorax and abdomen in a manner transversus abdominis originates. Cranially this
similar to that on other parts of the body. It ex fascia gradually disappears; on the thorax it
tends cranially, dorsally, and laterally to the attaches to the ribs lateral to the m. iliocostalis.
shoulder and neighboring parts of the brachium. The principal lamina of the thoracolumbar fascia
The ventral part uses the sternal region to gain gives rise laterally to the m. serratus dorsalis
the neck and also sends connections to the cranialis. It becomes extremely strong as the
superficial fascia of the medial surface of the superficial leaf of the fascia spinotransversalis.
thoracic limb. Caudally, direct continuations are It passes over the m. splenius, where it en
found in the superficial gluteal fascia and, by croaches upon the fascia colli profunda by means
means of the flank, on the cranial crural portions of a distinct border. In the lumbar region, the
to the lateral and medial crural fasciae, and fi deeper layer of the principal lamina of the
nally, in the pubic region, to correspondingly thoracolumbar fascia is the stronger leaf. At
superficial fascial parts. There are no attach about the level of the lateral border of the
ments with the dorsal ends of the thoracic and epaxial musculature, it gives rise to portions of
lumbar vertebrae. Thus, as on the neck, the the mm. obliquus externus abdominis and ob
fascia can be picked up with folds in the skin. liquus internus abdominis. As the relatively deli
There are ventral fascial leaves to the prepuce cate fascia trunci profunda passes to the lateral
and to the mammary glands. The superficial wall of the abdomen and thorax, it continues on
trunk fascia covers the mm. trapezius andlatis- the superficial surface of the m. obliquus ex
simus dorsi, as well as parts of the pectoral mus ternus abdominis and on the thoracic serrations
cles, omotransversarius, deltoideus, and triceps. of the m. serratus ventralis, with which it inti
In relation to the underlying structures, all parts mately fuses. Over the m. serratus ventralis
of the superficial fascia are extremely displace thoracis and under the shoulder, the deep tho
able; only on the shoulder is this mobility racic fascia is connected with the deep cervical
limited. Wherever there is great mobility, in fascia, these two fasciae meeting on the super
well-nourished animals large quantities of sub ficial surface of the mm. serratus ventralis cervi
fascial fat are deposited. cis and scalenus. In the abdominal region and in
The deep external fascia of the trunk (fascia the caudal thoracic region the deep fascia of the
trunci profunda) is a strong, shining, tendinous trunk descends over the external surface of its
membrane; it begins on the ends of the spinous pelvic and abdominal tendons and is more or less
processes of the thoracic and lumbar vertebrae, firmly united with them. Insofar as it has rela
from the supraspinous ligament. It passes over tionships with the abdominal tendon of the rec
the epaxial musculature to the lateral thoracic tus muscle, the deep abdominal fascia also takes
and abdominal wall, to fuse with the fascia of part in the formation of the superficial leaf of
the opposite side at the linea alba. In the sternal the rectus sheath. Caudal to the lumbar region,
region it passes under the pectoral musculature the deep fascia of the trunk becomes the deep
on the sternum and costal cartilages. Caudally it gluteal fascia, and from the lateral abdominal
is attached to the ilium. wall it becomes the crural fascia. In the region of
The principal leaf of the thoracolumbar fascia the superficial inguinal ring, the deep trunk
is again divided into two superimposed leaves, fascia has special significance. At the commis
and in well-nourished dogs is covered with large sures of the crura it unites the diverging collage
amounts of fat (Fig. 3-42). It passes under the nous strands (cranial commissure) and tends to
mm. latissimus thoracis, trapezius, and rhom- prevent enlargement of the ring during hernia
boideus to the medial surface of the base of the tion. At the caudal commissure it blends with the
shoulder; here, following the mm. iliocostalis tendon of origin of the m. pectineus. From the
and longissimus, it becomes the strong, eventu crura, especially the medial one, the deep fascia
ally bilaminate fascia spinotransversalis. Espe extends on the processus vaginalis and its con
cially in the lumbar region, a strong deeper leaf tents, there being known as the external sper
is separated from the thoracolumbar fascia; this matic fascia.
is the aponeurosis of the mm. longissimus and Deep beneath the shoulder, the deep fascia of
iliocostalis. From its deep surface arise numerous the trunk is represented by the fascia spino
fibers for both muscles. Moreover, it sends an transversalis, which is somewhat independent
intermuscular septum between the two muscles; from the fascia thoracolumbalis; like its principal
M u scles of the T h o r a c ic L im b 197
leaf, it consists of a superficial and a deep part, be divided into those from the trunk to the
the superficial leaf being by far the stronger. shoulder and those from the trunk to the bra
Here under the shoulder it lies medial to the chium.
mm. rhomboideus, serratus ventralis, and latis The m. trapezius is a broad, thin, triangular
simus thoracis, and lateral to the mm. semi muscle (Figs. 3-43, 3-45). It lies under the skin
spinalis, longissimus, and iliocostalis. The fascia and the cervical cutaneous muscle in the neck,
spinotransversalis is that portion of the deep and crosses the interscapular region of the
trunk fascia which arises from the supraspinous shoulder. It arises from the median fibrous raphe
ligament of the first eight to ten thoracic verte of the neck and the supraspinous ligament of the
brae. The two leaves are fused for 0.5 to 1 cm. thorax. Its origin extends from the 3rd cervical
from their origin. vertebra to the 9th thoracic vertebra. The in
The stronger superficial leaf of the fascia sertion is on the spine of the scapula. It is di
spinotransversalis is the aponeurosis of origin of vided into a cervical and a thoracic portion by a
the m. serratus dorsalis cranialis. Cranially, it has tendinous band extending dorsally from the
a distinct border; caudally, it becomes weaker spine of the scapula.
and blends with the principal leaf of the thora The fibers of the comparatively narrow m.
columbar fascia. From this the m. serratus dor trapezius cervicis arise on the mid-dorsal raphe
salis caudalis arises. The cranial segment of the of the neck. They run obliquely cau doventrally,
superficial fascial leaf seems to lie transversely to the spine of the scapula, and end on the free
over the origin of the m. splenius; however, it edge of the spine. Only a small distal portion of
also sends a narrow tendinous strand ventrally the spine remains free for the attachment of the
over the mm. longissimus and iliocostalis to end m. omotransversarius. This muscle cannot be
on the transverse processes of the first thoracic separated from the ventral border of the trape
and last cervical vertebrae. zius near the spine.
The more delicate, deep leaf of the fascia The m. trapezius thoracis arises from the
spinotransversalis extends from a transverse supraspinous ligament and the dorsal spines of
plane through the third intercostal space to the the 3rd to the 8th or 9th thoracic vertebra, and
last few thoracic vertebrae, where it goes into by an aponeurosis which blends with the lumbo-
the deep layer of the thoracolumbar fascia. Its dorsal fascia. Its fibers are directed cranioven-
fibers extend transversely over the mm. semi trally and end on the proximal third of the spine
spinalis and longissimus to end with the lateral of the scapula.
tendons of the m. longissimus on the ribs. Crani The fibrous band which divides the m. trape
ally the m. splenius arises from this leaf. zius varies considerably. Sometimes it is lacking;
The superficial and deep fasciae of the tail sometimes it is broad and includes the dorsal
(fasciae coccygeae) arise from the correspond border of the middle part of the entire muscle;
ing leaves of the gluteal fascia. The superficial is sometimes it is interrupted. When it is present,
very insignificant; the thick, deep leaf provides it serves as a common attachment for the two
thick connective tissue masses for special en- parts of the m. trapezius.
sheathment of the long tendons of the mm. Action: To elevate the limb and draw it for
sacrococcygeus dorsalis lateralis and sacrococ ward.
cygeus ventralis lateralis. Innervation: Dorsal branch of the n. acces
sorius.
The m. omotransversarius (Figs. 3-43,3-45)
MUSCLES OF THE THORACIC LIM B lies at the side of the cervical vertebrae as a flat,
narrow muscle. It arises on the distal portion of
Extrinsic Muscles the scapular spine, as far as the acromion, and
from that part of the omobrachial fascia which
The extrinsic muscles of the thoracic limb covers the acromial part of the m. deltoideus. It
originate on the neck and thorax and extend to soon separates from the m. trapezius cervicis,
the shoulder or brachium as far distally as the el dips under the m. cleidocervicalis, and proceeds
bow joint. They include a superficial layer of over the mm. scalenus and intertransversarius
muscles lying directly upon the fascia of the cervicalis, which cover the transverse processes
shoulder and brachium, and a second, deeper of the cervical vertebrae dorsally, to the caudal
layer, being in part medial and in part lateral to end of the wing of the atlas. In large dogs it is at
the shoulder and brachium. According to the first as much as 4 cm. wide and 2 to 4 mm. thick;
points of attachment, the extrinsic muscles can cranially it becomes narrower and thicker. Its
198 Chapter 3. M yo lo g y
Bi ceps~
T r a p e z iu s a n d
Deltoideus De H o i d e u s
-j-ln fn a s p in a tu s
T e r e s m i n o r and Rh o mb o i d e u s
T r i c e p s , l ong h ead
Subs c apul ar i s' Teres m ajor
F i g . 3-43. Left scapula, showing areas of muscle attachment, lateral aspect.
C u ta n e u s tru n c i
i
C le id o c e r v ic a lis —
S te rn o c e p h a lic u s - -
O m o + r a r s v e r s a r i u s ------ ---
C la v ic u la r -tendon
D e l t o i d e u s - - ~-
C leidobrachialis ~ -
E x te n s o r g ro u p
A n c o n e u s ------
ventral border is limited by the transverse proc has well-defined serrations which are covered in
esses of the cervical vertebrae. part by the m. scalenus. Its three or four caudal
Action: To draw the limb forward. serrations interdigitate with those of the m.
Innervation: N. accessorius. obliquus externus abdominis.
The m. rhomboideus (Figs. 3-24, 3-43, 3-44), Action: Support of the trunk, to carry the
covered by the trapezius, fans out on the neck trunk forward and backward; inspiration;
and on the scapular region of the back between to carry shoulder forward and backward
the median line of the neck and the base of the with respect to the limb.
scapula. It is in part flat and in part thick, and is Innervation: Cervical portion: rami ventrales
divided into two portions. of nn. cervicales; thoracic portion: n. tho
The stronger cervical part, m. rhomboideus racalis longus.
cervicis, lies dorsolateral on the neck from the The m. sternocephalicus (Figs. 3-24,3-45) in
2nd or 3rd cervical vertebra to the 3rd thoracic the dog can be more or less clearly separated
vertebra. It arises on the tendinous raphe of the into mastoid and occipital parts. In large dogs
neck and the ends of the spinous processes of this flat muscle is 2.5 to 3.5 cm. wide at the ster
the first three thoracic vertebrae, and ends on num and 10 to 14 mm. thick. It arises as a unit
the rough medial surface and on the edge of the on the manubrium sterni and, covered only by
base of the scapula, including the scapular carti skin and the m. cutaneus colli, runs to the mas
lage close to the cervical angle. Near the scapula, toid part of the temporal bone and to the dorsal
in large dogs, it becomes as much as 1.5 cm. nuchal line of the occipital bone. At their origin
thick. From the cervical part, cranial to the 4th the muscles of the two sides are intimately
cervical vertebra, a lateral portion, the m. rhom joined, but they separate at or before the middle
boideus capitis, is given off as a straplike muscle of the neck, and each crosses under the external
to the occiput. In large dogs it is 2 cm. wide and jugular vein of its own side, and encroaches
1 to 2 mm. thick. The thoracic portion, m. rhom closely upon the ventral edge of the ipsilateral
boideus thoracis, in large dogs 4 to 8 mm. thick, m. cleidocervicalis.
arises on the spinous processes of the 4th to the The ventral portion, the m. sternomastoideus,
6th or 7th thoracic vertebra and inserts on the separates as a strong, elliptical bundle and,
medial and partly on the lateral edge of the base united with the m. cleidomastoideus in a strong
of the scapula. This portion of the m. rhomboid tendon, goes to the mastoid part of the temporal
eus is covered by the m. latissimus dorsi. The bone; the broader, thinner, dorsal segment, the
two portions are never clearly separated from m. sterno-occipitalis, attaches to the dorsal nu
each other and are often intimately bound to chal line as far as the mid line of the neck by
gether. means of a thin aponeurosis. Because of the di
Action: To elevate the limb, pull the limb and vergence of the two sternocephalic muscles,
shoulder forward or backward; to draw the there is a space ventral to the trachea in which
scapula against the trunk (in common with the bilateral mm. sternohyoideus and sternothy
all the extrinsic muscles). roideus appear. Here in the deep cervical fascia,
Innervation: Rami ventrales of nn. cervicales additional fibers for the m. sternomastoideus
et thoracales. may arise.
The m. serratus ventralis (Figs. 3-24, 3-44) The m. brachiocephalicus (Figs. 3-47,3-48,
covers the caudal half of the lateral surface of 3-52), lying on the neck under the m. sphincter
the neck and the cranial half of the lateral tho colli superficialis and platysma as a long, flat
racic wall; it is a very strong, fan-shaped muscle. muscle, extends between the brachium and the
It arises on the facies serrata of the scapula, its head and neck. Cranial to the shoulder the mus
fibers diverging to form an angle of about 150 cle is traversed by a clavicular remnant, a trans
degrees. It ends on the transverse processes of verse, often arched, fibrous intersection or plate,
the last five cervical vertebrae as the m. serratus the clavicular tendon (tendo clavicularis). The
ventralis cervicis, and on the first seven or eight vestigial clavicle is connected with the medial
ribs, somewhat ventral to their middle, as the m. end of the clavicular tendon and lies under the
serratus ventralis thoracalis. In large dogs the muscle.
muscle is 1.5 to 2 cm. thick near the scapula. In man, in whom the clavicle is completely
The terminal serrated edge of the cervical por developed, the m. cleidomastoideus extends
tion is not sharply defined; the individual slips from the medial end of the clavicle to the head;
insert between the m. longissimus cervicis and from the lateral end of the clavicle the pars cla
the m. intertransversarius. The thoracic portion vicularis of the m. deltoideus, which is closely
M u scles of the T h o r a c ic L im b 201
joined with the two other portions of the deltoid The m. latissimus dorsi (Figs. 3-48, 3 -4 9 ,3 -
muscle, extends to the arm. In the phylogenetic 52) is a flat, almost triangular muscle which lies
scheme, when the clavicle is reduced and short caudal to the muscles of the shoulder and bra-
ened, the origins of these muscles come closer chium on the dorsal half of the lateral thoracic
together until they fuse. The muscle now ex wall. It begins as a wide, tendinous leaf from the
tends from the arm to the head—the m. brachio- superficial leaf of the lumbodorsal fascia and
cephalicus. The m. cleidocephalicus, which ex thus from the spinous processes of the lumbar
tends up the neck from the clavicular tendon, is vertebrae and the last seven or eight thoracic
further divided into two portions. In the dog it vertebrae; and it arises muscularly from the last
divides into a superficial m. cleidocervicalis two or three ribs. Its fibers converge toward the
(pars cervicalis), which broadens and attaches to shoulder. The cranial border of the muscle lies
the dorsal part of the neck, and a deep m. cleido- under the m. trapezius thoracis, where it covers
mastoideus (pars mastoidea), which extends to the caudal angle of the scapula. The apical end
the mastoid part of the temporal bone. However, of the muscle encroaches upon the dorsal edge
the m. cleidobrachialis (pars brachialis of the of the deep pectoral and with it goes under the
brachiocephalicus), which runs from the clavicu shoulder and arm musculature, ending in an
lar tendon to the humerus, corresponds to the aponeurosis medially on the m. triceps; this
pars clavicularis of the m. deltoideus of man. aponeurosis partly blends with the tendon of the
The m. cleidobrachialis, 5 to 6 cm. broad and m. teres major to end on the teres tubercle and
5 to 8 mm. thick in large dogs, arises from a nar partly goes with the deep pectoral muscle to the
row part of the distal end of the humeral crest. medial fascia of the brachium. Laterally a tip of
It appears between the m. brachialis and m. bi the m. cutaneus trunci joins it; this is near the
ceps and, covering the shoulder joint cranially origin of the m. tensor fasciae antebrachii. Since
and somewhat laterally, ends on the clavicular the ventral border of the m. latissimus dorsi
tendon. With the m. pectoralis superficialis it gives off a bundle over the biceps to the m.
forms a flat border which corresponds to the col pectoralis profundus and with it inserts apo-
lateral sternal groove of the horse. neurotically on the crest of the major tubercle,
The m. cleidocervicalis is in an equally super the dog, like the cat, has a “muscular axillary
ficial position, and appears as a cranial extension arch” (Heiderich 1906 and Langworthy 1924).
of the m. cleidobrachialis from the clavicular Action: To draw the trunk forward and pos
tendon to the back of the neck. Nevertheless, sibly laterally; depress the vertebral col
there is no connection between fibers proximal umn; support the limb, draw the limb
and distal to the tendinous plate. Moreover, in against the trunk, draw the free limb back
the fascia of the triangle bounded by the mm. ward during flexion of the shoulder joint.
cleidocephalicus, pectoralis superficialis, and Innervation: Nn. pectorales caudales.
sternocephalicus there are scattered bundles The m. pectoralis superficialis (Figs. 3-46, 3 -
coming from the medial edge of the m. cleido 47, 3-48, 3-52) is the smaller of the two pectoral
cephalicus. The m. cleidocervicalis gradually be muscles and lies under the skin on the cranio-
comes broader and thinner as it goes to its ventral part of the thorax between the cranial
aponeurosis of insertion on the fibrous raphe of end of the sternum and the humerus. It arises
the cranial half of the neck. paramedially on the cranial end of the sternum
The m. cleidomastoideus is the deep cranial as far as the third costal cartilage. It runs later
continuation of the m. cleidocephalicus anterior ally and distally, and covers the m. biceps
to the clavicular tendon. It is covered by the brachii; then, with the m. cleidobrachialis, it
m. cleidocervicalis and m. sterno-occipitalis. It passes between the mm. biceps brachii and
reaches a width of 2.5 to 3 cm. and a thickness brachialis and ends, except for a small distal
of 7 to 10 mm., and is often split into two round part, on the entire crest of the major tubercle of
bundles throughout its length. By means of a the humerus. Cranially (in large dogs) the mus
strong tendon it ends on the mastoid part of the cle is 1.5 to 2 cm. thick; caudally it is thinner.
temporal bone with the m. sternomastoideus, Three divisions of the muscle are discernible.
dorsal to which it lies. Action: To support the limb, draw the limb in
Action: To draw the limb forward, draw the ward, draw the limb forward or backward
trunk backward, and, acting unilaterally, to according to its position, and draw the
fix the neck. trunk sideward.
Innervation: M. cleidocephalicus: n. accesso Innervation: Nn. pectorales craniales and also
rius, rami ventrales of the nn. cervicales; m. branches from nn. cervicales 7 and 8 (Lang
cleidobrachialis: n. axillaris. worthy 1924).
202 Chapter 3. M yo lo g y
My i oh y o i d e u s z Geni ogl os s us
, Styloglossus
Digastricus
Masseier
Hyoglossus
M a n d i b u l a r l ymph nodes
sMasseter
Thyrohyoideus
Pa ro ti d duct
/ Cricothuroideus
Stylohyoideus„
Parotid gland -
\ *
S t e r n o t h y ro i deus
Mandibular s o liv a rg gland /Trachea
M e d i a l r et r o p h a r y n g e a l In.
\
.Serratus ventralis
St er nomast o i deus )
Longus c a p i t i s
S ferno-occi pi falls z Tr apez i us
Ste rnohyoideus s' , Esophagus
Ext. j u g u l a r i/.- Common c a r o t i d a.
Cleidomas toideus^
Vagosympathetic trunk
C leidocervicaI i s Omotransversarius
C l a v i c u l a r tendon- — Supraspinatus
C l e i d o b r a c h i a I is - Subscapularis
~Scalenus
Del to i d e u s - "
fer’
i mus d o r s i
Su p n asp in atu si ,B r a c h i a l i s
in frasp in a tu s S u b s c a p u l a r i s - -/- - S u p r a s p in a t u s
T eres m in o r --D e e p p e c to ra l
_ T r ic e p s , a c c e s s o r y head-
T ric ep s, l a te ra l h e a d —
D elto id e u s- C onacobrachialis
-S u p e rfic ia l p e c t o r a l
B nachialis~
B ra ch io ce p h a licu s-
■ A n c o n e u s --------------------------------- Mf ■ B r a c h i o c e p h a li c u s
E x te n so r carpi r a d i a l i s -------
P ro n ato n teres
--E x ten so rs of
S u p in a to r -F lexors of
c a rp u s t d ig its
carpus f d ig its
F ig . 3-47. Left humerus, showing areas of muscle attach- F ig . 3-48. Left humerus, showing areas of muscle attach
ment, lateral aspect. ment, medial aspect.
204 Chapter 3. M yology
The m. pectoralis profundus (Figs. 3 -4 6 ,3 - strong muscular belly curves far around the neck
48, 3-52) is a broad muscle lying ventrally on the of the scapula so that it also appears on the me
thorax; it can be divided into a major portion dial surface of the shoulder. The entire muscle
and a minor superficial, lateral portion. It ex ends with a short, extremely strong tendon on
tends between the sternum and the humerus and the free edge of the major tubercle of the hu
corresponds to the pars humeralis of the same merus. In the distal third of the muscle, a strong
muscle of some other animals. It arises from the tendinous fold develops which extends into the
first to the last sternebra and, with a superficial terminal tendon. The end of the muscle appears
marginal portion as the pars abdominalis, from to be pennate. The caudal half of the muscle is
the deep fascia of the trunk in the region of the covered by a glistening tendinous sheet from the
xiphoid cartilage. Its fibers run cranially and lat spine of the scapula.
erally toward the brachium. It covers the ster Action: Extension of the shoulder joint and
num and the cartilages of the sternal ribs from forward advancement of the limb.
which it is separated by the aponeurosis of the • Innervation: N. suprascapularis.
mm. rectus abdominis and transversus costarum. The m. infraspinatus (Figs. 3-43, 3-47,3-50)
After going underneath the superficial pectoral, is covered largely by the m. deltoideus. It lies in
the major part of the muscle largely inserts, the infraspinous fossa and extends caudally
pardy muscularly and partly tendinously, on the somewhat beyond the fossa. It arises from the
minor tubercle of the humerus. An aponeurosis fossa, the scapular spine, and the caudal bor
goes over the m. biceps brachii to the major tu der of the scapula, and finally from the tendi
bercle. The superficial part, which originates nous sheet which covers it (shoulder aponeu
from the abdominal fascia, and which is crossed rosis and tendon of origin of the m. deltoideus).
laterally by the terminal fibers of the m. cuta- At the shoulder joint the fleshy muscle becomes
neus trunci, goes to the middle of the humerus. a strong tendon which crosses the caudal part of
There ther m. latissimus dorsi and the m. cuta- the major tubercle. The infraspinous bursa is
neus trunci attach to it. It then radiates into the found here. The muscle ends distal to the tuber
medial fascia of the brachium. cle. This tendon originates from the middle of
The muscle in large dogs is 2 to 2.5 cm. thick the muscle so that it is circumpennate in form.
in its cranial part; caudally it is thinner. The m. Proximal to the infraspinous bursa, which is
pectoralis superficialis covers it cranially. about 1 cm. in diameter in large dogs, there is
Action: To pull the trunk up on the advanced constantly found a second, smaller one.
limb; extend the shoulder joint; draw the Action: The muscle is the outward rotator and
limb backward. According to Slijper (1946), abductor of the humerus and a flexor or ex
the m. pectoralis profundus, along with the tensor of the shoulder joint, depending on
m. serratus ventralis, plays an important the position of the joint when the muscle
role in supporting the trunk, since its hu contracts. Its tendon functions as a lateral
meral insertion is considerably dorsal to its collateral ligament of the shoulder joint.
sternal origin. Innervation: N. suprascapularis.
Innervation: Nn. pectorales caudales, and also The m. teres minor (Figs. 3-43, 3-4 4 ,3 -4 7 ,
branches from nn. cervicalis 8 and thoraci- 3-51) lies distocaudally on the scapula on the
cus 1. flexor side of the shoulder joint, where it is cov
The lateral shoulder muscles. The lateral ered by the m. deltoideus and the m. infraspi
shoulder muscles, mm. supraspinatus and infra natus. It arises by an aponeurosis which lies on
spinatus, occupy the scapular fossae. Superfi the long head of the m. triceps, from the distal
cially, the m. deltoideus and the m. teres minor third of the caudal edge of the scapula, and pri
traverse the flexor angle of the shoulder joint lat marily from the infraglenoid tuberosity. It in
erally. serts by a short, strong tendon on a special
The m. supraspinatus (Figs. 3-43 and 3-47 to eminence of the humeral crest above the deltoid
3-50) is covered by the mm. trapezius cervicis tuberosity. It is covered on both sides by a ten
and omotransversarius. It fills the supraspinous dinous sheet.
fossa and curves over the lateral edge of the neck Action: Flexion of the shoulder joint.
of the scapula. It arises from the entire surface Innervation: N. axillaris.
of the supraspinous fossa, including the spine of The m. deltoideus (Figs. 3-43, 3 -4 5 ,3 -4 7 ,3 -
the scapula, and from the edge of the neck of the 50) is composed of two portions lying side by
scapula by numerous tendons from which the side. It lies superficially directly under the shoul
subscapularis also partly originates. Distally the der fascia between the scapular spine and the
M u sc les of the T h o r a c ic L im b 205
Scapula, s e r r a t e d fa c e —
-S u p ro & p m a tu s
T e re s m ajon-
T erdan o f in s e rt io n of
L a tissim u s d o r s i and Teres m aj.~ -H u m eru s, g r e a t e r tu b e rc le
T en so r fa sciae a n tebra ch H - - B ic e p s b ra c h ii
- -T ric e p s , m e d ia l h e a d
T ric e p s , loncf h ea d
H u m e ru s
b r a c h ia lis
- - S ca pula , sp in e
.S u p ra sp in a tu s -----
- I n f r a s p in a t u s
Scopula, a cra m ia n - — S ca pu la , c a u d a l b o r d e r
D eltoideus - ■ - T e re s m a jo r
h u m e ru s, g r e a t e r t u b e rc le - - T r ic e p s , lQn g h e a d
T ricep s, la t e r a l h e a d -----
T r ic e p s , lo n g head
B n a c h ta h s —
E x t e n s o r c a r p i r a d ia lis
B ic e p s -
In fra s p in a tu s -
T r ic e p s I
T ere s m in o r -
- Icn cj h e a d
T r ic e p s
a ccessa ry h e a d ' - a c c e s s o r y head
long h e a d - -
la t e r a l h e a d -------
♦V\ -i- - m e d ia l h ead
B r a c h ia l is — -
eps
- S u p ra s p m a tu s
C oro co bn a ch ta lis - G r e a t e r tu b e rc le
T e re s m m a r
In f r a s p in a t u s
T ni ceps,
medial head C o ra c o b ra c h ia h s
B r a c h ia lis
B r a c h ia lis
-B ice p s, tendon a f in s e rtio n
--A n ca n e u s
-B ra ch ia lis, tendon o f insertion
U lna' 'R a d iu s
Cephalic
Biceps - -
-Brachi ali s
Musculocutaneous n.-~
Medi an n. --
- - Medi al head
Ulnar n.~ - - Accessory hea d
>Triceps
Brachial v.' - - Lateral head
Loncj head
Superfi ci al p e c t o r a l -
Deep p e c t o r a l -
F ig . 3-52. Schematic plan of cross section through the middle of the arm.
M uscles of the T h o r a c ic L im b 209
tendon of insertion of the m. biceps brachii. Be tendon extends obliquely caudodistally over the
ginning at the tendon of origin, the muscle is medial side of the shoulder joint and thus lies in
covered by two extensive fibrous sheets which a groove close to the tendon of the m. subscapu
cover three-fourths to four-fifths of its length. laris. The muscle runs between the medial and
The narrower one is applied to the side of the accessory heads of the m. triceps brachii, ending
muscle next to the bone; the other is broader on the crest of the minor tubercle, as well as
and covers the cranial and medial surfaces. caudal to the crest between the medial head of
Pushed into the interior of the muscle is a strong the m. triceps brachii and the m. brachialis.
tendinous fold which, externally, is manifested From its insertion a delicate tendinous leaf ex
by a groove. The fold does not reach the proxi tends proximally over almost the entire muscle
mal tendon of origin; it makes the m. biceps belly.
brachii in the dog double pennate. The fibers of Action: Extension and adduction of the shoul
the m. biceps brachii run obliquely from both fi der joint.
brous coverings to the interior fibrous fold, so Innervation: N. musculocutaneus.
that their length is less than one-fifth that of the The caudal brachial muscles. The muscles
entire muscle. The m. biceps brachii in the dog which fill in the triangular space between the
is not composed of long fibers as is the case in scapula, humerus, and olecranon form a mighty
man; rather, it shows the first step toward the muscular mass. They are the extensors of the el
acquisition of a passive tendinous apparatus bow joint. The principal part of this musculature
(Kruger 1929), which in quadrupeds is neces is formed by the m. triceps brachii. The other
sary for the fixation of the shoulder joint when extensors of the elbow joint in the dog are the
standing. Distally from the interior fold, there m. anconeus and the m. tensor fasciae antebra
extends a tendinous strand in the groove be chii.
tween the m. extensor carpi radialis and m. pro The m. triceps brachii (Figs. 3-43 to 3-54)
nator teres; it crosses these muscles and spreads consists of four heads; caput longum, laterale,
out in the antebrachial fascia. It corresponds to mediale, and accessorium, with a common ten
the lacertus fibrosus of man. don to the olecranon. Where this tendon crosses
Action: Flexion of the elbow joint. the grooves and prominences of the proximal
Innervation: N. musculocutaneus. end of the ulna, a synovial bursa is interposed.
The m. brachialis (Figs. 3-47, 3-51, 3-52) The caput longum o f the m. triceps forms
arises muscularly from the proximal part of the a triangular muscle belly whose base lies on the
caudal surface of the humerus or proximal part caudal edge of the scapula and the apex on the
of the musculospiral groove. It extends laterally olecranon. The muscle arises partly fleshy and
as far as the humeral crest, and medially as far partly tendinously on the distolateral two-thirds
as the medial surface. It winds from the caudo of the caudal edge of the scapula and chiefly by
lateral to the cranial surface of the humerus in its tendon on the infraglenoid tuberosity. Its fibers,
course distally. At the distal third of the hu which are covered laterally by a rather weak and
merus it becomes narrower, goes over the flexor somewhat extensive fascia, converge toward the
surface of the elbow joint, lateral to the m. bi olecranon and end in a short, thick, round ten
ceps brachii, and ends partly fleshy on that part don. This tendon is attached to the caudal part
of the tendon of the m. biceps brachii which of the olecranon, but under the lateral head, it is
goes to the radial tuberosity. The remainder be supplemented by a fascial sheet which is strong
comes the tendon of insertion which goes to the distally and which radiates between the long
ulnar tuberosity between the two tendons of the head and lateral head in a proximal direction.
m. biceps brachii. The muscle is mostly covered This fascia also embraces the cranial edge of the
by the m. triceps. Medially it is covered by a muscle. The interior of the muscle reveals a
closely adherent fascial leaf which extends dis weak, tendinous strand which is parallel to the
tally to the m. extensor carpi radialis. surface. Between the terminal tendon and the
Action: Flexion of the elbow joint. cranial, grooved portion of the olecranon, there
Innervation: N. musculocutaneus, without the is a synovial bursa (bursa subtendinea olecrani),
participation of the n. axillaris (Reimers which may be over 1 cm. wide; here there is
1925). abundant fat. The muscle is interspersed with
The m. coracobrachialis (Figs. 3-44, 3 -4 8 ,3 - several tendinous bands and manifests distinct
49), short and rather thick, arises on the coracoid subdivisions. Near the scapula, the mm. deltoi
process of the scapula by a long, narrow tendon deus and teres minor are found laterally and the
which is surrounded by a synovial sheath. The m. teres major lies medially.
210 Chapter 3. M yo lo g y
The caput laterale o f the m. triceps is a strong, ceps brachii. It arises above the “axillary arch”
almost rectangular muscle lying between the from the thickened perimysium of the lateral
long head and the humerus. This muscle, which surface of the m. latissimus dorsi. It ends, in
blends with the accessory head and which lies common with the m. triceps brachii, in a ten
on the m. brachialis, arises on the humeral crest don on the olecranon, and independently in the
by an aponeurosis, which in small dogs is about antebrachial fascia. Occasionally one finds a
1 cm. wide. After emerging from the caudal bor synovial bursa between the muscle and the me
der of the m. teres major, its fibers run toward dial surface of the olecranon.
the olecranon and terminate in a broad, short Action: It supports the action of the m. triceps
tendon which blends partially with the tendon brachii and is the chief tensor of the ante
of the long head and partially with the deep leaf brachial fascia.
of the antebrachial fascia. Innervation: N. radialis.
The caput m ediale o f the m. triceps is a spin-
dle-shaped muscle which arises tendinously on
the crest of the minor tubercle between the The Antebrachial Muscles
point of insertion of the teres major and that
of the m. coracobrachialis. A strong, tendinous The muscles of the forearm embrace the
fascia extends over the proximal two-thirds of bones in such a way that the distal two-thirds of
the muscle. It attaches medially and independ the medial side of the antebrachial skeleton (es
ently on the olecranon. In addition, the tendon pecially the radius) is uncovered. The extensors
blends with that of the long head and continues of the carpus and digits lie dorsally and laterally.
into the antebrachial fascia. The bursa subten- The carpal and digital joints of the thoracic limb
dinea olecrani is underneath the tendon. have equivalent angles; that is, their extensor
The cap u t accessoriu m o f the m. triceps, ir surfaces are directed dorsally or cranially. On
regularly rectangular in cross section, lies on the the palmar side are the flexors of the joints. The
caudal side of the humerus between the other mm. pronator teres and supinator serve to turn
heads of the m. triceps brachii and the m. bra the forepaw about the long axis; these are found
chialis. It arises from the proximal caudal part in the flexor angle of the elbow joint. Because
of the neck of the humerus, and becomes ten most of the muscles appear on the palmar side,
dinous at the distal third of the humerus. The the antebrachium of the dog appears to be com
tendon is elliptical in cross section and blends pressed laterally. Since the muscle bellies are lo
with that of the long and lateral heads and thus cated proximally and the slender tendons dis-
inserts on the olecranon. The common tendon tally, the extremity tapers toward the paw.
lies over the subtendinous bursa. The dorsolateral antebrachial muscles. The
Action: Extend the elbow joint. dorsolateral group of antebrachial muscles are
Innervation: N. radialis. represented chiefly by the extensors of the car
The short, strong m. anconeus (Figs. 3-45, 3 - pal and digital joints. These are the mm. exten
47, 3-4 8 , 3-51, 3-54) lies on the caudal side of sor carpi radialis, extensor digitorum communis,
the distal half of the humerus between the epi- extensor digitorum lateralis, extensor carpi ul-
condyles. It arises on the lateral condyloid crest, naris, extensor pollicis longus et indicis proprius,
the lateral epicondyle, and, since it almost com and abductor pollicis longus. To these are added
pletely fills the olecranon fossa, part of the me the mm. brachioradialis and the supinator in the
dial epicondyle also. It ends on the lateral sur flexor angle of the elbow joint. The majority of
face of the proximal end of the ulna and is mostly these muscles arise directly or indirectly from
covered by the m. triceps brachii. It covers the the lateral (extensor) epicondyle of the humerus.
proximal surface of the elbow joint capsule and The m. brachioradialis (Figs. 3-47, 3-55, 3 -
one of its outpocketings. 57), much reduced and occasionally lacking, is a
Action: The m. anconeus, with the m. triceps long, narrow muscle in the flexor angle of the el
brachii, extends the elbow joint, and helps bow joint; in large dogs it is 0.5 to 0.75 cm. wide
tense the antebrachial fascia. and about 1 mm. thick. It is cranial in position
Innervation: N. radialis. between the superficial and the deep antebra
The m. tensor fasciae antebrachii (Figs. 3-49, chial fascia, and is intimately bound to the
3-52, 3-53) is a flat, broad, straplike muscle superficial leaf of the latter fascia. It arises on the
which, in large dogs, is only 2 mm. thick; it lies proximal end of the lateral condyloid crest of the
on the caudal half of the medial surface and on humerus directly above the m. extensor carpi
the caudal edge of the long head of the m. tri radialis. It extends cranially at first beside the m.
M u sc les of the T h o r a c ic L im b 211
F ig . 3-53. Left radius and ulna, showing areas of muscle F ig . 3-54. Left radius and ulna, showing areas of muscle
attachment, medial aspect. attachment, lateral aspect.
212 Chapter 3. M yo lo g y
B ic e p s -
B n o c h io r o d ia lis -
E x te n so r c o r p i-
r a d ia lis
R a d iu s -
B - Extensor ca rp i u ln a ris
E x te n so r ca rp i - Mr
r a d ia lis — E x ten sor d ig it o ru m
la f e r a l is
E x te n so r d ig i torum
Fic. 3-55. Forearm with antebrachial fascia, cranial aspect. c o m m u n is
/ Ib d u c .t o r p o l l i c i s - - ,
lo n g u s
- E x t e n s o r p o llic is lo n g u s et
in d i c is p r o p r iu s
extensor carpi radialis, then turns more medially Action: Extension of the carpal joint and flex
and extends distally in the groove between the ion of the elbow joint.
m. extensor carpi radialis and the radius. Be Innervation: N. radialis.
tween the third and the distal fourth of the bone The m. extensor digitorum communis (Figs.
it ends on the periosteum by a thin aponeurosis. 3-47, 3-56 to 3-58, 3-64 to 3-66) lies on the
Action: Rotation of the radius dorsolaterally. craniolateral surface of the radius between the
Innervation: N. radialis. m. extensor carpi radialis and the m. extensor
The m. extensor carpi radialis (Figs. 3 -4 7 ,3 - digitorum lateralis. It arises on the lateral epi
56 to 3-59, 3-66) is a long, strong, fleshy muscle condyle somewhat in front of and above the at
lying on the cranial surface of the radius medial tachment of the ulnar collateral ligament of the
to the m. extensor digitorum communis. It is the elbow joint, and with a smaller portion from the
first muscle encountered after the free surface antebrachial fascia. At its origin it is fused
of the radius, when one palpates from the medial deeply with the m. extensor carpi radialis by a
to the dorsal surface. The m. extensor carpi radi common aponeurosis which separates into two
alis arises on the lateral condyloid crest of the parts distally, one for each muscle. After the ap
humerus, united with the m. extensor digitorum pearance of a corresponding number of superfi
communis for a short distance by an intermus- cial tendinous bands, the slender belly divides
cular septum. It forms a muscle belly which into four bellies and tendons distally; at first
fades distally and splits into two flat tendons at these lie so close together that the whole tendon
the distal third of the radius. This muscle in the appears to be undivided. At the same time the
dog reminds one of the relations prevailing in deep muscle fibers extend over to the medial
man: an incomplete division into a weaker, more tendon. The compound tendon, enclosed in a
superficial, medial m. extensor carpi radialis common synovial sheath, extends distally on the
longus, and a stronger, deeper, more lateral m. m. abductor pollicis longus and passes through
extensor carpi radialis brevis. The deep muscle the lateral distal sulcus of the radius, where it is
is limited on its deep surface by a fascial leaf covered by a strong indistinct transverse liga
which extends from the lateral epicondyle to its ment. After the tendon crosses the extensor sur
terminal tendon. Both tendons are closely ap face of the carpal joint, the individual tendons
proximated as they extend distally along the ra separate from each other and pass on the exten
dius. By way of the middle sulcus of the radius, sor surface of the corresponding metacarpal
they gain the extensor surface of the carpus, bones and phalanges to the distal phalanges of
where they lie in a groove formed by the dorsal digits II to V, inclusive. Here each tendon
transverse carpal ligament. They are often sur broadens into a caplike structure and ends on
rounded by a synovial sheath. The tendons sep the dorsal portion of the edge of the horn of the
arate; one inserts on a small tuberosity on meta distal phalanx, covered by the crura of the dor
carpal II (m. extensor carpi radialis longus) and sal elastic ligaments. The m. extensor digitorum
the other on metacarpal III (m. extensor carpi communis is composed of digital extensors II,
radialis brevis). From the aponeurosis covering III, IV, and V. Each tendon, at the distal end of
the medial surface of the m. brachialis arises a the proximal phalanx, receives bilaterally thin
fascial leaf which extends over the proximal me check ligaments which cross obliquely from the
dial surface of the belly of the m. extensor carpi palmar mm. interossei. The tendons of the lat
radialis, as does a fascial leaf from the m. biceps eral digital extensor unite with the tendons of
brachii. The most proximal part of the muscle the common digital extensor on digits III, IV,
lies on the joint capsule, which forms a bursa and V. Thus, all extensor tendons are deeply em
like pocket at this point. In about half of all bedded in the dorsal fibrous tissue of the digits.
specimens the tendons are completely or almost Under the origin of the m. extensor digitorum
completely surrounded by a cpmmon tendon communis there extends an outpouching of the
sheath which extends from the beginning of the elbow joint capsule. The separation of the ter
tendon to the proximal end of the metacarpus. A minal portion of the muscle is usually described
synovial bursa may exist at the proximal row of as distinct, although an undivided muscle is sim
carpal bones under both tendons or only under ulated. On the other hand, the tendons may fuse
the lateral tendon. A second bursa is occasion in part with one another; this is especially true
ally found under the lateral tendon at the distal for the tendons of digits IV and V. The muscle
row of carpal bones. In other specimens, in branch for digit II is the longest and becomes
place of the synovial sheath, one finds loosely tendinous at the middle of the antebrachium.
meshed tissue. The three remaining muscle branches reach only
214 Chapter 3. M yology
to the middle third of the antebrachium. The digitorum communis medially to the third and
synovial sheath which surrounds the tendon fourth metacarpophalangeal joints; on the proxi
bundle of this muscle and that of the m. extensor mal phalanx of digits III and IV they unite with
pollicis longus et indicis proprius begins shortly the corresponding tendons of the common digit
after the muscle has become tendinous (in large al extensor; often they also unite with one or
dogs 3 to 4 cm. above the carpus). It reaches at both of the check ligaments which come from
least to the middle of the carpus, often to the the m. interossei. They end principally on the
proximal end of the metacarpus. Its fibrosa fuses distal phalanges of digits III and IV. The ten
with the periosteum of the radius and with the dons of the lateral digital extensor are only about
joint capsule of the carpus, that is, with the dor one-third the width of those of the common dig
sal carpal ligament. Its mesotendon, which ap ital extensor.
pears at its medial border, first covers the tendon In about one half of all specimens, both ten
of the m. extensor pollicis longus et indicis pro dons are enclosed in a common synovial sheath,
prius and then the four tendons of the m. exten which in large dogs begins 2.5 to 3 cm. above
sor digitorum communis. At the metacarpopha the carpus and often reaches the metacarpus.
langeal joint the tendon glides on the sesamoid In others there is no distinct synovial sheath, but
element which is embedded in the joint capsule; rather there is a space under both tendons
this sesamoid has an ossified nucleus, whereas bounded by the fascia. In exceptional specimens
those at the proximal interphalangeal joints re the tendon for digit III is independent and arises
main cartilaginous. from the fascia distal to the carpus (Ziegler
Action: Extension of the joints of the four 1929).
principal digits. Action: Extension of the joints of digits III, IV,
Innervation: N. radialis. and V.
The m. extensor digitorum lateralis (Figs. 3 - Innervation: N. radialis.
47, 3 -5 6 to 3-58, 3-65, 3-66) is somewhat simi The strong m. extensor carpi ulnaris (Figs. 3 -
lar to the common extensor in strength; in the 56, 3-57, 3-61, 3-65, 3-66) lies on the caudolat-
antebrachium it lies laterally on the radius be eral side of the ulna, and is directly under the
tween the m. extensor digitorum communis and fascia. It arises on the lateral or extensor epi-
the m. extensor carpi ulnaris. It covers the m. condyle of the humerus behind the ulnar col
abductor pollicis longus. The muscle has two lateral ligament of the elbow joint by a long,
bellies. It arises on the cranial edge of the ulnar relatively strong tendon. At the middle of the
collateral ligament of the elbow joint, and on antebrachium the strong distal tendinous band
the head and lateral tuberosity of the radius. A of the lateral surface, which eventually goes into
band from the ulnar collateral ligament runs un the strong terminal tendon, takes on the delicate
der the muscle, then separates into two branches distal tendinous leaf of the medial surface. On
in its distal half, each half going into a tendon. the medial surface of the terminal tendon, fibers
The tendon adjacent to the common digital ex of the deeper muscle mass radiate into the broad
tensor is the weaker one and comes from a slen tendon as far as the carpus; the tendon passes
der, distal fascial sheet; the other tendon arises laterally over the carpus, being held in place by
from a considerably stronger, distal fascial leaf connective tissue without a sulcus in which to
which lies next to the m. extensor carpi ulnaris. glide. It ends laterally on the proximal end of
The tendons lie close together and usually are metacarpal V. From the accessory carpal bone,
enclosed in a common synovial sheath. They two fiber bundles arise from the antebrachial
pass through the groove between the distal ends fascia and cross each other to blend with the
of the radius and ulna, over the dorsolateral bor tendon of the m. extensor carpi ulnaris at the
der of the carpus to the metacarpus, and then carpus. A tendinous fold, which parallels the sur
diverge from each other. The tendon of the face, is concealed in the interior of the muscle.
stronger caudal belly extends from metacarpal V Under the tendon of origin of the muscle in
to the proximal phalanx of digit V, unites with older dogs, there is constantly a synovial bursa,
the corresponding tendon of the m. extensor dig 1 to 2 cm. in diameter; a second bursa is found
itorum communis and ends with it on the distal occasionally, between the tendon and the distal
phalanx as well as on the dorsal surface of the end of the ulna.
proximal ends of the proximal and middle pha Action: Extension of the carpal joint with
langes. The tendon of the weaker belly divides weak lateral rotation.
at the carpus into two branches which extend Innervation: N. radialis.
obliquely under the tendons of the m. extensor The m. supinator (Figs. 3-47, 3-53, 3-59) is
M u sc les of the T h o r a c ic L im b 215
cra n ia l
M e d i a n a.,v. t n. - -
Lateral digital extensor
Pronator quadratus-
-Volar i nterosseous a. <t v.
Volar a n t e b r a c h i a l a. 4- v.-
-Abductor pollicis longus
Deep dicj. flexor? radial head-
-Extensor carpi ulnaris
F l e x o r c a rp i r adi al is -
Deep d i g i t a l f l e x o r ;
Deep dig. flexoi? humeral head ulnar head
--Accessory volar
interosseous a. 4 v,
Superficial digital fl e x o r
Ulnar n.
caudal
F ig . 3-57. Schematic plan of cross section of the forearm between the proximal and middle thirds.
216 Chapter 3. M yology
Tendon to 3 na d i j i t
C
F ig . 3-58. Tendons on the dorsum of the left forepaw.
A. Dorsal aspect.
B. Lateral aspect, with tendons of extensor digitorum communis removed.
C. Two common variations.
M u sc les of the T h o r a c ic L im b 217
broad and flat and is almost completely covered don of the m. extensor carpi radialis, passes into
by a delicate fascia. It lies laterally in the flexor the medial sulcus of the radius and crosses the
surface of the elbow, covered by the m. extensor medial border of the carpus under the short
carpi radialis and the digital extensors. It lies di collateral ligament. Finally, the tendon in
rectly on the joint capsule and radius. It arises serts medially on the proximal end of meta
by a short, strong tendon on the ulnar collateral carpal I, where a sesamoid bone is embedded
ligament of the elbow joint, and on the lateral in it.
epicondyle. It extends obliquely distomedially Where the tendon goes over that of the m. ex
and, covering the proximal fourth of the radius, tensor carpi radialis, there is usually a bursa or a
ends on its dorsal surface as far as the medial short synovial sheath.
border. The muscle pushes a short distance Action: Abduction and extension of the first
under the border of the m. pronator teres. digit; medial deviation of the forepaw.
Action: Rotation of the paw so that the palmar Innervation: N. radialis.
surface faces medially. The caudal antebrachial muscles. The cau
Innervation: N. radialis. dal group of antebrachial muscles consists of the
The m. extensor pollicis longus et indicis flexors of the carpus and digits: mm. flexor carpi
proprius (Figs. 3-54, 3-59, 3-66) is an exceed radialis, flexor carpi ulnaris, flexor digitorum
ingly small, slender, flat muscle on the lateral superficialis, and flexor digitorum profundus. To
side of the antebrachium, where it is covered by these are added the small mm. pronator teres
the m. extensor carpi ulnaris, and the lateral and and pronator quadratus, which do not extend
common digital extensors. After arising from beyond the antebrachium. Most of these muscles
about the middle third of the dorsolateral border come from the medial or flexor epicondyle. They
of the ulna, adjacent to the m. abductor pollicis form the caudal part of the antebrachium.
longus, it runs distally, parallel to the ulna. Its When viewed medially, the muscles appear
fibers run obliquely distomedially. The muscle in the following order, beginning cranially: mm.
gradually crosses under the extensors, so that its pronator teres (only the proximal third of the
extremely delicate tendon appears medial to the forearm), flexor carpi radialis, flexor digitorum
common extensor tendons at the carpal joint, profundus (only the distal half of the forearm),
where the two tendons are surrounded by a flexor digitorum superficialis, and caput ulnare
common synovial sheath. On the dorsal surface of the m. flexor carpi ulnaris (only in a very in
of metacarpal III the tendon divides into two significant and jjroximal segment of the ante
parts. The medial portion expands over meta brachium). Seen from the palmar aspect, the
carpal II to the distal end of metacarpal I, where caput ulnare of the m. flexor carpi ulnaris is
it is buried in the fascia. The lateral portion, after medial to the m. flexor digitorum superficialis.
passing under the tendon of the m. extensor dig However, at the distal half of the antebrachium,
itorum communis to digit II, unites with it at the the caput humerale of the m. flexor carpi ulnaris
metacarpophalangeal joint. Rarely another weak pushes between these two. In the dog, the caput
tendon goes to digit III, and occasionally the humerale is deep; in other animals, when viewed
muscle splits into two bellies. from the medial aspect, it is covered by the m.
Action: Extension of digits I and II and ad flexor digitorum superficialis. The m. extensor
duction of digit I. carpi ulnaris is next to the m. flexor carpi ulnaris
Innervation: N. radialis. on the lateral surface of the forearm.
The m. abductor pollicis longus (Figs. 3-54, The m. pronator teres (Figs. 3-48, 3-53, 3 -
3-57, 3-59, 3-60, 3-67), almost completely 57, 3-59, 3-60), round in cross section, crosses
covered by the digital extensors, lies in the the medial surface of the elbow joint. It lies
lateral groove between the radius and the ulna. under the skin and fascia largely on the proximal
It arises on the lateral surface of the radius and third of the radius. It arises from the medial
ulna, and the interosseous membrane. Its fibers, epicondyle in front of the m. flexor carpi radialis.
which are directed obliquely medially and dis The body of the muscle extends obliquely crani-
tally, blend into a narrow but strong tendinous odistally and, upon forming a strong tendinous
band which proceeds along the dorsomedial band, ends distal to the m. supinator on the
border of the muscle and which becomes the medial border of the radius as far as its middle.
terminal tendon toward the carpus, after it has Its internal surface is provided with a strong,
bridged the gap between the tendons of the m. proximal tendinous band.
extensor digitorum communis and the m. ex Action: It rotates the forearm so that the dorsal
tensor carpi radialis. Its tendon crosses the ten surface tends to become medial. It may
218 C h a p te r 3 . M yo lo g y
Biceps - — S r a c h ia h s
- O lecranon
- v5u p i n a t o r
P r on o f o r te re s
R a d iu s , d o rs a l s u r f a c e -
A b d u c t o r p o llic is lo n q u s
---- E x t e n s o n p o llic is lo n q u s ef
in d ic is p n o p riu s
E x te n s o r c a r p i r a d ia h s —
- - L a t e r a l collateral
I, qam en t
S upm c to r -
— E x te n s o r d iq itc r u m la te ra lis
T r ic e p s , m e d ia l h e ad—
- ~ - H u m e r us, m e d ia l a p ic o n d ^ le
F l e x o r c a r p i u ln a ris ,- -
- F l e x o r d ig it o r u m p r o f u n d u s , h u m e ra l head
u ln a r head
Pn on o f o r te re s
- - Radius
- F l e x o r d iQ i t o r u m p ro f, ra d ia l head
F le x o r d ig ito r u m p ro f, hu m era l h e a d --
- - A b d u c t o r p o llic is lo n q u s
F le x o r ca rp i u ln a r is , u l n a r head -
A b d u c t o r p f l l i c i a b r e v i s etO pponens p o l l i c i s - -
v m 11 - E x t e r s o n p o l l ic is lo n q u s
F le x o r d ig it o r u m s u p e rf, te n d o n to 2 nd d i g i t — e t in d ic is p r o p r i u s
- - In terosseous m
F le x o r d g i to ru m p r o f u n d u s , tendon to 1 st d i g i t -----
- 2 nd m e t a c a r p a l bone
F l e x o r ClCjitonurn s u p e r f . ------ E x t e n s o r d ig it o r u m
P r o x im a l com m u m s
A n n u l a r h e ja m e n ts Mi d d l e
D is fa !
F le x o r d to ru m p ro fu n d u s
Fic. 3 60. Muscles on the medial surface of the left forearm and forepaw.
220 Chapter 3. M yology
function only as a flexor of the elbow joint border of the palmar surface of the middle
(Zimmermann 1928). phalanx after being “perforated” by one of the
Innervation: N. medianus. deep flexor tendons passing to a distal phalanx.
The m. flexor carpi radialis (Figs. 3-57,3-60, Each branch of the superficial tendon, at the
3-65, 3-67) lies in the medial part of the ante metacarpophalangeal joint, forms a tubelike en
brachium directly under the skin and ante closure around the deep flexor tendon. The
brachial fascia, where it covers the m. flexor dig proximal edge of this sleeve projects a short dis
itorum profundus. It arises on the medial epi tance proximally beyond the articular surfaces
condyle behind the radial collateral ligament of of the sesamoid bones. Distal to the sesamoids,
the elbow joint between the m. pronator teres the tube is so split for the passage of the deep
and the m. flexor digitorum profundus. It ex tendon that the superficial tendon appears to be
tends distally between the m. pronator teres and in two branches when it is viewed from the pal
the m. flexor digitorum superficialis and, form mar side. The deep part of the sleeve, however,
ing a short, thick fusiform belly, merges into a accompanies the enclosed tendon farther and
flat tendon near the middle of the radius. It re attaches to the palmar, proximal edge of the
ceives a delicate supporting fascia from the middle phalanx throughout its breadth. The
radius throughout its entire length. At the flexor four terminal tendons of the muscle are as a rule
surface of the carpus it runs through the palmar of equal strength (in large dogs, about 5 mm.
carpal ligament, where it is enclosed in a syno wide and 0.5 mm. thick). The branch to digit V
vial sheath. At the metacarpus, it splits into two is much weaker. At the metacarpophalangeal
strong tendons which end on the palmar side of joint and at the proximal and middle digital
metacarpals II and III, very close to the proxi joints, the branches of the superficial and deep
mal articular surface. The end of the muscle digital flexor tendons are bridged by the three
bears a lateral tendinous band and a delicate well-defined proximal, middle, and distal trans
medial one. A projection from the joint capsule verse ligaments.
extends under the muscle at its origin. Beneath the origin of the superficial digital
Action: Flexion of the carpal joint. flexor there is a synovial bursa 2 to 2.5 cm. long
Innervation: N. medianus. in large dogs. This communicates with a second
The strong, flat m. flexor digitorum superfi bursa beneath the origin of the caput humerale
cialis (Figs. 3-57, 3-60, 3-61, 3-64, 3-65,3-67) of the m. flexor carpi ulnaris. Each of the four
in dogs, in contrast to that in other animals, lies terminal tendons has a long digital synovial
directly beneath the skin and antebrachial fascia sheath. This sheath is described more fully
in the caudomedial part of the antebrachium. It below, in the discussion of the m. flexor digito
covers the m. flexor digitorum profundus and rum profundus.
the humeral head of the m. flexor carpi ulnaris. Action: Flexion of the proximal and middle
It arises by a short but strong tendon on the digital joints of the four principal digits and
medial or flexor epicondyle cranial to the thereby of the whole forepaw.
humeral head of the m. flexor carpi ulnaris and Innervation: N. medianus.
somewhat proximal to the flexor digitorum pro The m. flexor carpi ulnaris (Figs. 3-53,3-57,
fundus. The fleshy muscle belly reaches far dis 3-61, 3-63, 3-67) consists of two bellies, which
tally and becomes tendinous only a short dis converge into a tendon ending on the accessory
tance above the carpus. Its tendon is strong, carpal bone. The muscle lies caudolaterally on
elliptical in cross section, about 1 cm. wide and the antebrachium, with its weaker ulnar head
0.5 cm. thick. This tendon runs over the flexor most superficial and lateral to (and, in part,
surface of the carpus medial to the accessory upon) the m. flexor digitorum superficialis. The
carpal bone, but is not enclosed in the carpal much stronger humeral head is in the second
synovial sheath, as it crosses the palmar carpal layer of the palmar musculature beneath the m.
transverse ligament superficially. By means of flexor digitorum superficialis and upon the m.
this thick ligament it is separated from the deep flexor digitorum profundus.
flexor tendon. In the proximal third of the meta The rather flat ulnar h ead (caput ulnare),
carpus the tendon splits into four parts, which which is straplike in small dogs, arises medially
diverge to the second to fifth metacarpophalan on the palmar border of the proximal end of the
geal joints. Lying in a palmar relationship to the ulna and is covered at its origin by the terminal
corresponding terminal tendons of the deep tendon of the medial head of the m. triceps bra
flexor tendon, each extends over the respective chii. Above the middle of the antebrachium the
proximal phalanx and ends on the proximal ulnar head becomes a flat tendon which extends
M u sc les of the T h o r a c ic L im b 221
distally, lateral and palmar to the m. flexor digi on the medial epicondyle of the humerus, imme
torum superficialis covering the humeral head. diately caudal to the tendon of origin of the m.
Toward the accessory carpal bone the tendon flexor carpi radialis and covered by that of the
gradually dips beneath the terminal tendon of m. flexor digitorum superficialis, and the hu
the humeral head and ends independently on the meral head of the m. flexor carpi ulnaris. The
accessory carpal bone. The strong antebrachial strongly constructed body of the muscle lies on
fascia fuses with it throughout its length. the caudomedial side of the antebrachium in
The much stronger hum eral h ead (caput hu- such a way that it appears partly enclosed be
merale) arises on the medial or flexor epicondyle tween the radial head medially and deeply, and
of the humerus by a short, strong tendon which the ulnar head laterally. Near the carpus, or at
is a close neighbor of that of the m. flexor digi the border between the carpus and antebra
torum superficialis. It ends by an equally short, chium, the tendons of the humeral head fuse into
strong tendon on the accessory carpal bone. This a flat, but strong, main tendon which is grooved
strong, flat muscle, as much as 3 cm. wide and 1 on its palmar surface. Just distal to the groove
cm. thick, in the dog, in contrast to that in other the weaker tendons of the radial and ulnar heads
mammals, is almost completely covered by the converge to form the deep flexor tendon.
m. flexor digitorum superficialis. Only the lateral The radial head (caput radiale) of the m.
edge of its distal half and its terminal tendon en flexor digitorum profundus lies on the caudome
croach upon the antebrachial fascia. Both sur dial surface of the radius, among the mm. prona
faces of its body are covered by a tendinous tor quadratus, pronator teres, flexor carpi radi
sheet. The palmar sheet is almost entirely a dis alis, and the humeral head of the flexor digitorum
tal one, the dorsal tendinous sheet is equally ex profundus. It arises, as the weakest division of
tensive proximally and distally, and is provided the entire muscle, from the medial border and
with a narrow but strong tendinous sulcus for a small distance also on the caudal surface of
which, near the middle, appears to be displaced the proximal three-fifths of the radius. Near the
somewhat medially. Thus the muscle has a com carpus it forms a thin, flat tendon, which runs
plicated fiber structure. from a slender tendinous sheet on the proximal,
Beneath the origin of this muscle is found a caudal border of the muscle. This joins the
synovial bursa which communicates with the strong tendon of the humeral head at the proxi
one beneath the origin of the m. flexor digitorum mal border of the carpus. After being united for
superficialis. A second bursa, beneath the termi a short distance with the principal tendon, it
nal tendon, in large dogs extends proximally 1 to again splits away to insert on digit I.
1.5 cm. from the accessory carpal bone. According to Kajava (1922), the muscle does
Action: Flexion of the forepaw with abduction. not correspond to the m. flexor pollicis longus of
Innervation: N. ulnaris. man but is only a division of the m. flexor digi
The m. flexor digitorum profundus (Figs. 3 - torum profundus. Exceptionally, the branch of
53, 3-57, 3-60, 3-63, 3-64, 3-67) consists of the deep tendon going to the first^digit is inde
three heads and, generally speaking, forms the pendent in the dog; in such a case the tendon
deepest layer of the caudal musculature of the proceeds from the palmar carpal fascia (Zieg
forearm. It is covered by the mm. flexor carpi ra ler 1931).
dialis, flexor digitorum superficialis, and flexor The ulnar head (caput ulnare) of the m. flexor
carpi ulnaris. Its bellies, along with the m. pro digitorum profundus is stronger than the radial
nator quadratus, lie directly on the caudal sur head. It is a flat muscle at the caudal side of the
face of the radius and ulna. It consists of ulna, located among the m. extensor carpi ul
humeral, radial, and ulnar heads, which repre naris, flexor carpi ulnaris, and the humeral head
sent completely separate muscles whose tendons of the m. flexor digitorum profundus. It is
fuse to form the strong, deep digital flexor ten covered superficially by both the m. flexor
don. The homologization of these three muscles and the m. extensor carpi ulnaris. It arises on the
is still in dispute (Kajava 1922). caudal border of the ulna from the distal portion
The humeral head (caput humerale) of the m. of the medial ridge of the olecranon to the distal
flexor digitorum profundus, as the strongest di fourth of the ulna. Its fibers run obliquely distally
vision of the entire muscle, consists of three and caudally to a strong, broad tendinous sheet
bellies which are difficult to isolate. It is pro which accompanies the palmar edge of the mus
vided with tendinous sheets and bands and thus cle almost from the level of the elbow joint. At
has a very complex fiber arrangement. The three the distal fourth of the antebrachium the muscle
bellies arise by a common short, strong tendon ends in a tendon which is soon united with the
222 Chapter 3. M yology
common tendon of the m. flexor digitorum pro they also enclose the superficial flexor tendons.
fundus. Here their fibrosa fuses with the proximal trans
The deep flexor tendon crosses the flexor sur verse ligaments. To each collateral border of the
face of the carpus in a groove which is converted deeply situated and flattened superficial flexor
into the carpal canal by the transverse palmar tendons, there extends a mesotendon. Distally
carpal ligament. The tendon is very wide and, the synovial sheaths also enclose the middle
because of great thickening of its edges, forms a transverse ligaments of the flexors so that they
palmar groove. In the proximal portion of the receive mesotendons extending to their proximal
metacarpus, from its medial border, the deep edges. Farther distally the deep branches alone
flexor tendon gives off the round, weak tendon are surrounded by synovial sheaths. The distal
to digit I. Shortly thereafter the principal tendon transverse ligaments, however, do not push into
divides into four branches, for digits II to V, the synovial spaces, but fuse with the palmar and
which run distally, covered by the correspond lateral walls of the synovial sheaths.
ing grooved branches of the m. flexor digitorum A ction: The m. flexor digitorum profundus is
superficialis. At the level of the sesamoids of the the flexor of the forepaw (carpus and digital
metacarpophalangeal joints of digits II to V they joints).
pass through the tubular sheaths formed by the Innervation: The radial head as well as the
branches of the superficial digital flexor tendons. deep and medial portions of the humeral
They emerge from the palmar sheaths, extend head, n. medianus; the lateral portion of the
over the flexor surface of the distal digital joints, humeral head and the ulnar head, n. medi
and end on the tuberosities of the distal phalan anus and n. ulnaris (Agduhr 1915).
ges of digits II to V. The m. pronator quadratus (Figs. 3-53,3-57,
The two digital flexor tendons on each of the 3-62) fills in the space between the radius and
four main digits are held in place by three trans the ulna medially. It is rhomboidal in oudine,
verse (annular) ligaments. The proximal one lies and covers the interosseous membrane, a por
at the metacarpophalangeal joint and runs be tion of the medial surface of the ulna, and the
tween the collateral borders of the sesamoid caudal surface of the radius, except for the proxi
bones. The middle one lies on the proximal pha mal and distal ends. It is covered by the m.
lanx, at about its middle, and the distal one lies flexor digitorum profundus. Its fibers run from
immediately distal to the proximal interphalan the ulna obliquely, distally and medially, to in
geal joint. The distal one is lacking in digit I. sert on the radius.
Synovial apparatus. Beneath the origin of Action: Turn the forepaw inward.
the m. flexor digitorum profundus is a synovial Innervation: N. medianus.
bursa. The terminal tendon of the muscle, as it
passes through the carpal canal, is partly or The Muscles of the Forepaw
wholly surrounded by a synovial sheath. This ex
tends from the distal end of the radius to the In the forepaw are the tendons of the antebra
metacarpus. It may be replaced by a sac with chial muscles which insert on the metacarpal
rough, weak walls, without synovia. In the digits bones and phalanges, as well as the special mus
the digital synovial sheaths are formed around cles which are confined to the palmar surface of
the individual branches of the deep flexor ten the forepaw. Digits I to V are thus covered by a
don. The branch going to the first digit has its large number of muscles. A portion of these spe
own sheath, which extends from the middle of cial muscles lies between the large flexor ten
metacarpal I to the tuberosity of the distal pha dons; another portion lies between these and
lanx. The principal tendons going to digits II to the skeleton, either lying directly upon the four
V, however, bear synovial sheaths which are also large metacarpal bones or occurring as special
common for the superficial flexor tendons. These muscles of digits I, II, and V.
extend from the ends of the metacarpal bones to The muscles lying between the flexor ten
the tuberosities of the distal phalanges. These dons. These muscles include the m. interflex-
common synovial sheaths begin, in large dogs, 1 orus, m. flexor digitorum brevis, and the mm.
to 1.5 cm. proximal to the metacarpophalangeal lumbricales.
articulations immediately proximal to the sesa The weak m. interflexorius (Fig. 3-63) is the
moid bones in the region of the proximal ends of longest of the group. It arises at the level of the
the rings formed by the superficial flexor ten distal fourth of the antebrachium from the pal
dons. At their origin the sheaths enclose only the mar tendinous sheet of the lateral superficial
deep flexor tendons. Somewhat farther distally, belly of the humeral head of the m. flexor digi-
M u s i :l e s of the T h o r a c ic L im b 223
Olecranon- - O le c ra n o n -
B ic e p s -
— Flexo r c a rp i u ln a ris,
E x tens o r c a r p i - - u ln a r h ea d P ro n a to r te re s —
u ln a r is
P ro n a to r g u a d ra tu s —
Flexor d ig it o r u m —
-Flexor digitorum p ro f, ulna r head
s u p e r fic ia lis
Flexo r c a rp i u ln a ris,- F le x o r d ig it o r u m —
h u m e ro l h e a d p ro f, ra d io ! head
R a d iu s - ■
F le x o r c a r p i u ln a ris -
h u m e ra l h ea d F le x o r d ig it a r u m -
prof. h u m e ra l head
-Flexon ca rpi ra d ia lis
Tendon o F -
- C a r p a l Fa scia , F le x o r ca rp i radialis
cu t e d g e
A ccessory carpal
A b d u c to r d i g i t i -
g u m ti A b d u c t o r p a llic is-
-A b d u cto r p o llic is la n g us
Flexor d ig i t i q u in t i ~ b r e v is et O pponenspoll-
- - F le x o r d ig i forum
4 th in t e r o s j eous m. p r o f to I st d ig it
-L u m b n c o le s mm
A n n u la r lig am en ts'
P ro x im a l- ~
M id d le - -
D is t a l- - ■Flexor d ig i to ru m Fic. 3-62. Deep antebrachial muscles, caudomedial aspect.
proF. to 2 d ig it
torum profundus. As a slender, rounded muscle 3-67) are four in number. They lie on the palmar
belly, it runs to the carpal joint lying between side of the four large metacarpal bones at the
the digital flexors. It crosses under the palmar depth of the tendon branches of the flexor digi
carpal transverse ligament with the deep flexor torum profundus. They are relatively strong
tendon, and its thin tendon splits into two (or and border on one another. They arise from
three) branches at the middle of the metacar the proximal ends of metacarpals II, III, IV, and
pus. These accompany the branches of the m. V, and from the joint capsule, and cover the en
flexor digitorum superficialis for digits III and tire palmar surfaces of these metacarpal bones.
IV, and occasionally digit II, and fuse with them. After coursing a short distance, each muscle di
Gurlt (1859) homologizes this muscle with the vides into two branches, which attach by tendons
m. palmaris longus of man, whereas Ellenberger to the sesamoids of the respective metacarpo
and Baum (1943) designate it as the m. palmaris phalangeal joints. A portion of each tendon ex
longus accessorius. The name m. interflexorius is tends over the collateral borders of the joint and
derived from Agduhr (1915) and Pitzomo runs distally on the dorsal surface of the proxi
(1905). According to Kajava (1923), the muscle mal phalanx to unite with the tendon branch of
could be designated m. interflexorius profundo- the common extensor tendon.
sublimis. Morphologically each of these muscles results
Action: Flexion of the forepaw. from the fusion of two muscles. According to
Innervation: N. medianus. Kajava (1923), they represent the mm. flexores
The m. flexor digitorum brevis (Fig. 3-63), breves profundi, which are placed dorsal to the
the weakest of this group, is a delicate, only ramus palmaris profundus of the n. ulnaris and
slightly fleshy muscle, which arises distal to the are innervated by it. Each of the four muscles is
carpus from the palmarolateral surface of the collaterally covered by a considerable tendinous
superficial flexor tendon branch for digit V. It fascia which extends far distally. According to
goes into a flat tendon, which occasionally takes Forster (1916), each muscle is invaginated by a
the place of the whole muscle. It ends on the tendinous sheet which comes from the diverging
transverse ligament of the metacarpophalangeal portions and which extends proximally, causing
joint. the duplicity of each muscle to be much more
Gurlt (1859) compares this with the m. pal marked.
maris brevis of man; according to Kajava (1923), Action: Flexion of the metacarpophalangeal
however, one would have to regard this as a joints.
muscle mass in the carpal pad. Ellenberger and Innervation: Deep branch of the n. ulnaris.
Baum (1943) describe this muscle as the m. pal The special muscles of digit I. The rudimen
maris brevis accessorius. tary first, or medial, digit has three special mus
The mm. lumbricales (Figs. 3-61, 3-64) are cles: an outward rotator, a flexor, and an inward
three small muscles which are associated with rotator.
the tendons of the flexor digitorum profundus. The m. abductor pollicis brevis et opponens
The first muscle arises from the contiguous sides pollicis (Figs. 3-60, 3-63, 3-67) arises from a
of the flexor digitorum profundus tendons to the tendinous band which comes from the synovial
second and third digits; the second from the ten sheath of the superficial flexor tendon and goes
dons to the third and fourth digits; and the third to the sesamoid of the tendon of the m. abductor
from the tendons to the fourth and fifth digits. pollicis longus located on the medial side of the
They pass obliquely distally and laterally and carpus. It ends in the ligamentous tissue at the
end in thin tendons which are inserted on the metacarpophalangeal joint of digit I. According
proximal medial surfaces of the first phalanges to Kajava (1923), the muscle represents the ra
of the third, fourth, and fifth digits. The tendon dial head of the m. flexor pollicis brevis profun
of the first muscle inserts on the third digit, the dus.
second on the fourth, and the third on the fifth Action: Flexion of digit I.
(Leahy 1949). Innervation: Deep branch of the n. ulnaris.
Action: Flexion of the metacarpophalangeal The m. flexor pollicis brevis (Figs. 3-63, 3 -
joints. 67) is larger than the special abductor just de
Innervation: Deep branch of the n. ulnaris. scribed and lies between it and the m. adductor
The muscles lying on the palmar side of the pollicis. It arises on the palmar carpal ligament,
metacarpal bones. These include the interos runs obliquely to digit I, and ends on the sesa
seous muscles. moid bone or on the proximal phalanx.
The fleshy mm. interossei (Figs. 3-63, 3-64, (Text continued on page 230.)
M u sc les of the T h o r a c ic L im b 225
- In t e r f le x o r iu s tendon to
3 r° d i g i t , cut
Flexor diqitorum-
superf. -U lna
— In f e r f le x o r iu-S
- - F le x o r ca^p r a d ia lis
- F I e xo n d ig i f o r u m
p ro fu n d u s (c u t)
Accessory
c a r p a l bore A b d u c t o r p a i l i c i s b r e v is
et O ppo nens p o l l i c is
A b d u c t o r d i g i t i q u in f i-
- Flexor pollicis brev.s
- -A d d u c to r d ic jit i
A d d u c f o r d ig ih q u in ti -
Second/
In t e r o s s e o u s m.-
in te ro s s e o u s m
- ~ q fh m e t a c a r p a l bone
F l e x o r d ig ito ru m p ro fu n d u s te n d o n --
J ~'d in t e ro s s e c u s m
F le x o r diqi torum p ro fu n d u s-
tendon +o 3 rd J i a i t , c u t
I u m b r ic a lis —
- ~Ext ensor diqi t o r u m communi s
Flexun a c/itorum p ro fu n d u s - -
t e n d o n to j/.th d i g i t
Flexor d iq i t o r u m superf.- - ■
- Proximal dorsal sesamo, d bone
Flexor d 'C / it o r u m p r o f u n d u s -1 st p h a la n x
Dors e la s t i c l i g a m e n t
D i s i a ! annular hqament, c u t ------
- 2 nd p h a la n x
------ - 3 rd p h a l a n x
Fic. 3 -64. The fourth digit, medial aspect. (Proximal annular ligament removed.)
M u sc les of the T h o r a c ic L im b 227
cranial
,A cc e s s o r y cephal i c v.
Pr oxi ma l col lateral radi al a., med. branch i/ P r o x i ma l colla teral r a d i a l a., lat. branch
Superfi ci al r adi al n., med. branch^ _ * Superficial radial n., la t branch
Me di
-Subfascial bursa
Super f i ci al dig
Ulnar n!
cau dal
Fic.. 3-65. Schematic plan of cross section of forepaw through accessory carpal bone.
228 Chapter 3. M yo lo g y
Abductor digiti V-
Interossei
Extensor ca rp i ulnaris-
- -Abductor pollicis l ongus
Int enossei -
■Adductor pollicis
Deep d ig it a l f le x o r I V —
Action: Flexion of digit I. They lie palmarly on the ramus profundus of the
Innervation: Deep branch of the n. ulnaris. n. ulnaris and are innervated by it.
The m. adductor pollicis (Figs. 3-63, 3-67) is
the strongest muscle of digit I. It arises as a T he F a s c ia e o f t h e T h o r a c ic L im b
small, fleshy muscle body between the special
flexor and the m. interosseus of digit II on the The superficial and deep fasciae of the neck
palmar carpal ligament and ends on the lateral and thorax extend laterally over the shoulder
surface of the proximal phalanx of digit I. and there form the superficial and deep fasciae
The special muscles of digit V. The fifth of the shoulder and brachium. The whole limb
digit, like the first, also has three special mus is covered by this double system of connective
cles: an adductor, a flexor, and an abductor. tissue, the parts of which take their name from
The m. adductor digiti quinti (Figs. 3-63, 3 - the portion of the limb that they cover.
67) arises from the palmar carpal ligament and The superficial fascia on the shoulder and
extends as a slender belly obliquely in a lateral brachium covers these portions of the extremity
direction to end on the medial surface of meta as a bridge laterally from the superficial neck
carpal V and the proximal phalanx of digit V. and superficial trunk fasciae. In the distal bra
Its proximal portion lies on the mm. interossei chial region, however, it is completely closed
3 and 4 and its distal portion lies between mm. into a cylinder, since that portion of the super
interossei 4 and 5. ficial fascia in the region of the sternum goes
Action: Adduction of digit V. over to the medial brachial surface. Cranially
Innervation: Deep branch of the n. ulnaris. and laterally this covers the mm. brachiocephali-
The m. flexor digiti quinti (Figs. 3-6 3 ,3 -6 7 ) cus, deltoideus, and triceps brachii, on which it
arises on the ligament from the accessory carpal unites strongly with the deep leaf of the brachial
bone to metacarpal IV, runs obliquely over the fascia. It covers the m. pectoralis superficialis
m. interosseus of the fifth digit laterally, and by medially. The v. cephalica is covered laterally by
a thin tendon joins that of the m. abductor digiti the superficial fascia and, as it crosses the flexor
quinti. surface of the elbow joint, it lifts the fascia into
Action: Flexion of the fifth toe. a fold. Beyond the elbow joint the closely ap
Innervation: Deep branch of the n. ulnaris. plied superficial fascia is extremely delicate and
The strong m. abductor digiti quinti (Figs. 3 - less easily movable with respect to the deep tis
63, 3-67) arises on the accessory carpal bone. It sue. On practical grounds one also differentiates
ends by means of a tendon which unites with the special superficial fascia of the forearm, car
the m. flexor digiti quinti on the lateral sesamoid pus, metacarpus, and digits, under which the cu
and frequently by a thin tendon on the proximal taneous vessels and cutaneous nerves extend
phalanx of the fifth digit. It lies directly under over long distances before they actually enter
the skin on the palmar carpal ligament. On its the skin.
deep surface the fusiform body bears a delicate, The deep fascia of the lateral surface of the
proximal tendinous leaf. shoulder and brachium is called the fascia omo-
Action: Abduction of digit V. brachialis lateralis. From the deep fascia of the
Innervation: Deep branch of the n. ulnaris. neck it runs along the m. rhomboideus; the su
The special muscle of digit II. The second perficial leaf of the deep fascia of the back runs
digit has only one special muscle, an adductor. over the mm. latissimus dorsi and trapezius to
The m. adductor digiti secundi (Figs. 3 -6 3 ,3 - the lateral surface of the shoulder. Here it firmly
67) arises on the palmar carpal ligament between attaches to the spine of the scapula after it has
the m. interosseus 2 and the m. adductor digiti covered the mm. infraspinatus, supraspinatus,
quinti, runs distal (between the mm. interossei 2 deltoideus, and triceps brachii, or as much of
and 3), and ends by means of a tendon on the these muscles as make their appearance later
proximal end of the proximal phalanx of digit II. ally. It extends distally as far as the elbow joint,
Action: Adduction of digit II. attaching, between the mm. deltoideus and bra
Innervation: Deep branch of the n. ulnaris. chialis on one side and the m. cleidobrachialis on
Forster (1916), includes the adductors of the the other, to the crest of the major tubercle of
first, second, and fifth digits under the name, the humerus. Distally, where the m. brachialis is
mm. contrahentes digitorum, which in the dog free in the triangle between the mm. triceps
are represented separately but which, when bet brachii and brachiocephalicus, the fascia is espe
ter developed, constitute a “contrahentes leaf.” cially strong; here it extends under the cephalic
M u sc les of the P e l v ic L im b 231
vein and surrounds the muscle loosely with in- M U SC LES OF TH E PELVIC LIM B
termuscular septa under the m. brachiocephali-
cus. Farther medially it covers the m. biceps The M uscles of the Loin, Hip, and Thigh
brachii. The deep fascia is thinner and more
firmly attached to the long and lateral heads of The pelvis and thigh are covered on all sides
the m. triceps brachii than it is elsewhere. The by muscles which are for the most part common
lateral omobrachial fascia, including the origin to both of these body regions, so that the two
of the m. tensor fasciae antibrachii, passes over groups cannot be sharply differentiated. The
the m. cleidobrachialis into the fa s c ia omobra- so-called hip muscles act mostly on the hip joint,
chialis inedialis, which arises from the inner sur but a few act also on the sacroiliac joint. The
face of the mm. subscapularis and teres major, muscles of the thigh act primarily on the knee
and passes distally over the m. triceps brachii joint. The loin and hip muscles are divided into
and the medial surface of the humerus to merge three groups. The sublum bar or inner loin mus
into the deep antebrachial fascia at the elbow cles lie on the ventral surfaces of the lumbar ver
joint. The fa scia antebrachii covers the muscles tebrae and ilium: mm. psoas minor, iliopsoas,
of the forearm as a closely applied tube which is and quadratus lumborum. The rump muscles lie
thickest medially. Between the extensor and on the lateral side of the pelvis: mm. gluteus su
flexor muscles it sends septa to the periosteum perficialis, medius, and profundus, piriformis,
of the radius and ulna; these septa enclose the and tensor fasciae latae. The inner pelvic mus
individual muscles, as well as small groups of cles are in part inside the pelvis: mm. obturator
muscles. The fascia is intimately united with the internus, gemelli, and quadratus femoris.
extensors, more loosely covers the flexors, and The muscles of the thigh are designated as to
is firmly fused to all free portions of the bones of their cranial, caudal, and medial positions. To
the antebrachium. In the distal antebrachial re the caudal group belong the mm. biceps femoris,
gion it is intimately joined with the connective semitendinosus, and semimembranosus (some
tissue found between the tendons. In the groove times called the hamstring muscles). They form
between the tendons of the mm. extensor and a strong fleshy mass on the caudal side of the fe
flexor carpi ulnaris proximal to the accessory mur. The m. biceps femoris is lateral, the m.
carpal bone, the fascia invaginates more deeply. semitendinosus caudal, and the m. semimem
At the carpus it becomes the fa s c ia o f the fore branosus medial. To the cranial group belongs
paw (fascia manus), which, as the dorsal and pal the m. quadriceps femoris, which forms the
mar deep fascia, ensheathes all tendons and fleshy foundation of the cranial half of the thigh.
superficial muscles of the forepaw distally and A small m. capsularis and the m. sartorius crani-
attaches to all projecting parts of bones. Even alis are associated with it. The medial group in
the cushions of all of the pads are closely united cludes the mm. gracilis, pectineus, adductores,
with the deep fascia. From the cushion of the obturator externus, and sartorius caudalis.
carpal pad an especially strong band of the fas The sublumbar muscles. The sublumbar
cia extends directly laterally and a little proxi- muscles arise on the ventral surfaces of the cau
mally toward the distal end of the ulna; on the dal thoracic and lumbar vertebrae and insert on
medial side another such band, the palmar car the os coxae and femur; they lie on one another
pal transverse ligament, goes to the medial bor in several layers.
der of the carpus, bridging over the flexor ten The m. psoas minor (Figs. 3-68 to 3-71) runs
dons. On the dorsal surface of the carpus are toward the pelvis ventromedially as it lies be
found transverse supporting fibers. Finally, on tween the iliac fascia ahd peritoneum ventrally
the dorsal surface of the metacarpus, there is and the mm. iliopsoas and quadratus lumborum
formed a triangular, fibrous leaf which extends dorsally. Its muscular belly is weaker than that
from metacarpal I obliquely laterally and dis of the m. iliopsoas. It arises from the tendinous
tally to the branches of the common digital ex fascia of the m. quadratus lumborum at the level
tensor tendon for digits II, III, and IV. Palmarly, of the last thoracic vertebrae and on the ventral
on the metacarpophalangeal joints, thickened surface of the last thoracic vertebra and the first
portions of the deep fascia (transverse ligaments) four or five lumbar vertebrae; it is separated from
attach primarily to the sesamoid bones. its fellow of the opposite side by an interval
232 Chapter 3. M yology
gradually increasing caudally so that the bodies last three thoracic vertebrae to the transverse
of the lumbar vertebrae become visible. The processes of the lumbar vertebrae as far as the
strong, flat tendon fused with the iliac fascia seventh; it is covered by tendinous leaves dor
comes out of a shining, tendinous leaf at the fifth sally and ventrally. It ends on the medial sur
lumbar vertebra; it runs to the iliopectineal line face of the wing of the ilium between the articu
and inserts on this line as far as the iliopectineal lar surface and the caudal ventral iliac spine.
eminence. The lateral portion of this muscle overhangs the
Action: To steepen the pelvis, or to flex the transverse processes of the lumbar vertebrae, so
lumbar part of the vertebral column. that it also comes to lie on the ventral surface of
Innervation: Lateral branches of the rami ven the tendon of origin of the m. transversus ab
trales of lumbar nerves 1 to 4 or 5. dominis. *
The m. iliopsoas (Figs. 3-69, 3-73) repre Action: Fixation of the lumbar vertebral col
sents a fusion of the m. psoas major and the umn.
m. iliacus. It lies ventral to the m. quadratus Innervation: Rami of the ventral branches of
lumborum and dorsal to the m. psoas minor, the lumbar nerves.
which it covers laterally and also medially cau The rump muscles. The rump muscles ex
dal to its point of transition into tendon. It is tend between the ilium and the thigh; they are
narrow and tendinous at its origin on the trans arranged in several layers.
verse processes of lumbar vertebrae 2 and 3, The m. tensor fasciae latae (Figs. 3-70,3-72)
where it lies lateral to the m. quadratus lumbo is a triangular muscle which attaches proximally
rum. It also attaches by means of the ventral to the tuber coxae. With three more or less dis
aponeurosis of this muscle on lumbar vertebrae tinct slips, it radiates distally and caudally over
3 and 4, and, finally, on the ventral and lateral the m. quadriceps femoris. It arises (partly super
surfaces of lumbar vertebrae 4 to 7. After this ficially and partly deeply) from the aponeurosis
portion of the m. iliopsoas passes over the ilium of the m. gluteus medius and deeply from the
as the m. psoas major it receives the m. iliacus tuber coxae. Laterally it covers the origin of the
from the smooth ventral surface of the ilium be caudal belly of the m. sartorius. Distally it con
tween the iliopectineal line and the lateral bor tinues into the deep leaf of the fascia lata on a
der of the ilium. The two muscle masses (m. horizontal line running cranially from the tro
psoas major and m. iliacus) composing the m. chanter major. Over this deep aponeurotic leaf
iliopsoas can be easily isolated. The m. iliopsoas runs the m. biceps femoris and a superficial leaf
attaches to the trochanter minor of the femur of this same fascia. Since this fascia runs over the
between the m. rectus femoris, laterally, and the m. quadriceps femoris to the patella, this muscle
m. pectineus, medially. extends the stifle.
Action: To draw the pelvic limb forward by Action: Tension of the fascia lata, and thus
flexion of the hip joint; when the femur is flexion of the hip joint; extension of the
fixed in position, flexion and fixation of the stifle joint.
vertebral column; when the leg is extended Innervation: N. gluteus cranialis.
backward, it draws the trunk backward. The m. gluteus superficialis (Figs. 3-72, 3-73,
Innervation: Branches of the rami ventrales of 3-75), the most superficial gluteal muscle, is a
the lumbar nerves. rather small, flat, almost rectangular muscle. It
The m. quadratus lumborum (Figs. 3-68, 3 - extends between the sacrum and first coccygeal
69, 3-71) is the most dorsal of the sublumbar vertebra proximally and the trochanter major
muscles. It lies directly on the bodies of the last distally. The gluteal fascia covers this muscle
three thoracic and all the lumbar vertebrae, as loosely; it fuses with the muscle more intimately
well as under the proximal portions of the last only at the proximal two-thirds of its cranial
two ribs and the transverse processes of the portion. The muscle arises from the gluteal
lumbar vertebrae. It is covered ventrally from fascia and thereby from the tuber sacrale of the
the first lumbar vertebra by the m. psoas minor ilium; neighboring portions come from the coc
and also, from the fourth lumbar vertebra, by cygeal fascia. The thick caudal portions come
the m. psoas major. It has a thoracic and a lum from the lateral part of the sacrum, from the first
bar portion. The thoracic portion of this muscle, coccygeal vertebra, and from more than half of
which is rather strong in the dog, consists of in the proximal part of the sacrotuberous ligament.
completely isolated bundles which become ten Its fibers converge distally and laterally and be
dinous; these bundles extend more or less dis come tendinous; the tendon runs over the tro
tinctly caudolaterally from the bodies of the chanter major and inserts on the weak trochanter
M u s c le s o f th e P e lv ic L im b 233
- - Internal intercostal
Transversus abd:
cut
D iaph ra g m
A o rtic h iatus-
--X II rib
R etractor costae--
Qu a d n a t u s lumborum
Transversus a b d o m i n i s -------
I nt er v e r t e br a I f i b r o -
cartilage between I V
and V lum bar
Psoas m a jo r
Quadratus lumborum
Iliocostalis lumborum
Psoas minor
Ps o a s minor, cut
Psoas major
M iddle glu te a l
-------- L e s s e n t r o c h a n t e r of
External obturator fem ur
'\ jjo r r u
Q u a d ra tu s
lu m b o ru m
Psoas m in o r
-Q u a d ra tu s lu m b o ru m
-P soas mo or
Tendon o f P j o a s m i n o r
I
-I lia c u s
I
M u sc les of th e P e l v ic L im b 235
tertius. This tendon fuses with the aponeurosis The strong, fan-shaped m. gluteus profundus
of the m. tensor fasciae latae. The m. gluteus (Figs. 3-70, 3-74, 3-78, 3-86) is the deepest of
superficialis covers portions of the mm. gluteus the gluteal muscles. It is completely covered by
medius and piriformis, and also the sacrotuber the mm. gluteus medius and piriformis; at the
ous ligament. In large dogs it is 5 to 7 cm. wide same time it extends, for a considerable distance,
and more than 1 cm. thick caudally. A thin, deep caudal to the deep portion of the gluteus medius.
portion was seen by Ziegler (1934), lying under It takes its origin laterally from the shaft of the
the caudal border of the muscle and having a ilium near the ischiatic spine. Its fibers converge
special origin on the sacrotuberous ligament. over the hip joint distolaterally and form a short,
Beneath its terminal tendon on the trochanter strong tendon which ends cranially on the tro
major, in about one-third of the specimens, chanter major distal to the insertion of the m.
there is a synovial bursa approximately 1 cm. gluteus medius. Underneath the tendon of inser
wide. tion, there is often a small synovial bursa.
Action: Extension of the hip joint. Action: Extension of the hip joint, with some
Innervation: N. gluteus caudalis. abduction.
The strong m. gluteus medius (Figs. 3 -7 0 ,3 - Innervation: N. gluteus cranialis.
72, 3-73, 3-76) lies on the gluteal surface of the The inner pelvic muscles. The so-called
ilium, from which it takes its principal origin. It “small pelvic association” includes a group of
also arises from the iliac crest and from both short muscles which lie caudal to the m. gluteus
angles (tuberosities) of the ilium. Some fibers profundus and the hip joint. They extend from
also come from the dorsal sacroiliac ligament the inner and outer surfaces of the ischium to
and from the deep surface of the gluteal fascia, the femur. These are the mm. obturator internus,
which is fused with the muscle caudal to the the gemelli, and the quadratus femoris.
iliac crest. A large part of the muscle lies under The m. obturator internus (Figs. 3-71, 3-73,
the gluteal fascia and skin, and only caudally is 3-78) is the strongest of this group. As a fan
it covered by the superficial gluteal muscle. In a shaped muscle it covers the obturator foramen
craniodistal direction it extends over the m. internally. It arises medial to the foramen on the
gluteus profundus and ends by a short, strong pelvic surfaces of the rami of the pubis and
tendon on the free end of the trochanter major. ischium, and from the ischiatic arch. Its fibers
The muscle is 2.5 to 3.5 cm. thick and 7 to 9 cm. converge laterally and, extending under the
wide. From the caudal border of the m. gluteus sacrotuberous ligament, are drawn over the
medius there is split off a narrow but strong, lesser sciatic notch in a distinct, gliding surface
deep belly; this does not, however, correspond directly behind the ischiatic spine. At the same
to the m. gluteus accessorius of other animals. It time the muscle turns almost at right angles, dis
arises on the transverse processes of the last tolaterally, and forms a strong, flat tendon which
sacral and first coccygeal vertebrae and on the is embedded deeply between the edges of the
sacrotuberous ligament, and ends on the tro broader mm. gemelli which lie beneath. The
chanter major by a narrow, strong tendon which overhanging portions of the mm. gemelli, bear
sinks into the principal tendon. Only a small, ing a shining distal tendinous sheet, run from the
proximal portion of the caudal border of the edges of the lesser sciatic notch into the obtura
muscle can be seen under the principal muscle. tor tendon, so that the triple tendinous appara
Action: Extension of the hip joint. tus ends undivided in the trochanteric fossa.
Innervation: N. gluteus cranialis. Caudally the tendon of the m. obturator externus
The flat m. piriformis (Figs. 3-73, 3-76, 3 - accompanies it. Where the muscle glides over
86), which, in large dogs, is 2 to 2.5 cm. wide the ramus of the ischium, there is a very thin-
and 1 to 1.25 mm. thick, joins the m. gluteus walled synovial bursa, 1.5 to 2 cm. wide, which
medius caudally and is completely covered by surrounds the edges of the tendon and may ex
the m. gluteus superficialis. It arises with this tend out on the underlying mm. gemelli. A sec
muscle on the transverse process of the last ond bursa, 2 to 3 mm. wide, lies under the mm.
sacral vertebra and on the sacrotuberous liga obturator and gemelli tendons in the trochan
ment. It runs to the outer surface of the tro teric fossa between the trochanter major and the
chanter major where its narrow, flat tendon ends joint capsule.
on the weak trochanter tertius, with the tendon Action: Outward rotation of the hip joint.
of the m. gluteus superficialis, which covers it. Innervation: N. ischiadicus.
Action: Extension of the hip joint. The mm. gemelli (Figs. 3-70, 3-73, 3-78)
Innervation: N. gluteus caudalis. represent a muscle which has been formed by
236 Chapter 3. M yology
the fusion of two parts and which lies between femoris. The deep head, covered by the super
the terminal portions of the mm. obturator ficial portion, arises under the medial tendinous
internus and obturator externus caudal to the m. origin of the latter from the ventral side of the
gluteus profundus behind the hip joint. It over lateral angle of the ischiatic tuberosity by a long,
hangs the tendon of the m. obturator internus strong tendon. The slender muscle, at the mid
cranially and caudally. The mm. gemelli arise dle third of the femur, passes over the lateral
together on the outer surface of the ramus of the surface of the thigh at the caudal edge of the
ischium in the arch under the gliding surface for principal portion; here it broadens to unite with
the m. obturator internus and are covered super the middle head as the caudal branch of the m.
ficially by a shining fascia which forms the floor biceps femoris, at the level of the popliteal space.
of the groove for the obturator tendon. Alto The m. abductor cruris caudalis runs along its
gether this tendon apparatus ends undivided in free edge. In the region of the femorotibial joint
the trochanteric fossa. the entire m. biceps femoris, with slightly di
Action: Outward rotation of the hip joint. verging fibers, runs on the lateral surface of the
Innervation: N. ischiadicus. vastus lateralis and the gastrocnemius group,
The short, fleshy m. quadratus femoris (Figs. and, as an aponeurosis, radiates into the fascia
3-70, 3-73, 3-76) arises on the ventral surface of lata and the fascia cruris. By means of this apo
the ischium medial to the lateral angle of the neurosis the cranial head, by fusing with the
ischial tuberosity. It is surrounded by the origins covering of the m. quadriceps femoris, attaches
of the mm. adductor magnus et brevis, semi to the patella and the straight patellar ligament
membranosus, semitendinosus, biceps femoris, and by this to the tibial tuberosity. The middle
and obturator externus. Medial to the m. biceps and caudal branches radiate into the fascia lata
femoris, it extends in an almost sagittal direction and crural fascia as these enclose all of the
cranially; it bends slightly laterally and runs cranial and lateral muscles of the thigh and crus
distally to reach the distal portion of the tro to insert on the tibial crest. The deep surface of
chanteric fossa between the mm. obturator ex the muscle is covered by a distinct perimysium.
ternus and adductor magnus et brevis. It ends The covering appears strengthened distally as a
immediately above the trochanter tertius. distinct tendon. Where the muscle passes over
Action: Extension and outward rotation of the the deep surface of the m. abductor cruris cau
hip joint. dalis (which accompanies the caudal border of
Innervation: N. ischiadicus. the m. biceps femoris) the tendon reaches a
The caudal muscles of the thigh. The cau width of 5 mm. It lies under the abductor and,
dal thigh muscles are grouped about the ischial by means of the crural fascia, runs distally along
tuberosity and some of them run to the lateral the m. gastrocnemius. It curves in front of the
side of the thigh—the mm. biceps femoris and main part of the calcanean tendon to end on the
abductor cruris caudalis; others run to the medial medial tuberosity of the tuber calcanei after it
side of the thigh—the mm. semitendinosus and has united with a thick, tendinous strand com
semimembranosus. ing from the mm. semitendinosus and gracilis.
The m. biceps femoris (Figs. 3-70, 3-72, 3 - The calcan ean tendon (tendo calcaneus), also
82, 3-83, 3-87) is a large, long muscle, lying in known as A chilles’ tendon, consists of all those
the lateral part of the buttock and thigh, and ex structures attaching to the point of the heel, or
tending from the region of the ischial tuberosity the tuber calcanei of the fibular tarsal bone. The
to the middle of the tibia. It arises by two un tendons of the mm. flexor digitorum superficialis
equal heads, a cranial, superficial one, and a cau and gastrocnemius are its main components,
dal, much smaller, deep one. The principal although the mm. biceps femoris, semitendino
superficial head arises from the ventrocaudal sus, and gracilis also contribute to its formation.
end of the sacrotuberous ligament and partly From the proximal end of the strand which aids
from the lateral angle of the ischial tuberosity, in forming the calcanean tendon, fibers pass to
where it is firmly united with the m. semitendi the medial lip of the femur. Heavy, connective
nosus in an intermuscular leaf. The fibers of this tissue fibers of the interfascicular septa of the m.
muscle run nearly parallel. New fiber bundles, biceps femoris enter the tendon and, without
which arise from a strong, tendinous leaf on the the aid of muscle fiber attachment, fasten it
deep surface of the muscle, are added distally. A intimately so that a single functional unit results.
portion of these laterally located fibers can be Action: Extension of the hip, stifle, and tarsal
differentiated from the main mass of the cranial joints. The caudal part of the muscle raises
head as the middle branch of the m. biceps the crus when the extremity is not bearing
M u s c le s o f th e P e lv ic L im b 237
tM i d d ! e cjluteal
238 Chapter 3. M yology
weight; this part, therefore, flexes the stifle such, attach to the medial surface of the tibia in
at these times. front of the flexor muscles. Separate strands of
Innervation: Cranial part: ramus distalis of the both tendons, however, extend in the fascia in
n. gluteus caudalis. Middle and caudal an arc cranially and upward toward the rough
parts: ramus muscularis proximalis of the n. surface on the distal end of the tibial crest. From
tibialis (Skoda 1908, Ziegler 1934, Nickel, the caudal edge of the tendon of the m. semi
Schummer, and Sieferle 1954). This double tendinosus (close to its origin) a tendon is sepa
innervation supports the theory that the m. rated which unites with a distinct, strong strand
biceps femoris of the dog is a m. gluteobi- from the tendon of the m. gracilis. The conjoined
ceps; its cranial belly is to be regarded as tendons extend on the medial surface of the m.
split off from the m. gluteus superficialis. gastrocnemius medialis to the calcanean tendon
The straplike m. abductor cruris caudalis and thus to the tuber calcanei. It is related to the
(Figs. 3-72, 3-86, 3-87), 10 mm. wide and 1 corresponding tendon of the m. biceps femoris.
mm. thick, lies under the caudal edge of the m. The strands are attached to each other caudal to
biceps femoris. It arises by a long, flat tendon, the main part of the calcanean tendon and the
which lies on the aponeurosis of the deep sur digital flexor tendons by the fascial bridge, which
face of the m. biceps femoris on the ventrocaudal distally becomes progressively thicker.
edge of the sacrotuberous ligament near the Between the proximal and the middle third of
ischial tuberosity. It extends under both heads of the semitendinosus, and visible on the free sur
the m. biceps femoris on the lateral surface of face, there is a delicate, but complete, tendinous
the mm. quadratus femoris, adductor, and semi inscription. This intercalation is probably the re
membranosus. At the level of the popliteal space, mains of the tendon of the m. caudofemoralis,
it appears superficially between the mm. biceps which in lower mammals divides the semi
femoris and semitendinosus and, keeping close tendinosus transversely. This muscle has dis
to the edge of the m. biceps femoris, crosses the appeared but its tendon remains as the inscrip
shank in an arc and goes into the crural fascia, tion; the proximal portion of the muscle is to be
where it often extends beyond the end of the m. regarded as a division of the m. gluteus super
biceps femoris to the digital extensors. Because ficialis. The two portions of the m. semitendino
of its different innervation, this muscle cannot be sus are provided with individual nerves.
looked upon as a division of the m. biceps fem Action: Extension of the hip, stifle, and tarsal
oris. joints; flexion of the stifle joint in the free
Action: With the caudal branch of the m. bi non-weight-bearing limb.
ceps femoris, it abducts the limb. Innervation: Ramus muscularis proximalis of
Innervation: N. ischiadicus. the n. tibialis (special branches for the part
The m. semitendinosus (Figs. 3 -7 0 ,3 -7 2 ,3 - found proximal to and the part found distal
82, 3-87) is 2.5 to 3.5 cm. thick, and has four to the tendinous inscription).
edges in cross section. It lies in the caudal part The m. semimembranosus (Figs. 3-70, 3-73,
of the thigh between the cranial and lateral parts 3-80, 3-85, 3-87) is entirely fleshy, has parallel
of the m. biceps femoris and the medial and fibers, and is oval to triangular in cross section.
cranial parts of the m. semimembranosus. It ex It crosses the m. semitendinosus medially and
tends in an arc between the ischial tuberosity lies between the mm. biceps femoris and semi
and the proximal segment of the shank, where it tendinosus laterally, and the mm. adductor and
forms a large part of the caudal contour of the gracilis medially. It arises caudal and medial to
thigh. Its distal end diverges from the m. biceps the m. semitendinosus on the lower surface of
femoris to the medial side of the m. gastroc the rough portion of the tuber ischiadicum. It
nemius. The m. semitendinosus arises on the soon splits into two equally strong bellies which
caudal and ventrolateral parts of the lateral extend in a slight arc to the medial side of the
angle of the ischiatic tuberosity between the stifle joint. The cranial belly, which is 3 to 3.5
mm. biceps femoris and semimembranosus. It cm. thick, joins the m. adductor magnus et
extends distally at the caudal edge of the m. bi brevis and ends in a short, flat tendon on the
ceps femoris and diverges from it at the popliteal aponeurosis of origin of the m. gastrocnemius. It
space to the medial side of the shank, where it attaches also to the distal end of the medial lip of
follows the m. semimembranosus. By means of a the femur which runs distally on the shaft as a
strong, flat tendon it passes under the aponeuro rough line to the medial condyle. The caudal
sis of the m. gracilis. Both of these aponeuroses belly partly covers the cranial belly from the
represent portions of the crural fascia and, as lateral side. It goes into a somewhat longer, nar-
Gl ut eus s u p e r f i c i a l i s
G l u t e u s medius
- S a r t o r i us
- Tensor fa s c ia e la ta e
-B icep s fe m o ris
- Sem i m e m b r a n o s us
Sem i tendinasus
A bductor crur-s c a u d a l is
- - G lu te u s m e d iu s
- - Gluteus s u p e r f i c ialis
--T e n s o r fosciae la ta e
- J e m t m em b r o n o s u s
-S em itendinosus
Removed
Biceps
S a rto riu s
Q u a d ra tu j fe m o ris
Gem e l l I
239
240 Chapter 3. M yology
rower tendon, which extends more distally than The m. vastus medialis, 4 to 5 cm. wide and
the m. semitendinosus over the m. gastrocnemius up to 2.5 cm. thick, arises in the proximal fifth
medialis to end under the tibial collateral liga of the femur on the intertrochanteric crest cra
ment of the femorotibial joint on the margin of nially. It arises medially on the line for the vas
the medial condyle of the tibia. tus medialis and, somewhat farther distally, on
Action: Extension of the hip and stifle joints the proximal portion of the medial lip. It is lat
with the stifle adducted. erally compressed and, with portions of the vas
Innervation: Ramus muscularis proximalis of tus intermedius, covers the distal portion of the
the n. tibialis. rectus medially. Laterally and cranially it bears
The cranial muscles of the thigh. The cra strong distal tendinous leaves; medially it bears
nial thigh muscles extend between the pelvis a strong proximal one. Between these two tendi
and the femur proximally and the patella and nous systems the muscle fibers extend rather ob
tibial tuberosity distally. The four subdivisions liquely. Only above the patella do its tendinous
of the m. quadriceps femoris form the bulk of elements fuse with those of the rectus femoris.
the group. To this is added the insignificant mus The muscle ends on the patella, covered by the
cle of the joint capsule of the hip, the m. capsu- cranial belly of the m. tensor fasciae latae, and
laris, and the m. articularis genus proximal to the its principal portions encroach upon the mm.
stifle joint. sartorius and pectineus. The vastus intermedius
The strong m. quadriceps femoris (Figs. 3-72 sends many fibers into the tendinous leaf.
to 3-75, 3-79, 3-80, 3-82, 3-87) covers the fe The stronger m. vastus lateralis arises on the
mur cranially, laterally, and medially. Distally it craniolateral part of the proximal fifth of the fe
forms a tendon which includes the patella within mur on the transverse line and on the line of the
it and ends on the tibial tuberosity as the straight vastus lateralis to the lateral lip. It covers the
ligament of the patella; it fuses with the fascia rectus femoris laterally to some extent. Later
lata and thereby with the aponeurosis of the ally, at the origin, it bears a strong tendinous leaf
mm. biceps femoris and sartorius. The quadri from which the majority of the muscle fibers ex
ceps muscle consists of the rectus femoris (cra tend obliquely medially and forward. The upper
nial), vastus lateralis (lateral), vastus medialis fibers go to the rectus femoris and radiate into
(medial), and sometimes a fourth belly, the vas its lateral aponeurotic covering as far as the ter
tus intermedius, directly underneath the rectus minal tendon. The vastus lateralis is inseparably
femoris and covering the cranial surface of the united with the rectus femoris and its terminal
femur. tendon except proximally. The caudal fibers of
The m. rectus fem oris, 2 to 3 cm. thick in large the muscle are more longitudinal in direction;
dogs, is round in cross section proximally and is they extend from the lateral lip rather directly
laterally compressed distally; it is enclosed be to the stifle joint where their tendinous portion
tween the vasti in such a way that it overhangs unites intimately with the joint capsule. There
them somewhat cranially. It arises by a short, are firm connections with the fascia lata and
strong tendon from the iliopubic eminence of with a strong aponeurotic leaf which pushes be
the ilium, and appears between the m. sartorius tween the vastus lateralis and the m. biceps fem
and the m. tensor fasciae latae. Covered crani oris to attach to the bone.
ally and medially by the cranial belly of the m. The m. vastus intermedius is the weakest por
tensor fasciae latae, it extends between the vas tion of the quadriceps. It arises with the vastus
tus lateralis and vastus medialis to the patella, lateralis, which covers it, and also from the lat
which is included in its strong tendon as a sesa eral part of the proximal fourth of the femur. Ly
moid bone. This tendon continues distally as the ing under the rectus, directly on the femur, its
straight ligament of the patella, over the stifle terminal tendinous leaf radiates into the m. vas
joint, to insert on the tibial tuberosity. Proximal tus medialis.
to and at the sides of the patella, islands of carti Bursae. Under the tendon of origin of the m.
lage are found buried in the tendon. (For details, rectus femoris, a small, synovial bursa is occa
see the description of the stifle joint in Chapter sionally found. There is almost constantly a
2.) The tendons of the vastus lateralis, vastus bursa (0.5 to 1.5 cm. in diameter) between the
medialis, and the m. tensor fasciae latae fuse in distal third of the muscle and the femur. A small
timately with the patellar portion of the rectus (0.5 cm.) bursa is usually present under the ter
tendon. Each collateral surface of the rectus minal tendon of the vastus medialis and the vas
bears a strong, tendinous leaf; the lateral surface tus lateralis. In addition, the patellar joint cap
takes on tendinous strands from the vastus later sule has a considerable proximal out-pocketing
alis. under the tendon of the quadriceps.
Gemel l i , Int. o b t u r a t o r
•t Ext . o b t u r a t o r
M iddle gluteal
Pi n i f o r m i - s - l -
<S u p e r f i c i a l g l u t e a l -
11 i o p s o a s
Quadratus fem oris--
Vastus lateralis- - *■ — / -Add uctor longus
l/astu s m ed ialis
A dductor - - ■
m a g n u s et b r e v i s - Pectin eus
■ ™ Se mi m e m b ra n o s u s
Gastrocnemius, - - 4 I
. , , . I * 4 -G astrocnem ius,
la te r a l head f I ,. , ,
r \ , medial head
S u p e r f d i g i t a l fl.~
Popliteus
- Mi ddl e g l u t e a l
-Deep g l u t e a l
-Superf. g l u t e a l
-Capsularis coxae
— Vastus la te ralis
A r t i cu l a r i s g e n u s
-G astrocnem iu s
F abe llae
Popliteus
^ — Long. dig. e x t
F ig . 3 -7 4 . Left femur, showing areas of muscle attach F ig . 3-75. Left femur, showing areas of muscle attach
ment, cranial aspect. ment, lateral aspect.
241
2 42 Chapter 3. M yology
Action o f the m. quadriceps: Extension of the tae it also bounds the muscle laterally. The cra
stifle joint, tension of the fascia cruris. nial belly of the m. sartorius is visible proximally
Innervation: N. femoralis. on the lateral aspect of the thigh in front of the
The m. capsularis coxae (Figs. 3-70, 3 -7 4 ,3 - m. tensor fasciae latae. From the cranial border
77, 3-81) is a small, spindle-shaped muscle 2 to it slowly turns to the medial surface of the thigh,
4 cm. long in the region of the hip joint. It is to pass into the medial femoral fascia immedi
placed laterally and caudally to the rectus fem ately above the patella. There is a firm union
oris, and is covered laterally by the cranial bor with the tendon of the rectus femoris and the
der of the m. gluteus profundus. It arises with vastus medialis.
the rectus femoris on the iliopubic eminence The caudal belly lies close beside the cranial
and extends cranially and laterally over the cap belly, entirely on the medial surface of the thigh.
sule of the hip joint to the neck of the femur, It arises on the bony ridge between the two ven
where it attaches to the common ridge between tral spines of the ilium, between the m. iliopsoas
the lateral and the medial vastus muscles. and the lumbar portion of the m. iliocostalis on
Action: Flexion of the hip joint. the one side, and the mm. tensor fasciae latae
Innervation: N. femoralis. and gluteus medius on the other. It runs over the
The m. articularis genus (Fig. 3-74) is a small, medial surfaces of the vastus medialis and stifle
short muscle which arises from the cranial sur joint, and forms an aponeurosis which blends
face of the femur just proximal to the trochlea with that of the m. gracilis. Radiating into the
for the patella. It inserts on the tibial tuberosity. crural fascia, it ends on the tibial crest.
It is separated from the main portion of the m. Action: To flex the hip; to advance and adduct
quadriceps femoris by a delicate intermuscular the free thigh.
septum. It is closely related to the proximal Innervation: N. femoralis (cranial branch) for
pouch of the stifle joint capsule as it courses dis both bellies.
tally. The m. gracilis (Figs. 3-70, 3-79, 3-82,3-87),
Action: Extension of the stifle joint and pos in the dog, forms an extensive, broad muscular
sibly tension of the proximal pouch of the sheet which is found in the superficial layer of
stifle joint capsule. the caudal portion of the inner surface of the
Innervation: N. femoralis. thigh. Caudally the muscle becomes rather
The medial muscles of the thigh. The ad thick, and here it can be seen to a small extent
ductors are a strong group of muscles on the me from the lateral aspect. By means of its broad
dial side of the thigh. Proximally, the long belly aponeurosis, which covers the medial surface
of the m. tensor fasciae latae lies on the medial of the m. adductor magnus et brevis and also
side; distally, the m. semitendinosus takes part the median subpubic tendon, the muscle arises
in the caudal border of the medial surface of the from the pelvic symphysis. The subpubic ten
thigh. The adductors are arranged in superficial don is a thick, flat tendon below the symphysis
and deep layers. The superficial group includes pelvis in the region of origin of the bilateral ad
the mm. sartorius and gracilis; the deep group is ductors. Its line of origin presents a distally con
represented by the mm. pectineus, adductor, vex arc which passes from the pecten ossis pubis
and obturator externus. Between the m. sarto to the ischial arch. This is about 4 cm. in its
rius and the m. gracilis a broad gap exists proxi symphyseal length and 2 cm. in its maximal
mally. In its depth the mm. adductor and pectin width. Tendinous fibers extend from one side of
eus can be seen caudally; the rectus femoris this median, unpaired tendinous plate to the
and vastus medialis cranially. Superficially, this other. The aponeurosis of origin of the m. gracilis
gap is closed by the medial femoral fascia. from the subpubic tendon is wider cranially than
The m. sartorius (Figs. 3-70, 3-79, 3 -8 2 ,3 - it is caudally. The m. gracilis passes over the m.
86, 3-87) is a long, flat muscle which extends in adductor magnus et brevis as well as both bellies
two, peculiar, straplike strands, each 3 to 4 cm. of the m. semimembranosus. At approximately
wide on the cranial contour of the thigh from the edge of this latter muscle it goes into a strong,
the region of the tuber coxae to the medial sur flat tendon which pushes under the m. sartorius,
face of the stifle joint. passes over the popliteal space with the m. gas
The cranial belly arises on the iliac crest and trocnemius medialis and the end of the m. semi
the cranial ventral iliac spine, as well as from the tendinosus, to end along the entire length of the
lumbodorsal fascia. Along with the caudal belly tibial crest. This end aponeurosis also spreads
it bounds the m. quadriceps femoris cranially out into the crural fascia, and from its caudal
and medially, and with the m. tensor fasciae la (Text continued on page 247.)
M u sc les of the P e l v ic L im b 243
G lu te u s medius,
p a r s caudal is
Pi r i fo r m is
S acro tu b ero u s
lig a m e n t
„ 'G lu te u s p r o f u n d u s
G em ellu s
G lu te u s
G luteus p ro f
m ed iu s - O b t u r a t o r in i
te n d o n
Glu t e u s Gem e ll us
p ro fu n d u s
' Q u a d r a t u s fe m o ris
'O b tu ra to r e x te rn u s
- G lu t e u s m e d /u s
uraton
e xte nnus
--G lu te u s p ro fu n d u s
-Caps u I a r i s Coxae
Head o f fe m u r
-A cd uc+ o r lo n g u e
--Q u o d ra tu s fe m o r is
■ — O b tu ra to r ex tern us
Fic. 3 77. Muscles of the hip joint.
A. Ventral aspect.
B. Obturator extemus, lateral aspect.
G lu te u s p r o fu n d u s \
Tendon o f O b t u r a t o r e x t .
Tendon o f O b t u r a t o r in t.'
Gem e l l i '
O bturator in te rn u S '
„ S a rto riu s , c ra n ia l p o r t
, Rectus f e m o r is
, Head o f f e m u r
, Vastus m e d ia lis
- - T e n s o r fasciae
la ta e
A d d u c to r
G r a c i I is - - Pe c t i n e u s
- F o s c ia l a t a
Semi tendinosus
- Va-stus medialis
-A d d u c to r
rnagnus er brevis
S e m ite n d in o s u s
G a s tr o c n e m iu s ■
-P o p lite u s
Rectus femoris -
- A d d u c to r
V a s tu s -
l a te r a lis
Semimembranosus
'R e ctu s f e m o r i s
-V o s fu s
la t e ra I is
Vostus m e d ia li s - - R ectus
fe m o ris
■Vastus
la te ra lis
Vastus i n t e r m e d i u s --
border it sends a well-developed reinforcing extends cranial to the m. obturator externus and
band to the calcanean tendon of the semitendi inserts on the lateral lip of the femur near the
nosus. According to Frandson and Davis (1955), third trochanter. Its tendon covers the end of
the caudal part of the gracilis, the part which at the m. quadratus femoris.
taches to the tuber calcis after joining the cal The m. adductor magnus et brevis arises along
canean tendon of the semitendinosus, may rup the entire pelvic symphysis and on the neigh
ture in racing dogs. Their findings suggest that boring parts of the ischiatic arch, but primarily
the caudal part of the gracilis is an important ex from the subpubic tendon and pelvic symphysis.
tensor of the tarsus in the racing greyhound. The strong muscle belly of the large portion of
Action: Adduction of the thigh, extension of the adductor extends obliquely under the m.
the hip joint. gracilis and accompanies the distal portion of
Innervation: N. obturatorius. the m. pectineus. Under the m. sartorius it
The m. pectineus (Figs. 3-70, 3-73, 3 -7 9 ,3 - presses so closely against the vastus medialis that
81, 3-87) belongs to the deeper group of adduc the m. pectineus with its tendon is compressed
tors, although in front of the m. gracilis it en to a thin leaf. Here the m. adductor magnus et
croaches in part upon the medial fascia of the brevis receives a glistening tendinous leaf which
thigh. It is closely applied to the m. adductor unites firmly with the tendon of the m. pectin
magnus et brevis cranially, but is separated from eus. The muscle ends over the whole length of
the m. sartorius by the femoral fascia and ves the lateral lip, from the trochanter tertius to a
sels. Its spindle-shaped body is circular in cross place near the origin of the m. gastrocnemius
section and may reach a thickness of 2 or 3 cm. laterale. Its tendon, with that of the m. pectin
It arises tendinously on the prepubic tendon and eus, blends with the periosteum of the popliteal
from the abdominal muscles which sink into this surface of the femur. Laterally the muscle is cov
tendon; it also has a fleshy origin on the iliopec ered by the m. biceps femoris. It extends be
tineal eminence. Distally the muscle becomes tween the m. quadratus femoris and the m.
flatter after it has passed under the m. sartorius. semimembranosus to the caudal surface of the
It fills the narrow space between the vastus me femur. From the broad muscle body of the ad
dialis and the adductor in such a way that its ductor magnus et brevis a less distinct portion
wide and long tendon coming from tendinous can regularly be separated.
sheets on two surfaces of the muscle passes over Action: Adduction and flexion of the hip joint.
to the caudal surface of the femur after turning Innervation: N. obturatorius.
obliquely. The medial border of the tendon is The m. obturator externus (Figs. 3-70, 3-73,
thickened and ends on the raised ridge of the 3 -77) is a fan-shaped muscle which covers the
medial femoral condyle in common with the obturator foramen externally. It arises on the
cranial belly of the m. semimembranosus. The ventral surface of the pelvis adjacent to the
thinner, principal portion of the tendon goes pubic symphysis. It is separated from the ischial
into the periosteum on the popliteal surface of symphysis by the mm. adductores. On some of
the femur medial to the insertion of the m. ad its pennate parts, the muscle body bears deli
ductor magnus et brevis. cate, tendinous strands from the origin of the
Action: Adduction of the thigh, outward rota muscle. The fibers converging toward the tro
tion of the stifle joint. chanteric fossa appear under the ramus of the
Innervation: N. obturatorius. ischium and form a strong tendon. The m. quad
The mm. adductores (Figs. 3 -7 0 ,3 -7 3 , 3-79, ratus femoris, which takes a more sagittal course,
3-81, 3-87) form the caudal part of the deep is crossed on its deep surface in such a way that
group of muscles next to the m. pectineus. In a small part of the obturator externus appears
the dog they are represented by two separable between the mm. quadratus femoris and gemelli
muscles: the small fusiform m. adductor longus, laterally. It ends between these in the trochan
and the much stronger m. adductor magnus et teric fossa.
brevis. Both extend distally and laterally from Action: To rotate the limb outward.
the pelvic symphysis to the femur. It is uncer Innervation: N. obturatorius.
tain if there actually is a homology between this
muscle and the similarly named muscle of man.
Femoral Triangle and Associated Structures
The m. adductor longus encroaches upon the
proximal part of the m. pectineus. It is com
pletely covered by the other part of the adduc The opening caudal to the abdominal wall for
tor. From its origin on the pubic tubercle, it the passage of the iliopsoas muscle and its con-
248 Chapter 3. M y o lo g y
/l i o p s o a s
Adductor lonqus
- - ■/--A d d u c t o r l o n g u s
A dductor' m a q n u s '
et b r e v i s '
C a p s u ia n s c o x a e
Q u a d ra tu s f e m o r i s
- A d d u c t o r m a g n us et brevis
tained femoral nerve is known as the muscular whereas the medial surface is essentially left
lacuna (lacuna muscularis). It is bounded later free. Flexor and extensor groups are not sep
ally and caudally by the os coxae, medially by arated on the crus as they are on the antebra
the rectus abdominis, and cranially by the ilio chium. Cranially and laterally are found exten
p ectin eal arch (arcus iliopectineus). This arch, sors of the digital joints and flexors of the tarsus;
composed of blended iliac and transversalis fas caudally lie flexors of the digital joints and exten
cia, separates the muscular lacuna from the vas sors of the tarsus. These functional muscle
cular lacuna. groups are mixed on the crus because the tarsal
The vascular lacuna (lacuna vasorum) lies joint is set at an angle opposite to that of the dig
craniomedial to the muscular lacuna, separated ital joints. The tarsal joint has its flexor surface
from it by the iliopectineal arch. It is bounded dorsally, whereas each of the digital joints has its
cranially by the caudal muscular border of the extensor surface dorsally; therefore, the muscles
internal abdominal oblique and the inguinal lig lying over the dorsal surface must be flexors of
ament and medially by the rectus abdominis. It the tarsus and extensors of the digital joints.
is separated from the superficial inguinal ring, The craniolateral muscles of the crus. The
which lies only a few millimeters craniolateral flexors of the tarsal joint which lie on the cranio
to it, by the inguinal ligament. It contains the lateral side of the crus are the mm. tibialis crani
femoral artery and vein and the saphenous alis, peroneus (fibularis) longus, extensor digito
nerve. The transversalis fascia which surrounds rum longus, extensor digitorum lateralis, and
the femoral vessels as they pass through the vas extensor hallucis longus. They are separated
cular lacuna forms the fem oral sheath. The fun- topographically into two groups according to the
nel-shaped femoral sheath of man is divided into predominating muscles, not according to their
three compartments: a lateral one for the artery, function. One is the long digital extensor group;
an intermediate one for the vein, and a medial the other is the peroneal group. The former in
one partly occupied by lymph vessels and fat. It cludes the mm. tibialis cranialis, extensor digi
is into this medial compartment, known as the torum longus, and extensor hallucis longus,
fem oral canal in man, that a hernia may occur. which lie for the most part on the cranial aspect
The base of the femoral canal, in the vascular la of the tibia. The fibular group includes the mm.
cuna, is the fem oral ring. It is closed by a con peroneus longus, peroneus brevis, and extensor
densation of extraperitoneal connective tissue digitorum lateralis, which lie on the lateral part
and covered internally by parietal peritoneum. of the crural skeleton.
The femoral canal in the dog is considered by The m. tibialis cranialis (Figs. 3-83, 3-86, 3 -
some authors to include the sheath and its con 88, 3-89, 3-94) is a superficial, strong, some
tents (lymph vessels, femoral artery and vein, what flattened muscle lying on the cranial sur
and saphenous nerve). face of the crural skeleton. It arises medial to the
The fem oral triangle (trigonum femorale) is sulcus muscularis on the cranial portion of the
the shallow triangular space through which the articular margin of the tibia and on the laterally
femoral vessels run to and from the leg. It is arched edge of the tibial crest. From its origin
bounded cranially by the thin caudal belly of the the muscle, which is about 3 cm. wide, passes
m. sartorius and caudally by the m. pectineus. over the craniomedial surface of the crus, and
Its floor or lateral boundary is formed proximally near its distal third becomes a thin, flat tendon.
for a short distance by the m. iliopsoas. The m. This tendon extends obliquely over the tarsus to
pectineus and m. vastus medialis complete this the rudiment of metatarsal I, which is very often
boundary. Superficially, the femoral artery and fused with the first tarsal bone and to the proxi
vein and the lymphatics are covered by the me mal end of metatarsal II. The threadlike tendon
dial femoral fascia and skin. Because of the su of the m. extensor hallucis longus encroaches
perficial position of the artery in the femoral closely upon the lateral edge of this tendon
triangle it is a favorable site for taking the pulse throughout its extent as far as the metatarsus,
of the dog. both turning toward the medial edge of the tar
sus. It becomes increasingly removed from the
long digital extensor, as this passes to the meta
The Muscles of the Crus (True Leg, Shank, tarsus lateral to the axis of the foot. On the distal
or Gaskin) part of the tibia all three tendons are bridged
over by the broad proximal transverse ligament
On the crus the muscles lie on the cranial, lat which extends obliquely upward in a medial di
eral, and caudal surfaces of the crural skeleton, rection. The long digital extensor tendon passes
250 Chapter 3. M y o lo g y
. -Biceps f e m o r i s
.-Gran. t i b i a l
BicepsJ
■-Fi buiani s
Q ua dr i c e ps f emon i s-
lonc/us
Santa rius-
■Latdicj. ext.
Gracilis—
Flex, hal luci s
Ionc/us
Semitendinosus- - -
-Ext. hallucis
l ongus
F ig. 3-82. Left tibia and fibula, showing areas of muscle F ig. 3-83. Left tibia and fibula, showing areas of muscle
attachment, cranial aspect. attachment, lateral aspect.
M u sc les of th e P e l v ic L im b 251
under the distal transverse ligament as it crosses joint capsule. The tendon of origin of the long
the tarsus. This is a collagenous loop coming digital extensor is underlaid on its deep surface
from the fibular tarsal bone. The end of the ten by a pouch (3 to 4 cm. long) of the meniscotibial
don often shows variations which frequently are portion of the stifle joint capsule. This is the so-
associated with variations in the m. extensor called capsular synovial bursa. Since this bursa
hallucis longus (Grau 1932). The fascia is thick extends slightly around the caudal border of the
ened into a strand on the medial side of the mus tendon, it is also spoken of as a synovial sheath.
cle which reaches from the tibial crest to the The bundle of the four tendons of the long digital
proximal transverse ligament and farther, a extensor is surrounded by a synovial sheath
strand continues to the proximal end of meta which extends from the beginning of the tendon
tarsal III. In its entirety, this strand may, per at the proximal transverse ligament to the distal
haps, be comparable to the m. peroneus tertius border of the tarsus, where the branches diverge.
of the horse. The terminal tendons of the mm. Accordingly, the sheath passes under both trans
tibialis cranialis and extensor hallucis longus are, verse ligaments; the mesotendon is sent out from
according to Walter (1908), surrounded by a the lateral surface.
synovial sheath between the proximal transverse Action: Extension of the digits; flexion of the
ligament and the middle of the tarsus. tarsus.
Action: Rotation of the hindpaw in a lateral di Innervation: N. peroneus.
rection, extension of digit II. The m. extensor hallucis longus (Figs. 3-82,
Innervation: N. peroneus. 3-88, 3-89, 3-92) is a delicate muscle band,
The m. extensor digitorum longus (Figs. 3- elliptical in cross section, and covered by the
75, 3-88, 3-89, 3-92), spindle-shaped, and at mm. extensor digitorum longus and peroneus
most 2 to 2.5 cm. thick, lies in the group of digital longus. It lies directly on the tibia. It arises on
extensors on the tibia between the m. tibialis the cranial border of the fibula between the
cranialis and the m. peroneus longus. Proximally proximal and the middle third, and on the inter
it is covered by the m. peroneus longus; distally osseous membrane cranial to the m. peroneus
it lies free medial and caudal to the m. tibialis brevis. The muscle extends obliquely distomedi
cranialis. It arises in the extensor fossa on the ally under the long digital extensor, on the m.
lateral epicondyle of the femur and passes peroneus brevis and then on the tibia. It is ac
through the sulcus muscularis of the tibia. The companied by the a. and v. tibialis cranialis. Near
muscle belly bears a strong distal, cranial, tendi the distal fourth of the crus it goes into a fine
nous leaf toward which all fibers converge; near tendinous strand which continues between the
the tarsus it goes into its terminal tendon. This long extensor tendon, laterally, and the tendon
tendon runs along the tendon of the m. tibialis of the m. tibialis cranialis, medially. The tendi
cranialis and that of the m. extensor hallucis nous strand of the m. extensor hallucis longus
longus, laterally. At the distal fourth of the crus passes over the dorsal surfaces of the tarsus and
it is held in place by the proximal transverse lig metatarsal II to the metatarsophalangeal joint of
ament. Toward the tarsus it diverges from the the second digit, where it is lost in the fascia
other tendons and at the bend of the tarsus it is beside the corresponding branch of the long ex
fastened by a second weaker, transverse liga tensor tendon. It occasionally broadens into the
ment. At the same time the extensor tendon, aponeurosis of the metatarsus, and, exception
from the very beginning, is split into four ally, it ends on the rudiment of the hallux (first
branches; these extend distally along the dorsal digit). If the first digit is present, the muscle may
surfaces of the metatarsal bones and digits II, give off a tendinous branch to it. In about one
III, IV, and V. Each ends on a distal phalanx third of all specimens the muscle shows varia
after it has received the collateral branches of tions (see Grau 1932), which are often associated
the m. interosseus just proximal to the proximal with variations in the m. tibialis cranialis. Often
digital joint. As in the pectoral limb, the dorsal this muscle is stronger in smaller breeds than it
elastic ligaments attach with them on the distal is in larger ones. Occasionally the muscle is
phalanx. A broadened portion of the tendon of fused completely with the long digital extensor.
the m. extensor digitorum lateralis unites with Action: Extension of digit II; extension of digit
the branch for the fifth digit on the proximal I also if it attaches to it.
phalanx. At the level of the proximal digital joint Innervation: N. peroneus.
there is embedded in each branch a small, dorsal The short-bellied m. peroneus [fibularis]
sesamoid element; the corresponding sesamoid longus (Figs. 3-83, 3-88 to 3-91, 3-94) is the
of the metatarsophalangeal joint is located in the principal representative and the most superficial
2 52 Chapter 3. M yology
muscle of the fibular group. It lies in the proxi obliquely on their deep sides by the tendon of
mal half on the lateral surface of the crus, be the lateral digital extensor. It unites with the
tween the m. tibialis cranialis and the m. flexor long digital extensor and the m. interosseus of
hallucis longus. It covers the proximal portions the proximal phalanx of the fifth digit. The ten
of the m. extensor digitorum lateralis and the m. don of the lateral digital extensor is enclosed in
peroneus brevis. In large dogs it may be 2 cm. a synovial sheath, having a medial mesotendon
thick. It arises on the lateral condyle of the tibia, in common with that of the m. peroneus brevis.
the fibular collateral ligament of the femorotibial In large dogs this begins 1.5 to 2.5 cm. above the
joint, and the proximal end of the fibula. Near lateral malleolus, reaches almost to the middle of
the middle of the tibia it becomes an elliptical the tarsus, and almost always communicates
tendon which is enclosed in a tough fascial mass with the capsule of the talocrural joint.
with the tendons of the mm. extensor digitorum Action: Extension and abduction of digit V.
lateralis and peroneus brevis. From the lateral Innervation: N. peroneus.
surface of the tarsus it runs over the sulcus of the The m. peroneus [fibularis] brevis (Figs. 3 -
lateral malleolus. Running distally, cranial to the 83, 3-89, 3-91, 3-94) is the deepest muscle of
fibular collateral ligament, it crosses the tendon the fibular group. It first appears distally under
of the m. extensor digitorum lateralis and the m. the lateral digital extensor between the m. pero
peroneus brevis superficially; then, making use neus longus and the m. flexor hallucis longus. It
of the groove in the fourth tarsal, it passes in a arises from the lateral surface of the distal two-
sharp curve underneath the tendinous m. abduc thirds of the fibula and tibia, almost as far as the
tor digiti quinti to the plantar surface of the me lateral malleolus. Here the muscle becomes
tatarsus, which it crosses transversely. It ends on completely tendinous, although the relatively
the proximal end of metatarsals I, II, and V, and strong tendon begins far proximad as a narrow
on the rudiment of the first digit opposite the in strong leaf. After crossing under the long fibular
sertion of the m. tibialis cranialis. The synovial collateral ligament of the tarsus and the tendon
sheath of the m. peroneus longus begins 3 to 4 of the m. peroneus longus, it runs further be
cm. above the lateral malleolus and is provided tween the m. peroneus longus and the tendon of
with a mesotendon on its deep surface. It the lateral digital extensor, to attach to the proxi
reaches approximately the end of the tarsus and mal end of metatarsal V. The tendon of the
may be divided. Beneath the plantar end of the peroneus brevis is included in a common syno
tendon there is a synovial bursa which com vial sheath with the tendon of the m. extensor
municates with the joint capsule between the digitorum lateralis. An insignificant bursa lies
third and fourth tarsals. under the tendon at its insertion.
Action: Rotation of the hindpaw so that the Action: Flexion of the tarsal joint.
plantar surface faces laterally. Innervation: N. peroneus.
Innervation: N. peroneus. The caudal muscles of the crus. On the cau
The m. extensor digitorum lateralis (Figs. 3 - dal side of the crus lie the extensor of the tarsal
82, 3 -8 8 to 3-92) is the weakest muscle (1 cm. joint—the m. gastrocnemius; the flexors of the
wide; 2 to 3 cm. thick) in the fibular group. It digital joints—the mm. flexor digitorum super
lies between the m. peroneus longus and the m. ficialis and profundus; and the flexor of the knee
flexor hallucis longus on the m. peroneus brevis joint—the m. popliteus. The m. gastrocnemius is
and fibula. It arises on the proximal third of the superficial and largely encloses the m. flexor
fibula. Superficially it has a strong distal, tendi digitorum superficialis. It covers the deep, digit
nous covering which, at the middle of the crus, al flexor group consisting of the mm. flexor digi
becomes a thin tendon. This tendon lies between torum profundus, tibialis caudalis, and popliteus.
the cranially located tendon of the m. peroneus The m. gastrocnemius (Figs. 3-73, 3-86, 3-
longus and the caudally located tendon of the 88 to 3-91, 3-94) is divided into a lateral and a
m. peroneus brevis, as these cross the proximal medial head covered by strong tendinous leaves
lateral surface of the tarsus. As a rule the tendon and infiltrated by tendinous strands. The lateral
of the lateral digital extensor lies in front of that h ead (caput laterale) arises by a large tendon on
of the m. peroneus brevis. Thus, the caudal the lateral plantar tuberosity of the femur. The
groove of the lateral malleolus is crossed in m edial head (caput mediale) arises on the me
common by these two tendons. Both the long dial plantar tuberosity.
fibular collateral ligament of the tarsal joint and Each tendon of origin has a prominent sesa
the tendon of the m. peroneus longus are crossed moid bone, the lateral and the medial fabella,
M u sc les of the P e l v ic L im b 253
respectively; these are cartilage covered on the row of tarsal bones, forming four branches, each
side toward the femur, and there articulate with of which, in large dogs, is 5 mm. wide. These ex
the corresponding condyle on a small, flat area. tend distally over metatarsals II, III, IV, and V.
The two heads of the m. gastrocnemius almost At the metatarsophalangeal joints, the branches
completely enclose the m. flexor digitorum are enclosed, in common with the corresponding
superficialis; they fuse with each other distally, branches of the deep flexor tendon, in a trans
forming a flat (in large dogs 5 to 6 cm. wide) verse ligament. Here they form the cylinders, as
muscle, the duplicity of which is accentuated on the thoracic limb, for the passage of the
distally by a middle tendinous plate. After cross tendons of the deep digital flexor. The synovial
ing the superficial flexor tendon laterally, the apparatus and the transverse ligaments corre
tendon of the m. gastrocnemius attains its deep spond with those of the thoracic limb. Under
surface and ends on the tuber calcanei. Proxi neath the tendon on the tuber calcanei, and
mally the muscle is covered laterally by the m. extending proximally and distally from the tuber,
biceps femoris and medially by the mm. semi there is an extensive synovial bursa, the bursa
tendinosus, semimembranosus, and gracilis. calcan ei. Its proximal portion lies between the
Distally it lies under the fascia and skin. Beneath superficial digital flexor tendon and the tendon
the m. gastrocnemius, directly on the tibia and of the m. gastrocnemius; its distal portion lies
fibula, are located the m. popliteus and the heads between the superficial flexor tendon and the
of the m. flexor digitorum profundus. plantar ligament.
The calcanean tendon (tendo calcaneus The portion of the muscle on the crus, as far
[Achillis]) is the aggregate of those structures as the tuber calcanei, corresponds to the m.
which attach to the tuber calcanei. The tendon plantaris of man, and its pedal portion corre
of the m. gastrocnemius, the main component, is sponds to the m. flexor digitorum brevis of man.
crossed medially by that of the superficial digital Action: Flexion of the digits; extension and fix
flexor, which first lies cranial to the m. gastroc ation of the tarsus; flexion of the stifle joint.
nemius but attains its caudal surface at the Innervation: N. tibialis.
tuber calcanei. Joining these two tendons are The strong m. flexor digitorum profundus lies
those of the mm. biceps femoris and semitendi on the caudal surface of the tibia, covered by the
nosus. m. gastrocnemius and the superficial digital
Action: Extension of the tarsal joint, flexion of flexor. It consists of the large, laterally located
the stifle joint. m. flexor hallucis longus, located between the
Innervation: N. tibialis. fibular group and the gastrocnemius group, and
The m. flexor digitorum superficialis (Figs. the weaker, medially located m. flexor digitorum
3-73, 3-88 to 3-91), infiltrated by tendinous longus. The latter muscle, short and spindle-
strands, lies on the mm. flexor digitorum pro shaped, lies on the medial side, behind the tibia,
fundus and popliteus. It is enclosed to a great between the m. popliteus and the m. flexor hal
extent by the heads of the m. gastrocnemius and lucis longus. In hoofed animals the m. tibialis
is compressed to an angular body which flattens caudalis, whose tendon joins that of the m. flexor
distally and becomes broader. A small portion of hallucis longus, constitutes a third head of the
the proximal segment and a narrow portion of deep digital flexor muscle. The caudal tibial mus
the distal border appear beyond the m. gastroc cle is independent of the two heads of the deep
nemius and thus encroach upon the fascia. The digital flexor in carnivores.
muscle is multipennate, because of the numer The m. flexor hallucis longus (Figs. 3-84, 3 -
ous tendinous folds which course through it. 88 to 3-91, 3-93), laterally located and large,
Proximally, it is firmly united with the lateral arises from the caudal surface of the proximal
head of the m. gastrocnemius. It arises with the three-fifths of the fibula, the proximal caudo-
gastrocnemius on the popliteal surface of the lateral border of the tibia, and along the whole
femur and on the lateral fabella. At the middle length of the popliteal line. It also arises from
of the tibia the tendon of the m. flexor digitorum the interosseous membrane. It has many tendi
superficialis winds medially around the tendon nous strands, resulting in formation of a multi
of the m. gastrocnemius to gain its caudal sur pennate muscle. In large dogs it is 2.5 to 3 cm.
face. On the tuber calcanei it broadens like a cap wide and 1 to 1.5 cm, thick. Caudally it is
and inserts collaterally on the tuber calcanei, covered by a strong tendinous leaf from which
united with the crural fascia and the calcanean the principal tendon arises and runs distally.
tendon of the mm. biceps femoris and semiten At the level of the middle row of tarsal bones
dinosus. The tendon divides twice, at the distal it fuses with the weaker tendon of the m.
(Text continued on page 262.)
254 Chapter 3. M y o lo g y
--Sartorius
Caudal tib ial- Popl i f a u s
--G racilis
F ib u la ris b re v is --
F ig . 3-84. Left tibia and fibula, showing areas of muscle F ig . 3-85. Left tibia and fibula, showing areas of muscle
attachment, caudal aspect. attachment, medial aspect.
M u sc les of the P e l v ic L im b 255
C o cc ygeus -
Dorsal la t. sacrococcycjeo I
P i r i f o r mis
In te r tra n sve rsa riu s
Levatorani
Ventral lat S a c r o c o c c j g e 01
Deep g l u t e a l '
S a c ro t u b erous lig a m e n t
M id. g l u t e a l -
E x te rn a l anal sp h in cte r
T endon o f - S ciatic tu b e ro s ity
Int. o b t u r a t o r
V astus l o t e r a l i s ------
— S em itendinosus
Sem im em branosus
— G a s tro cn e m iu s
Cra n tib ia I—
Flexor hallucis
L o n cj d ig ita l e xte n so r--
longus
Tendon of Fi b u l a r i s longus-
Tendon of Superf . d ig i f a I f l e x o r -
T e n d o n o f L a t . d i g i t a l extensor-
Fi bu la-
E x te n s o r h a llu c is lo n c j u s ^ ^
■T ib i a
B ic e p s f e m o r i s - -
F ib u ia n is lo n g u s - - - - P o p li f e u s
'T e n d o n o f
E x t d g . lon g.
-T ib ia lis
c r a n ia l is -L a te ra l c o ll a t e r a l l i g a m e n t
-Tendon o f E xt. d i g i t , lo n g u s
- G a s t r o c n e m iu s
F i b u l o r i s lon qu s
-fle x o r h a llu c is lo n g u s
E x te n s o r d ig ito ru m - f l e x o r d i g it , s u p e r f .
lo n g u s
T i b ia lis -
c r a n ia lis - - Tendon o f Fib. long.
E x te n s o r d ig ito ru m - E x t d i g i t lateral.
lo n g u s
- Fi b u I a n ts
b re v is
proxi m ol-
Transverse hgg.
F ib u la ris longus
d is ta l
-fib u la r is b r e v is
xt d ig it la te ro l.
•lexor hallucis
lo n g u s
-Ext. d ig ito r u m
-F ib u la ris lo n g u s
la te ra lis
--F le x o r h a llu c is
E x te n s o r d i^ it. la te ra lis
-F ib u la
F ib u la ris b re v is E x t e n s o r h a llu c is
lo n g u s
- - T ib ia
- P o p lite u s
Tibialis c o u d o lis
-F le x o r d ig ito ru m
lo n g u s
- F l e x o r h a llu c is
lo n g u s
I- - -Ex t e n s o r d i g i t o r u m
la te ra l^ te n d o n
- -F ib u I a r is b r e vis
- T ib ia lis c a u d a l is
-Gastrocnemius
Medial collateral
lig a m e n t
- P o p h te u s
- .F le x o r d ig i to r u m
lo n g u s
_ F le x o r d ig it o r u m
s u p e rfic ia lis
F l e x o r h a l l u c s lon gu s
E x t dig. lo n g
D
G a s tr o c n e m iu s ,
l a t e r a l head
F ib u la r is longus
Te n d o n of
gastrocnemius
-i-E ax ti dig. l a t
- - Flexor d i g i t su pe rf
Tendo c a lca n e u s
- - A b d u c to r d ig it: g u m t i
- Q ua dratus p la n ta e
Flexon d ig ito r u m -F le x o r d i g i t p r o f
lo n q u s
Flexor d iq i torum 5 th i n t e r o s s e o u s m
s u p e r f i c i a l i-s
Flexor d ig itoru m
p ro fu n d u s
C
Fic. 3 -91. Muscles of left crus and hindpaw.
A. Superficial muscles, plantar aspect.
B. Deep muscles of crus, plantar aspect.
C. Deep muscles, lateral aspect.
260 Chapter 3. M y o lo g y
brezis
- - F l e x o r d iq prof.
F i e x o r h a l l u c i s longus
Flexor dig superf. ( c u t)
F le x o r d i q lo n q u s L u m b r ic a le s
- F le x o r d iq s u p e r f
Q u a d r a t u s p la n t a e to 2 nd d i g i t
A b d u c to r d g i t i (Quinti
F le x o r d ig it p r o f
-Pna interosseous m
Lumbricales-
- F l e x o r d ig superf.
to 3 rd d i g i t
"■ in terosseous m.
2 nd i n terosseous m
flexor digitorum longus to form the deep flexor 88, 3-90) is a strong, triangular muscle lying in
tendon. Becoming wider and flatter, it divides the popliteal space. It covers the joint capsule
into four branches at the middle of the metatar and the medial half of the proximal third of the
sus; these behave as do the tendons of the m. tibia. It is covered caudally by the mm. gastroc
flexor digitorum profundus of the thoracic limb. nemius and flexor digitorum superficialis; it en
In the region of the extensor surface of the tar croaches medially upon the m. semitendinosus
sus, and proximal to it, a synovial sheath is and fascia. The popliteus arises on the plantar
formed, which communicates with the capsule surface of the lateral condyle of the femur by a
of the talocrural joint; its mesotendon passes to long tendon which contains a sesamoid bone.
the caudal surface of the tendon. This tendon invaginates the joint capsule and
The weak m. flexor digitorum longus (Figs. 3 - crosses under the fibular collateral ligament of the
84, 3-88, 3-90, 3-93) is a short, flat muscle lying femorotibial joint. The triangular muscle extends
medial to the m. flexor hallucis longus and lateral obliquely medially over the popliteal space to
to the m. popliteus. At its origin it is narrow; it the medial border of the tibia and ends on its
arises on the head of the fibula, the popliteal proximal third, as well as on the popliteal line.
line, and the fascial leaf separating it from the m. Action: Flexion of the stifle joint; inward rota
flexor hallucis longus. Above the middle of the tion of the leg.
tibia it becomes a fine tendon which runs along Innervation: N. tibialis.
the caudomedial border of the tibia with the
even finer tendon of the m. tibialis caudalis. Both Muscles of the Hindpaw
tendons pass through the groove of the medial
malleolus, with the tendon of the tibialis caudalis The muscles of the dorsal surface of the hind
lying cranial to that of the m. flexor digitorum paw. Only one muscle, the m. extensor digito
longus. In this position they accompany the rum brevis, is found on the dorsal surface of the
tibial collateral ligament of the tarsus to the level hindpaw.
of the tibial carpal, where the tendon of the m. The weak m. extensor digitorum brevis
flexor digitorum longus unites with the tendon (Figs. 3-92, 3-94) is a flat muscle, 2.5 to
of the m. flexor hallucis longus to form the deep 3 cm. wide, on the dorsum of the foot; it lies on
flexor tendon. The synovial sheath of the muscle the distal row of tarsal bones and on the meta
begins 1 to 1.5 cm. above the medial malleolus tarsal bones. It is covered by the tendons of the
and extends on the tendon to its union with the m. extensor digitorum longus, the fascia, and
main tendon. A variable mesotendon passes to skin. It consists of three heads, of which the
the tendon, laterally. middle is the longest. The heads arise from the
Action: Flexion of the digits and stifle joint; distal part of the fibular tarsal bone and on the
extension of the tarsus. ligamentous masses on the flexor surface of the
Innervation: N. tibialis. tarsus. Of the three terminal tendons, the lateral
The m. tibialis caudalis (Figs. 3 -8 4 ,3 -8 8 ,3 - one goes to digit IV, the intermediate one to
90), deep and medially placed, is completely digits III and IV, and the medial one to digits II
separated in the dog from the heads of the flexor and III. Only the branch going to digit II radi
digitorum profundus, in contrast to the condi ates directly into the corresponding branch of
tion in the hoofed animals. As an insignificant the long digital extensor. All others unite on the
spindle-shaped muscle, it lies between the m. proximal phalanx with the tendinous branches of
popliteus and the m. flexor hallucis longus. It is the mm. interossei. The latter go to the dorsum of
covered by the m. flexor digitorum longus and the paw and thereby unite secondarily with the
lies directly on the caudal surface of the tibia. It digital extensor tendons.
arises on the medial part of the proximal end of Action: Extension of the digits.
the fibula and soon forms a very delicate tendon Innervation. N. peroneus.
which extends distally in front of the somewhat The muscles of the plantar surface of the
stronger tendon of the m. flexor digitorum hindpaw. The muscles of the plantar surface of
longus. It ends, as in man, on the medial liga the hindpaw, the mm. interossei, adductor digiti
mentous masses of the tarsus. II, adductor digiti V, and lumbricales, behave as
Exceptionally, the muscle may be lacking. do the corresponding ones in the thoracic limb.
Action: Extension of the tarsus; outward rota The mm; abductor digiti V and interflexorii,
tion of the foot. which are united with the suspensory ligament
Innervation: N. tibialis. of the metatarsal pad, are modified. When the
The m. popliteus (Figs. 3-75, 3-84, 3 -8 5 ,3 - first digit (dewclaw) is lacking, its muscles are
M u sc les of the P e l v ic L im b 263
fascia passes over both the caudal and cranial the mm. adductor and semimembranosus after
contours of the thigh, to its medial surface. The going beneath the branches of the m. sartorius
lateral femoral fascia and the m edial fem oral and after bridging the femoral canal. Into this
fa s c ia thus join to form a cylinder on the thigh. fascia the m. gracilis sinks proximally; however,
The fascia lata passes from the m. biceps femoris this muscle, like the caudal belly of the m. sar
cranially and covers the individual portions of torius in particular, is also covered superficially
the m. quadriceps femoris toward the medial by a thin leaf of the deep medial femoral fascia,
surface as far as the vastus medialis. The medial which unites with the deeper lamella over its
fascia of the thigh reaches the medial surfaces of caudal edge. Distally, on the medial surface of
M u sc les of the P e l v ic L im b 265
the thigh, the a. and v. saphena and the n. Agduhr, E. 1915. Anatomische, statische und experimentelle
saphenus are included between the two leaves. Untersuchungen iiber N. medianus and N. ulnaris, bes.
deren motorisches Innervationsgebiet im Vorderarm von
The fascia lata is attached to the lateral lip of the Haustieren, nebst einigen Bemerkungen iiber die Musku-
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Baumeier, M. 1908. Zur vergleichenden Anatomie und Mor-
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tinuation of the lateral and medial femoral fas
83.
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82-121.
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Le Gros, Clark, W. E. 1946. An experimental study of the re Gliedmassen des Hundes. Dissertation, Dresden-Leipzig.
generation of mammalian striped muscle. J. Anat. 80:24- Winckler, G. 1939. Contribution a l’etude de la morphogenese
36. du muscle spinalis dorsi. Arch. Anat. Hist. Embr. 27:99.
Lockhart, R. D., and W. Brandt. 1938. Length of striated mus Ziegler, H. 1929. Muskelvarietaten bei Haustieren. Ztschr.
cle fibers. J. Anat. 72:470. Anat. u. Entw. 97:442-451.
Maximenko, A. 1929, 1930. Material zum Studium der Mm. Ziegler, H. 1931. Die Innervationsverhaltnisse der Becken-
serrati dorsales der Saugetiere. Ztschr. Anat. u. Entw. 89: muskeln bei Haustieren im Vergleich mit denjenigen
156-170, 92:151-177. beim Menschen. Morph. Jahrb. 68:1-45.
Miller, M. E. 1958. Guide to the Dissection of the Dog. 3rd Ziegler, H. 1934. Weitere Untersuchungen iiber den M.
Ed. Ithaca, N.Y. Pub. by author. glutaeobiceps von Hund (Canis familiaris) und Katze
Nickel, R., A. Schummer, and E. Seiferle. 1954. Lehrbuch der (Felis catus dom.) Morph. Jahrb. 73:385-391.
Anatomie der Haustiere. Band 1: Bewegungsapparat, Zimmermann, A. 1927. Uber die Quermuskeln der Rippen.
Berlin, Paul Parey. Allattani Kozlemenyek 24:53-60.
Pancrazi, G. 1928. Intorno al “foramen venae cavae” del dia- Zimmermann, A. 1928. Zur vergleichenden Anatomie des M.
framma dei mammiferi. Atti Soc. Nat. Modena 7:191- pronator teres. Verhdlg. Anat. Ges. 66:281-282.
192.
CHAPTER 4
TH E H EA RT A N D A RTERIES
St ernum, V segmenf
- P a r i e t a l serous peri ca r d i um
P a r ie ta l pleura
- - Fibrous pericardium
R. luncj, c a r d i a c l obe- -
- Pericard. mediastinal p l e u r a
Pulmonary pleura— ■- - H e a r t
i a p h n a g m a t i c lobe —
. p u l m o n a r y l i gament— s - j U -----D i a p h r a g m a t i c l obe
I I I ------ A o r t a
Esophagus- '
A zlj^ os v. - -
VI t h o r a c i c vertebra
Fic. 4-1. Cross section of body through the sixth thoracic vertebra.
H eart 269
Esophagus
Trachea
Bra c h i oc ep h a l ic t r u n k
A p i c a l lobe, r i g h t lung
L. s u b c l a v i a n a.
A p i c a l l obe, l e f t lung
- P u l mo n a r y t r u n k
Transverse s i n u s
Azygos v. -
- C a r d i a c l obe
C a r d i a c l a b e ~ 7'
- L . p u l m o n a r y vv.
R. p ul mo nar y vv.
Diaphragmatic lobe
Azygos lo be x x Esophagus
F ig . 4 -2 . Dorsal wall o f the pericardial sac, with adjacent structures, ventral aspect.
270 Chapter 4. The H eart and A rte rie s
R ig h t a u r ic le
L ig o m e n t u m a r t a r i o s u m , \ , A o rt i c a r c h
f ' R. p u lm o n a r y a.
L e f t pu I m o n o r y a -
- Precova
L e ft a u r ic le
—AZljijOS v.
L e f t p u l mono ry
C i r c u m f le * b r a n c h of
p u ! m em o ry VI/.
l e f t c o r o n a r y a.
G r e a t c o r o n a r y v .- - -R a t r i u m
P o s tc a v o
L e f t v e n tr ic le —
M iddle c a rd ia c v
O b liq u e y o f I a t r i u m
D o r s a l i n t e r v e n t n c u l a r br, o f
C o ro n a ry s in u s , ccronory Q
L. p r o x i m o I a t r i a l a.t , L. a t r i u r n
L. a u r i c l e , !
L. p u l m o n a r y vv.
L. pu I m o n o r y a Ob I i q u e v. o f l e f t a t r i u m
L. s u b c l a v i a n a.
, ' L e f t d i s t a l a t r i a l a.
A o r t ic a rch -
--- G r e a t c o r o n a r y v.
B ra c h ioc ep h a l ic - -
tru n k
- - C ir c u m f le x b ra nch
L e f t c o r o n a r y a .- -
R ig h t a u r i c le - -
G r e a t c o r o n a r y v. - - L. v e n t r i c / e
R ig h t v e n t r i c le ''
I '' Apex
V e n t r a l i n t e r v e n t r i c u l a r S u lc u s
Fic. 4-4. The heart, left lateral aspect.
272 Chapter 4. T he H ea rt and A r t e r ie s
R i g h t p u l m ono ry vv., R. p u lm o n a r y a.
P o s tc a v a ■ «
I , A z y g o s v.
• R. a u r i c l e
R ig h t v e n tric Ie --COnuS a r te r io s u s
' R i g h t corona ry a-
Apex I R i g h t p ro x im a l a. o f a t r i u m
L e f t vent r i d e i ‘ R ig h t a triu m
V e n tro t i n t e r v e n t r i c u l a r br. R i g h t d i s t a l a. o f a t r i u m
of I e f t corona r y a.
Fic. 4-5. The heart, right lateral aspect.
H ea rt 273
plane, into a cranioventral (right) and a caudo- covering it, the beat of the organ can be easily
dorsal (left) part. These in turn are partly divided heard and felt through the thoracic wall. Accord
transversely into the blood-receiving chambers, ing to Detweiler (1955), the heart can be felt
the atria, and the pumping chambers, the ven beating more forcibly against the left chest wall
tricles. The internal partitions are indicated on than against the right. The conformation of the
the surface of the organ by grooves. The caudo- dog has most to do with the ease with which the
dorsa! part of the heart, consisting of the left heart beat can be felt and heard. The heart beat
atrium (atrium sinistrum) and the left ventricle is most pronounced in thin, narrow-chested,
(ventriculus sinister), receives blood from the athletic type animals.
lungs and pumps it to all parts of the body (sys
temic circulation). The cranioventral part of the Size and Weight
heart, consisting of the right atrium (atrium dex-
trum) and right ventricle (ventriculus dexter), re The dog heart is frequently used in studies of
ceives blood from all parts of the body and cardiac hypertrophy, and several surveys have
pumps it to the lungs (pulmonary circulation). been made of the normal values. Herrmann
Although the terms left and right are not de (1925) includes data on 200 dogs and cites heart
scriptively accurate as to the portions of the weight to body weight ratios averaging 8.10
heart concerned in the systemic and pulmonary grams per kilogram of body weight for males
circulations in the dog, they will, in deference to and 7.92 for females. Northrup, Van Liere, and
custom, be used to designate these parts. Stickney (1957) analyzed Herrmann’s data and
found that the sex differences were not signifi
Orientation cant. However, when they studied the heart
weight to body weight ratios of an additional
The heart forms a part of the mediastinum, 346 adult dogs and 135 pups, they found that
the partition which separates the two pleural females have significantly smaller ratios than
cavities. It is the largest organ in this tissue-filled males. Small adults were found to have higher
septum located between the walls of the medi ratios than large adults, although pups had sig
astinal pleurae. Covered by its pericardium, the nificantly smaller ratios than those in any adult
heart extends from the third rib to the caudal category. The average ratio for 169 adult males
border of the sixth rib. Roentgenograms show was 7.74, and for 177 females it was 7.56. The
that variations in position occur among the range for 346 adult dogs was 4.53 to 11.13 gm./
breeds, strains, and individuals, and in the same kg. Latimer (1961) has presented data on the
animal according to age, condition, and the pres ratios between the weights of the walls of the
ence of pathological processes. A longitudinal right and left ventricles in 46 dogs. The right
axis through the heart tips forward about 45 de ventricular wall accounted for 22 per cent of the
grees from a vertical plane. The base, therefore, total heart weight, the left ventricular wall for
faces dorsocranially and the bulk of it lies 39 per cent.
above a frontal plane dividing the thorax into In the average young dog the heart weight is
dorsal and ventral halves. The apex points cau- approximately 1 per cent of the body weight.
doventrally where it lies slightly to the left and
caudal to a transverse plane through the most Surface Topography
cranial part of the diaphragm. It touches the
cranial surface of a transverse plane through the The coronary groove (sulcus coronarius)
thorax which bisects the caudal part of the marks, on the surface of the heart, the separation
seventh sternebra. Normally the lungs cover of the atria and ventricles. It contains much fat,
most of the surface of the heart. On the right which surrounds the coronary vessels. The coro
side the cardiac notch of the lung allows a vari nary groove encircles the heart except cranio-
able area of the heart, covered by its fibrous peri ventrally, where the dorsal part of the right ven
cardium and three layers of serosa, to contact tricle (pulmonary conus) intervenes.
the lateral thoracic wall. The cardiac notch of The interventricular grooves are indistinct sur
the right lung is V-shaped, with the apex di face markings of the separation of the right and
rected dorsally. This allows a greater exposure of left ventricles. Since they neither indent the
the heart on the right side than on the left, where substance of the heart nor contain as much fat as
the ventral border of the lung is usually not does the coronary groove, vessels are visible in
notched. The lung is thin adjacent to the lateral them; however, most vessels stream toward the
surfaces of the heart so that, in spite of the lung apex on the surface of the ventricles, rather than
274 Chapter 4. The H e a r t an d A r te r ie s
following the interventricular grooves. Obliquely precava, although occasionally it enters the
traversing the cranioventral surface of the heart atrium directly. The azygos vein drains blood
is the ventral (cranial) interventricular groove back to the heart from part of the lumbar region
(sulcus interventricularis ventralis). It begins on and the caudal three-fourths of the thoracic wall.
the caudodorsal side of the pulmonary conus, The precava returns blood to the heart from the
where it is covered by the auricular portion of head, neck, thoracic limbs, the ventral thoracic
the left atrium; it ends before reaching the apex wall, and the adjacent part of the abdominal
of the heart. The dorsal (caudal) interventricular wall. The right atrioventricular orifice (ostium
groove (sulcus interventricularis dorsalis) is a atrioventriculare dextrum) is the large opening
short, straight, shallow furrow which marks, on from the right atrium into the right ventricle.
the dorsocaudal surface of the heart, the approxi This opening and the valve which guards it are
mate position of the interventricular septum. described with the right ventricle.
Two surfaces of the heart are recognized. The Other features of the right atrium are the tu-
one which faces the diaphragm is the diaphrag berculum intervenosum, crista terminalis, fossa
m atic surface (facies diaphragmatica). This lies ovalis, limbus fossae ovalis, and the mm. pecti-
in an oblique plane, and is flattened. The other nati. On the dorsal wall of the right atrium is a
surface, the sternocostal surface (facies sterno- transverse ridge of tissue placed between the two
costalis), is rounded and, as the name implies, caval openings. This is the intervenous tubercle
faces toward the sternum and ribs. This consti (tuberculum intervenosum). It diverts the con
tutes about two-thirds of the surface area of the verging inflowing blood from the two caval veins
cone-shaped organ. A line drawn on the left side into the right ventricle. Just caudal to the interve
from the root of the pulmonary trunk to the apex nous tubercle on the medial wall of the atrium is a
would follow an ill-defined border, known as the slitlike depression, the fo ssa ovalis, which varies
left border (margo sinister), located at the junc from one to several millimeters in depth. The
tion of the sternocostal and diaphragmatic sur crescent-shaped ridge of muscle which projects
faces on the left side. A similar poorly defined from the caudal side of the intervenous tubercle
border on the right side is known as the right and deepens the fossa is the limbus fossae ovalis.
margin (margo dexter). The base of the heart In the fetus there is an opening at the site of the
(basis cordis) is the hilus of the organ. It is the fossa, the foram en ovale, which allows blood to
craniodorsal portion and receives the great veins pass from the right to the left atrium. The fora
and emits the great arteries. The atria also enter men usually closes during the first few postnatal
into its formation. The apex of the heart (apex weeks. Even though a small anatomical opening
cordis) is formed by the looping of a swirl of frequently persists, because the obliquity of the
muscle fibers at the apex of the left ventricle. It foramen enables the greater pressure in the left
forms the most ventrocaudal part of the organ. atrium to close the passage, it is closed physio
logically.
Atria The right auricle (auricula dextra) is the blind,
ear-shaped pouch of the right atrium which ex
The right atrium (atrium dextrum) (Fig. 4-6) tends cranioventrally. The internal surface of the
receives the blood from the systemic veins and wall of the right auricle is strengthened by freely
most of the blood from the heart itself. It lies branching, interlacing muscular bands, the mm.
dorsocranial to the right ventricle. It is divided pectinati. These are also found on the lateral
into a main part, the sinus venarum cavarum, wall of the atrium proper. Most of the pectinate
and a blind part which projects forward and muscles radiate from a semilunar crest, a thick
downward, the right auricle (auricula dextra). ridge of tissue, which is placed between the en
The main openings of the right atrium are four trance of the precava and the atrioventricular
in number. The coronary sinus (sinus corona- opening. This is the crista terminalis. It is also
rius) is the smallest of these, and enters the the dorsal separation of the sinus venarum cava
atrium from the left. Dorsal to it is the large rum and the auricle. On the external surface of
postcava (vena cava inferior), which enters the a dilated heart a poorly defined groove, the sul
heart from behind. The postcava returns blood cus terminalis, lies opposite the crista terminalis
from the abdominal viscera, part of the abdomi at the ventral part of the junction of the precava
nal wall, and the pelvic limbs. The precava (vena with the atrium. Small veins empty into the right
cava superior) is about the same size as the post atrium through openings in the pits between the
cava. It enters the heart from above and in front. mm. pectinati. Everywhere, the internal surface
In the dog the azygos vein usually enters the of the heart is lined with a thin glistening mem-
H ea rt 275
In te rv e n o u s tu b e rc le
f
f l i g h t pulm onary a 1 ,R ig h t p u lm o n a r y vv,
Lirnbus f o s s a e 0v o l i Sj I i ‘
I ' ,AzygoS v
Fasso o v o l i s \ i * i
' , i 1 R i g h t a t r iu m
R ig h t p u lm o n a r y s/.'v t >( J • /
, precava
P o s tc a va - „
,A o rtic a rch
-Peri c o r d : um
O pe ning o f
Cnsta te rm in a liS
c o ro n a ry S in u s
'"M m pectinoti
'R ig h t auricle
Conus a r t e r i o s u s
R ig h t v e n t r i c l e '
Septa l cusp of r ig h t
a t r io v e n t r ic u l a r v a lv e
Ven tral in t e r v e n t r ic u la r s u ic u s
L e ft vent ri cl e ‘
F ig. 4 -6, A dissection showing the interioi of th» right atrium right latei.il aspecl
276 Chapter 4. T he H eart and A r t e r ie s
brane, the endocardium, which is continuous the ascending aorta. There are three points on
with the tunica intima of the blood vessels enter its aortic margin, each of which conforms to the
ing and leaving the organ. attachments of two adjacent semilunar valvulae
The left atrium (atrium sinistrum) forms the of the aortic valve. The projections between the
left, dorsocaudal part of the base of the heart. attachments of the dorsal and right and the
The mm. pectinati are confined to its left auricle dorsal and left semilunar valvulae are best de
(auricula sinistra), which is similar to the right veloped and are composed of hyaline cartilage.
auricle in shape and structure. It lies caudal to Between the points, the margin of the aortic
the pulmonary conus and pulmonary trunk. Its fibrous ring is semilunar so that its scalloped cir
apex covers the proximal end of the left longi cumference is in the form of three arcs. To the
tudinal groove. The left auricle lies along the right of the aortic fibrous ring the fibrous base of
right auricle, caudal to it, for nearly its entire the heart is best developed. Here, in the interval
length, the two being separated by the pulmo between the right and left atrioventricular ostia,
nary trunk. Elsewhere on the surface of the it consists of the thickened parts of the aortic
heart there is no distinct indication of the sepa and left atrioventricular rings as they lie adja
ration of the two atria. Internally, the atria are cent to each other. Because of its triangular
separated by the interatrial septum (septum shape, it is called the right fibrous trigone (trigo-
interatriale). Five or six openings (ostia venarum num fibrosum dextrum). The left fibrous trigone
pulmonalium) mark the entrance of the pulmo (trigonum fibrosum sinistrum) is smaller than the
nary veins into the atrium. The two or three right and lies in the triangle between the aortic
veins from the right lung cross over the right and left atrioventricular ostia to the left of the
atrium and open into the craniodorsal part of the right fibrous trigone. Both trigones contain carti
chamber; the three veins from the left lung usu lage and are united through the medium of the
ally empty into its caudodorsal part. The veins opposed aortic and left atrioventricular fibrous
from the diaphragmatic lobes are larger than the rings. In man they are fibrous, hence the name
others and are most caudal in position. Fre fibrous trigone. In the ox and horse, bone largely
quently a thin concave flap of tissue is present replaces the soft tissue in these areas.
on the cranial part of the septal wall. This is the The pulmonary fibrous ring (annulus fibrosus
valve o f the foram en ovale (valvula foraminis pulmonalis) is similar to the aortic ring. It serves
ovalis). for the attachment of the pulmonary semilunar
valvulae and distally it blends with the media of
Fibrous Base the pulmonary trunk. It is much weaker than the
aortic ring. The muscle fibers of the conus arteri
The fibrous base of the heart, or “cardiac osus attach to its distal surface. Between the ap
skeleton” (Fig. 4-7, B), is the fibrous tissue, con posed surfaces of the pulmonary and aortic
taining some cartilage, which separates the thin annuli there is a short mass of collagenous tissue,
atrial musculature from the much thicker muscle the ligament o f the conus, which unites these
of the ventricles. Only the special neuromuscular two structures.
tissue, the atrioventricular bundle, extends con The atrioventricular fibrous rings (annuli fi
tinuously through the fibrous base from the atria brosi atrioventriculares) (Fig. 4-7, B) are thin
to the ventricles. Baird and Robb (1950) have rings of collagenous tissue to which the muscle
reconstructed the principal parts of the conduc fibers of the atrial and ventricular walls attach.
tion system of a puppy’s heart. Their dissections From their endocardial edges emanate the atrio
and reconstruction show the close spatial rela ventricular valves. Although the proximal parts
tionship which exists between the primary con of the ventricular musculature attach to their
duction and supporting tissues of the dog’s distal surfaces, the bulk of the fibers of this mus
heart. The cardiac skeleton is in the form of two cle at its origin lie peripheral to the rings as the
narrow fibrous rings (annuli fibrosi), one sur muscle tissue bulges proximally after arising
rounding each atrioventricular orifice, and two from them. These fibrous rings are so delicate
scalloped cuffs, or rings, one surrounding each they are best seen in longitudinal sections of the
arterial orifice. heart cut through the atrioventricular junctions.
Of the two arterial rings, the aortic fibrous
ring (annulus fibrosus aorticus) is the better de Ventricles
veloped. Peripherally the collagenous fibers
which form it give way to the yellow elastic The ventricles form the bulk of the heart. To
fibers composing the tunica media of the wall of gether they form a conical mass, the apex of
H eart 277
DORSAL
Dorsal i n t e r v e n t r i c u l a r b r ■ / L e ft v e n tr ic le
/ L e f t a tr io v e n tr ic u la r valve
R igh t fib r o u s t r i g o n e ~ L e f t f ib r o u s tr ig o n e
s C irc u m fle x b ra n c h
^ L e f t c o r o n a r y a.
f. a t r io v e n t r i c u l a r valve —
S e p ta l br.
R i g h t v e n t r ic le —
-V e n tr a l i n t e r v e n t r i c u l a r br
Dorsal s e m ilu n a r L e f t sem i l u n a r v a lvu la e
va lv u la of a o r t i c valve
''Vent, sem ilunar valvula of pulmonary valve
R i g h t c o r o n a r y a. R i g h t s e m ilu n a r va lv u la e
R ing f o r
l e f t a t r i o v e n t r i c u l a r v a lv e s
R. f ib r o u s tr ig o n e ~~
Ring f o r r a t r i o
v e n t r i c u l a r valve -----
Root
which is also the apex of the left ventricle. The constant papillary muscle of the conus. The
right ventricle is ventral and cranial to the left numerous chordae tendineae which go to the
and arciform in shape (Fig. 4-8). septal or dorsal cusp arise from the septal wall
The right ventricle (ventriculus dexter) (Fig. directly or from muscular ridges or papillae
4-9) receives the systemic blood from the right located peripheral to the septal or dorsal cusp,
atrium and pumps it to the lungs. Its oudine is when this valvula lies against the septum.
in the form of a curved triangle, as it is molded The trabeculae carneae are myocardial ridges
on the surface of the caudodorsally lying conical which project mainly from the outer wall of the
shaped left ventricle. It is crescentic in cross ventricle. Since they are endocardial-covered
section, and its long axis extends from the dorsal portions of the deepest muscular layer of the
interventricular groove to the pulmonary trunk, right ventricle, their long axes parallel the direc
which leaves the most craniodorsal part of the tions of these fibers. They run from the base and
ventricular mass. converge toward the apex. Those on the septal
The potentially large opening through which wall are largely adjacent to the ventral inter
the blood enters the right ventricle is the right ventricular sulcus and they largely parallel the
atrioventricular ostium (ostium atrioventriculare axis of the heart. Some of the fossae coalesce
dextrum). Blood leaves the chamber through the near the ventral interventricular groove so that
pulmonary ostium (ostium trunci pulmonalis) muscular columns or pillars are formed.
and is received by the pulmonary trunk. The The chordae tendineae are fibromuscular
conus arteriosus, or infundibulum, is the funnel- cords which arise from the apices of the papillary
shaped part of the right ventricle which lies be muscles or directly from smaller, blunter eleva
tween planes going through the two openings of tions on the wall. The cords branch, at their
the ventricle. It is embraced by the right auricle valvular extremity, as they approach the free
externally. The supraventricular crest (crista border of the cusps. The smaller cords blend
supraventricularis) is a blunt, obliquely placed with the tunica media of these cusps at the points
ridge of muscle between the origin of the conus of their free borders, and the larger strands fan
arteriosus and the atrioventricular opening. out on the ventricular surfaces of the cusps.
Both ventricles contain muscular projections The trabecula septomarginalis, formerly called
and small, usually deep and oblong depressions. moderator band, is a thin, delicate, branched,
The conical-shaped muscular projections which muscular strand which extends across the lumen
give rise to the chordae tendineae are called of the right ventricle. It usually leaves the septal
papillary muscles (musculi papillares). They are wall of the right ventricle near or from the base
the most conspicuous features of the ventricular of the largest papillary muscle and runs to the
walls. There are usually three main papillary peripheral wall. The extremity of the trabecula
muscles in the right ventricle, although great usually branches repeatedly as it blends with the
variation exists. Typically they arise from the muscular ridges of the outer right ventricular
apical third of the septal wall, 1 to 3 cm. from its wall. It serves the morphological role of con
junction with the outer wall. Their branched ducting a fascicle of Purkinje fibers from the
chordae tendineae go to the free border and ad right branch of the atrioventricular bundle across
jacent ventricular surface of the ventral cusp of the lumen of the cavity. Instead of the usual
the right atrioventricular valve. Commonly the single band, there may be two or more anasto
papillary muscle which lies farthest to the right mosing strands forming a loose plexus.
is larger than the others and, when it is, its apex The left ventricle (ventriculus sinister) (Fig.
is usually bifid. The papillary muscle which lies 4-10) is conical in shape, with its apex forming
farthest to the left may be bifid also. In such the apex of the heart. It receives the oxygenated
specimens the middle muscle is absent. A single blood from the lungs by way of the left atrium
compound papillary muscle may replace three and pumps it to most parts of the body through
main ones. Other variations are common. Near the aorta. The left atrioventricular orifice (ostium
the angle formed by the septal and the outer atrioventriculare sinistrum) is the large opening
wall, and caudodorsal to the most sinistral large between the left atrium and the left ventricle.
papillary muscle, is a small, blunt to conical pap The aortic orifice (ostium aortae), located near
illary muscle which gives rise to chordae tendi the center of the base of the heart, is the open
neae going to the most caudodorsal part of the ing from the left ventricle into the ascending
ventral cusp. Truex and Warshaw (1942), in the aorta. The left ventricle is characterized by its
12 dogs they studied, found only three large thick wall, conical shape, and its two large pap
papillary muscles from the septum and a small illary muscles. Its wall is three or four times
H ea bt 279
, L e f t v e n t r i c le
i
i R i g h t v e n tr ic le
Fic. 4-8. Cross section through the ventricles, craniodorsal aspect.
A o rtic arch |
I 'P u lm o n a r y tru n k
I
Precava,
i
R i g h t a u r i c l e ----- V e n t r a l s e m i l u n a r v a lv u l a
of p u l m o n a r y v a lv e
Coronary s u l c u s -
Septal c u s p o f r i g h t - - -V e n tra l in t e r
a t rio v e n t ric u la r valve v e n tric u la r su lc u s
Lateral c u s p o f r i g h t - -
a trio v e n tric u la r v a lv e
P a p i I l a r y m u s c l e s 1'
Left ven tricle
iT r a b e c u l a e c a r n e a e
Fu.. 4-9. A dissection showing the interior of the right ventricle, ventral aspect.
280 Chapter 4. The H e a rt and A r te r ie s
, 'L a u r i c le
R i g h t a u r i c l e >.
R s e m i I un a r - - - D o r s a l cusp o f I.
v a lv u la of atrioventricular
a o r t ic valve valve
V ent r a l c u s p o f
I. a t r i o v e n t r i c u l a r
v a l ve
/
Trabeculae carneae'
Fic 4-10. A dissection showing the interior of the left ventricle, left lateral aspec t
H ea rt 281
thicker than that of the right ventricle. Truex triculare) consists of an inconspicuous, small,
and Warshaw (1942), in 12 specimens, found the proximally located membranous part and a large,
mean thickness of the left and the right ventricle thick muscular part. The membranous part (pars
to be 13 and 4.2 mm., respectively. The in membranacea) of the interventricular septum is
creased thickness of the left ventricle is due pri the thinnest part and is the last part of the sep
marily to the approximately threefold increase tum to form embryonically. This is brought
in the number of fibers it contains, compared about in man (Odgers 1938) by the fusion of the
with the right ventricle. Part of it is due also to atrioventricular cushions and not by a down
the fact that the individual fibers of the left ward growth of the aortic (pulmonoaortic) sep
ventricle are slightly larger than those of the tum to meet the interventricular septum. In
right. Ashley (1945) points out that, in histolog man, this closure is completed by the end of the
ical material, the capillary to fiber ratio in the second month of gestation. In the dog the mem
dog’s heart is essentially 1 to 1. branous part can be seen by transmitted light
The two large papillary muscles (musculi under the septal cusp of the right atrioventricu
papillares) of the left ventricle come from its lar valve adjacent to the origin of the aorta.
outer wall. They are heavy, smooth rolls of myo When the foramen fails to close, a subaortic de
cardium which have compound apices and give fect or interventricular foramen is left. The
rise to the stout chordae tendineae of this cham muscular part (pars muscularis), which consti
ber. The dorsal papillary muscle (musculus pa tutes the bulk of the interventricular septum, is
pillaris dorsalis) lies near the dorsal interventricu formed by the myocardium of the combined
lar groove; the ventral papillary muscle (muscu walls of the two ventricles as they lie adjacent to
lus papillaris ventralis) is closer to the ventral each other. It is usually 1.5 to 2 cm. thick.
interventricular groove. Adjacent to the ventral
papillary muscle, near its attachment, is a fine Myocardium
network of muscular strands which come from
the septal wall. From findings in similar strands The myocardium, or heart muscle of the atria,
from other species (Truex and Warshaw 1942), is not divided into distinct layers, except at the
it is probable that these contain fascicles of interatrial septum. Here the deep fibers of the
Purkinje fibers derived from the left branch of two chambers fold in and lie adjacent to each
the atrioventricular bundle en route to the ven other, to form this partition, whereas the super
tral papillary muscle and general ventricular ficial fibers are common to both atria. Muscular
musculature. The strand nearest the atrioven fibers encircle the ostia of the systemic and the
tricular opening is larger than the others and ex pulmonary veins as they empty into the atria.
tends obliquely to the ventral papillary muscle Between the pectinate muscles the musculature
from the septal wall. It may be the only one is so thin that the heart wall is translucent at
present. Near the apex of the ventricle some of these places. The fixed point of the atrial mus
the fine threads which compose this network ex culature, like that of the ventricles, except for
tend under the endocardium which covers the the atrioventricular bundle, is the fibrous base of
crests of the muscular ridges. Other threads the heart.
bridge the sulci between the larger trabeculae The musculature of the ventricles in the dog
carneae in their courses to the ventral papillary was described by Thomas (1957). There are
muscle. The chordae tendineae from the ventral superficial, middle, and deep layers. All mus
papillary muscle go to both the ventral and the cle fasciculi of the superficial layer arise from the
dorsal part of the left atrioventricular valve. fibrous base and return to it. The superficial
Those which go to the ventral or septal part are layer is common to both ventricles. When the
shorter and arise closer to the ventricular wall heart is viewed from the base, these bundles run
than those which go to the dorsal or outer part toward the apex, showing a clockwise twist. At
of the valve. The dorsal papillary muscle is the apex of the heart they turn in and run toward
separated from the ventral muscle by a deep the base in such a manner that they cross, at
cleft. The origin, course, and termination of the right angles, the superficial fibers running down.
chordae tendineae from this muscle are similar They form the papillary muscles of the left ven
to those of the chordae tendineae from the ven tricle. The superficial fibers penetrate the middle
tral muscle. Both papillary muscles extend to layer of the right ventricle to become its deep
within a few millimeters of the apex of the ven layer, after which they are disposed as on the
tricle. left side. The middle layer forms the bulk of the
The interventricular septum (septum interven- ventricular walls. These are spiral or circular
282 Chapter 4. The H e a rt and A r te r ie s
muscle masses which interdigitate between the tal space at the level of the costochondral junc
two chambers. They primarily decrease the size tion, the so-called “tricuspid area.”
of the lumen of the ventricles, and the superficial The left atrioventricular valve (valva atrio-
and deep layers shorten and twist the organ. The ventricularis sinistra), or mitral valve (valva mi-
apex of the heart is quite thin, being formed by tralis), is basically similar to the right atrioven
muscle fasciculi of the superficial layer of the left tricular valve in form and structure, but it is
ventricle as they swirl in figure-of-eight fashion made on a heavier scale. The chordae tendineae
to form the apex of the chamber. as well as the papillary muscles from which they
arise are several times larger in the left ventricle
than they are in the right. Likewise their stratum
Atrioventricular Valves proprium contains considerably more collage
nous tissue. This stronger construction of the left
The atrioventricular valves (valvae atrioven- intake valve, as compared with the right, is nec
triculares) (Figs. 4 -7 , 4 -9 , 4-10) are irregular, essary as the blood leaving the left ventricle
serrated cusps which are located in the atrioven through the aorta is under about four times more
tricular ostia. They are the intake valves to the pressure than that which leaves the right ventri
ventricles. They prevent blood from returning cle through the pulmonary trunk (Dukes 1955).
to the atria during the systolic phase of the heart The division of the left atrioventricular valve
beat. Peripherally, they attach to the fibrous into cusps is indistinct. The part which arises
rings which separate the musculature of the atria from the fibrous ring adjacent to the septum is
from that of the ventricles. When the ventricles wider than that coming from the remainder of
contract, the valves are kept from being pushed the ring so that when the valve is open and
into the atria by the chordae tendineae. Accord viewed from the atrial side it is tricuspid in ap
ing to Trautmann and Fiebiger (1952), the stra pearance.
tum proprium of the atrioventricular valves con According to Detweiler (1955), murmurs as
sists of connective tissue rich in elastic fibers. sociated with disease of the mitral valve are gen
This is covered by a thin endocardium on each erally most intense at the fifth left intercostal
surface. The chordae tendineae attach to the space above the middle of the lower third of the
ventricular surfaces of the valvulae. The strong thorax, the so-called “mitral area.”
est cords can be followed as ridges under the en
docardium to their attached borders, whereas Aortic and Pulmonary Valves
the weakest cords go to the points of the serra
tions and disappear. The medium-sized cords go The aortic valve (valva aortae) (Figs. 4-7, 4 -
as far as the middle part of the ventricular sur 10) consists of right, left, and dorsal semilunar
faces of the valvulae before blending with the cusps, or valvulae (valvulae semilunares dextra
stratum proprium. Although some blood vessels et sinistra et dorsalis). These consist of a fibrous
have been described in the valves adjacent to tissue stroma covered on each surface by endo
their attached borders, the bulk of the nutrition thelium. Opposite their free borders they are at
of the valves is derived from the free blood in tached to the aortic fibrous ring. In the middle
the heart. of the free borders of the semilunar cusps are
The right atrioventricular valve (valva atrio- nodules (noduli valvularum aortae). Extending
ventricularis dextra) in man is also known as the from each nodule toward the periphery of the
tricuspid valve. The valve in the dog consists valvulae are the lunulae (lunulae valvularum
basically of two cusps. The cusp of the right semilunarium). These represent the areas of con
atrioventricular valve which attaches to the tact with the adjacent cusps when the valve is
fibrous ring adjacent to the septum is called the closed. The nodules close the space that would
septal or dorsal cusp (cuspis dorsalis). The cusp otherwise be left open by the coming together of
which attaches to the fibrous ring adjacent to the the three contiguous arcs. On the vessel side, or
ventral wall is the ventral cusp (cuspis ventralis). peripheral to each of the semilunar cusps, the
The extremities of the dorsal and ventral cusps wall of the aorta is dilated to form the three aor
become narrower and merge or, in some speci tic sinuses (sinus aortae). These, like the cusps,
mens, small secondary cusps are formed at these are right, left, and dorsal in position, with the
sites. right and left coronary arteries leaving the right
According to Detweiler (1955), tricuspid and left sinuses. The sinuses function in holding
valve lesions cause murmurs which are usually a reserve of blood which prevents the cusps from
heard most distinctly at the fourth right intercos being closely applied to the aortic wall. Thus fol
H ea rt 283
lowing the end of systole, or heart contraction, ventricularis). This is about 1.5 mm. in diameter
the pressure in the aorta is greater than that in in the dog, according to Baird and Robb (1950),
the left ventricle, and a back pressure is devel and shows little histological differentiation from
oped. This immediately forces the cusps to the bundle. Nonidez (1943) demonstrated that
gether, closing the valve, thus retaining the the atrioventricular node received a richer para
blood in the aorta and allowing the left ventricle sympathetic supply than did the sinoatrial node.
to fill from the atrium. The atrioventricular node begins in the septal
The valve of the pulmonary trunk (valva wall of the atrium about 5 mm. cranioventral to
trunci pulmonalis) (Figs. 4-7, 4-9) lies cranial to the opening of the coronary sinus and craniodor-
the aortic valve, and is similar to it in construc sal to the septal cusp of the right atrioventricu
tion. Since the blood pressure developed in the lar valve. From this apparently blind beginning
pulmonary trunk which it guards is not as great the atrioventricular bundle runs forward and
as that in the aorta, the development of the downward through the fibrous base of the heart.
cusps and related structures is not as great. All As it does so it divides into right and left
parts present in the aortic valve are represented branches. These lie closely under the endocar
in the pulmonary. The valve of the pulmonary dium of the septal wall of the right and left ven
trunk consists of right, left, and ventral semi- tricles. The right septal branch crosses the cavity
lunar cusps, or valvulae (valvulae semilunares of the right ventricle in the septomarginal tra
dextra et sinistra et ventralis). becula of this chamber. It arborizes in the outer
According to Detweiler (1955), the “aortic ventricular wall of the right ventricle, where it
area,” the area in which sounds associated with ends. The left septal branch is more diffuse than
lesions of the aortic valve may be heard, is lo the right one, and partly traverses the cavity of
cated at the fourth left intercostal space slightly the left ventricle to the outer wall of this cham
below a line drawn through the point of the ber in bundles which are smaller and more
shoulder. Sounds associated with functioning of branched than those on the right side. The rami
the pulmonary trunk valve can also best be fication of the conduction system of the heart is
heard at this place. Furthermore, such sounds more complicated than the previous description
are audible on each side and both farther for indicates. Baird and Robb (1950) found that
ward and more ventrally than are the sounds of there were transitions from the atrioventricular
the aortic valve. node to the atrial muscle and that various parts
of the atrioventricular bundle blended with the
Conduction System general ventricular musculature. Abramson and
Margolin (1936) say that in the dog, as in other
No part of the conducting system of a pre species, the myocardial branches as they leave
served dog’s heart can be adequately identified the subendothelial plexus tend to pass perpen
in gross dissection. It consists of three parts, dicularly to the endocardium in the left ventricu
which are closely integrated physiologically: (1) lar wall and obliquely in the right ventricular
the sinoatrial node, (2) the atrioventricular node, wall. They further state that the interventricular
and (3) the atrioventricular bundle. septum is traversed by myocardial Purkinje fi
The sinoatrial node (nodus sinuatrialis) ap bers which arise from the adjacent subendothe
pears to be the center which initiates the heart lial Purkinje networks. No one has demonstrated
beat and also regulates the interval between a structural link between the sinoatrial and
beats. It is located in the terminal crest at the atrioventricular node or bundle.
confluence of the precava, sinus venarum cava-
rum, and auricular orifice. When it is destroyed B lo o d V e s s e ls o f th e H eart
experimentally the heart beat slows or stops
(Dukes 1955). The sinoatrial node is composed Coronary Arteries
of Purkinje fibers little modified from those
which compose the atrioventricular conduction Four arteries supply the heart. These are the
system. Nonidez (1943) has shown that both the right coronary artery and the three branches of
sinoatrial and the atrioventricular node are sup the left coronary artery: the septal, the circum
plied by postganglionic parasympathetic nerve flex, and its continuation in or near the dorsal in
terminals. terventricular groove, the dorsal interventricu
The atrioventricular bundle (fasciculus atrio- lar.
ventricularis) begins as a mass of Purkinje fibers The right coronary artery (a. coronaria dex
known as the atrioventricular node (nodus atrio- tra) (Figs. 4-5, 4-7, A) is smaller than the left
284 Chapter 4. The H e a rt and A r te r ie s
coronary artery and measures about 1.5 mm. in coronary artery and adjacent vessels are small.
diameter and 5 cm. in length. It arises from the Donald and Essex (1954) found that, in those
right sinus of the aorta and makes a sweeping dogs which survived the ligation of the right
curve to the right and ventrocranially, lying in coronary artery, four demonstrable anastomoses
the fat of the coronary groove. Its initial part could be found after 30 days. These were as fol
is bounded by the pulmonary trunk and the lows:
conus arteriosus craniolaterally, and dorsally it (1) By means of the right (anterior) and left at
is covered by the right auricle. The right coro rial (auricular) arteries.
nary artery supplies the bulk of the outer wall (2) Through the large, often tortuous, connec
of the right ventricle. As a result of ligation stud tion of the (left) circumflex and the right coro
ies, Donald and Essex (1954) found that an aver nary artery.
age of 66 per cent of the free right ventricular (3) Through small connections between the
wall normally receives its blood through the ventral interventricular (anterior descending)
right coronary artery. According to Kazzaz and and the ventricular (lateral) branches of the right
Shanklin (1950), the right coronary artery rarely coronary artery.
extends as far as the borders of the right ventri (4) By means of several branches of the dorsal
cle and in no case goes beyond them. It also interventricular (descending branch of the left
sends branches to the right atrium, and small circumflex) and the last large ventricular (lateral)
twigs go to the initial parts of the aorta and pul branch of the right coronary artery.
monary trunk. In 20 per jent of specimens, ac The left coronary artery (a. coronaria sinistra)
cording to Moore (1930), an accessory right coro (Figs. 4-4, 4-7, 4-11) is a short trunk about 5
nary artery (a. coronaria dextra accessoria) arises mm. long and nearly as wide. It always termi
closely adjacent to the main right coronary ar nates in the circumflex and ventral interventric
tery from the right sinus of the aorta and runs ular branches and, in many specimens, in a sep
about 2 cm. out on the conus arteriosus, where it tal branch also.
largely becomes dissipated. The circumflex branch (ramus circumflexus),
According to Pianetto (1939), there are 4 to 9 about 1.5 cm. in diameter, lies in the coronary
ventricular branches of the main right coronary groove as it extends to the left, then winds dorso-
artery. Most of these are small vessels which ar caudally and to the right across the caudodorsal
borize on and in the right ventricular wall. They surface of the heart. On reaching or approach
run at right angles to the long axis of the cavity, ing the dorsal interventricular groove it turns to
and none extend as far as the ventral interven ward the apex of the heart and is known as the
tricular groove. One of these, the right marginal dorsal interventricular branch. The combined
branch (ramus marginalis dextra), is larger, lengths of the circumflex and dorsal interven
longer, and more branched than the others. It tricular branches is approximately 8 cm. The
supplies the middle part of the right lateral ven circumflex branch has ventricular and atrial
tricular wall. branches.
The atrial branches from the right coronary According to Kazzas and Shanklin (1950),
artery are variable in development. Kazzaz and there are 4 to 11 ventricular branches. Pianetto
Shanklin (1950), Meek et al. (1929), Moore (1939) found 2 to 6 principal ventricular
(1930), and Pianetto (1939) all describe an atrial branches. Most of the hearts examined in the
branch, larger than the others, which leaves the present study had five main ventricular branches
distal half of the artery, traverses the sulcus ter- leaving the circumflex branch, but great varia
minalis and, as it does so, supplies the sinoatrial tion was found. When the left ventricular
node. This vessel anastomoses with one or branches of the ventral interventricular branch
more atrial branches which arise from the cir are well developed they are the major source of
cumflex vessel. Usually one or two small atrial supply to the outer wall of this chamber, and the
rami leave the right coronary artery both proxi ventricular branches of the circumflex are con
mal and distal to the branch destined for the fined to a triangular area adjacent to the coro
sinoatrial node. The distal branches supply the nary and dorsal interventricular grooves. When
caudal part of the right atrium adjacent to the first or first few branches of the circumflex
the coronary sulcus. A terminal twig ends on the branch cross the superficial muscle fibers of the
postcava. The first branch or two which leave left ventricle obliquely, they are short; they are
the dorsal surface of the right coronary artery much longer when they parallel the superficial
supply the right auricle. muscle fibers as well as the left ventricular
The normal anastomoses between the right branches of the ventral interventricular vessel.
H ea rt 285
Usually the longest branch leaving the circum parallel the superficial muscle fibers of the left
flex lies adjacent and parallel to the dorsal inter ventricle they lie in grooves on its surface. Usu
ventricular branch. This is appropriately known ally the branch which arises near the end of the
as the left marginal branch (ramus marginalis proximal third of the parent vessel is longer than
sinister), as it follows the left border down the the others and supplies the apex of the heart.
surface of the left ventricle. Most of the left ventricular branches are quite
The atrial branches of the circumflex branch free of superficial collateral branches.
of the left coronary artery, which supply the left The right ventricular branches (rami ventric-
atrium and auricle, are small and variable. The ulares dextri), usually five in number, supply a
first branch arises deep to the great coronary strip, about 2 cm. wide, of the right outer ven
vein and, extending dorsally and toward the cen tricular wall adjacent to the ventral interventric
ter of the base of the heart, supplies the deep ular groove. The first branch is prominent as it
surface of the left atrium and the ventral part partly encircles the conus arteriosus adjacent to
of the interatrial septum. It is the largest of the the origin of the pulmonary trunk. This is the
right atrial branches and, according to Meek et conal branch. It anastomoses with the conal
al. (1929), its terminal branches may partly en branch of the right coronary artery. Most of the
circle the termination of the precava, where remaining branches are short and form small
they anastomose with the right atrial vessel to anastomoses with the branches of the right coro
the sinoauricular node. It may be the main nary artery and the dorsal interventricular
source of supply to this node. At least two other branch.
small branches cross the lateral sjirface of the The septal branch (ramus septalis) (Fig. 4-11),
great coronary vein or the corori'ary sinus and as found by Donald and Essex (1954) in the 125
supply the right atrium. specimens they studied, arose as follows:
The dorsal interventricular branch or caudal (1) Ventral interventricular branch, 48 per
descending coronary artery (ramus interven- cent.
tricularis dorsalis) is a continuation of the cir (2) As one of the three terminal branches of
cumflex branch in or near the" poorly defined the left coronary artery, 27 per cent;
dorsal interventricular groove. It is over 1 mm. (3) Left coronary artery, 19 per cent;
wide and about 3.5 cm. long, and has left, right, (4) Aorta, 5 per cent;
and septal branches. It is shorter than the ven (5) Circumflex branch, 1 per cent.
tral interventricular branch, as its terminal Immediately after entering the interventric
branches usually end at about the junction of the ular septum the septal branch usually runs ob
middle and distal thirds of the ventricular mass. liquely toward the apex of the heart, giving off
They may reach to the apex of the heart, or even major and minor branches along its course. Ini
extend beyond it. Usually three or four branches tially it runs obliquely to the right ventricular
supply the adjacent musculature of the left ven side of the septum. In this course it may parallel
tricle, and usually five or six larger and longer the atrioventricular fibrous ring as it lies under
branches arborize in the adjacent part of the the endocardium adjacent to the dorsal half of
right ventricle. The septal branches supply a the septal cusp of the right atrioventricular
narrow zone of the interventricular septum ad valve. The first half of the artery lies closely un
jacent to the dorsal interventricular groove. der the endocardium of the right ventricle; the
The ventral interventricular branch or cranial second half penetrates deeply into the septum.
descending coronary artery (ramus interventric- It supplies all the main papillary muscles of the
ularis ventralis) is about the same diameter as the right ventricle. According to Donald and Essex
circumflex branch, and averages 1.5 mm. in (1954), it supplies 70 to 75 per cent of the inter
width. It is about 7 cm. long as it winds obliquely ventricular septum. These authors and many
and distally from left to right across the sterno others have found that the anastomoses between
costal surface of the heart in the ventral inter the septal artery and the adjacent arteries of the
ventricular groove. It usually extends beyond canine heart are not sufficiently large or numer
the apex of the heart (Christensen 1962). It has ous to permit retrograde filling of the septal ar
left ventricular, right ventricular, and septal tery with injected dye. The dorsal and ventral
branches. interventricular branches supply the periphery
The left ventricular branches are long and of the interventricular septum. The ventral ves
large. Usually there are seven, which in general sel contributes much more blood than does the
decrease in size toward the apex. As they largely dorsal vessel (Christensen and Campeti 1959).
286 Chapter 4 The H e a rt and A rte rie s
F k;. 4 - J 1 Branches of the left coronary artery, ventral aspect Thenghl ventricle has been removed.
P ulm onary T run k 287
intrapericardial part of the vessel. The ventral, subdivides and supplies the cardiac and dia
lateral, and caudal surfaces of the proximal three- phragmatic lobes. The relations of the pulmo
fourths cf the vessel are covered by serous peri nary arteries within the lungs are described in
cardium and fat. The distal fourth of the vessel Chapter 14, Respiratory System.
serves for the attachment of the fibrous pericar
dium and can be examined without opening the
pericardial cavity. The pulmonary trunk, along PULMONARY VEINS
its entire medial surface, contacts the aorta. The
two vessels form a slight spiral as they obliquely The pulmonary veins (vv. pulmonales) are
cross each other. The ligam entum arteriosum valveless and return aerated (arterial) blood
is a connective tissue remnant of the fetal ductus from the lungs to the left atrium. Unlike the pul
arteriosus which arises near the bifurcation of monary arteries, the pulmonary veins from each
the pulmonary trunk and passes to the aorta. of the lobes of the lungs as a rule retain their sep
The right pulmonary artery (a. pulmonalis arate identity to the heart. An exception to this
dextra) is about 2 cm. long and 1 cm. in diam is frequently found in the veins from the right
eter. It leaves the pulmonary trunk at nearly a diaphragmatic and intermediate lobes, which
right angle and runs to the right, where at first it fuse just before entering the atrium. Not uncom
is in contact with the concavity of the arch of monly the vein from the left diaphragmatic lobe
the aorta and later with the right bronchus. It joins the venous trunk from the right side, a sin
lies obliquely across the base of the heart be gle opening in the left atrium thus serving for
tween the precava and postcava. Its first branch the return of the blood from the right and left
enters the apical lobe of the lung cranial to the diaphragmatic and the intermediate lobes. Other
apical bronchus (prebronchial). About 1 cm. dis variations are common; not infrequently two
tal to the origin of this branch the vessel divides veins may enter the heart directly from one of
into numerous vessels which supply the apical, the lobes. Usually, however, several veins con
cardiac, diaphragmatic, and intermediate lobes verge and empty into the heart by a single vessel
of the right lung. which is from a few millimeters to about 1.5 cm.
The left pulmonary artery (a. pulmonalis sinis in length. In medium-sized dogs all pulmonary
tra) is shorter and slightly smaller in diameter veins are over 5 mm. in diameter as they enter
than the right artery. It is partly covered dor the heart. The pulmonary veins may be divided
sally at its origin by the left bronchus. The artery into the right pulm onary veins (vv. pulmonales
then passes obliquely across the pulmonary vein dextrae) from the right lung, and the left pulmo
coming from the apical lobe and divides un nary veins (vv. pulmonales sinistrae) from the
evenly into two or more branches. The smaller left lung. The pulmonary veins within the lungs
branch or branches enter the apical lobe; the are described with the intrinsic structure of the
large one enters the bulk of the lung, where it lungs, in Chapter 14.
SY S T E M IC A R T ER IES
AORTA dally and to the left, the aortic arch (arcus aor-
tae). The remainder of the aorta, from the arch
The aorta leaves the left ventricle near the to its terminal iliac branches, is the descending
center of the base of the heart. It is a heavily aorta (aorta descendens). The descending aorta
walled vessel through which all the systemic may be divided further into a thoracic part
blood of the body passes. All of the large sys (aorta thoracica) and an abdom in al part (aorta
temic arteries arise directly from it. For descrip abdominalis).
tive purposes, it may be divided into an The ascending aorta, at its origin, is slightly
ascending and a descending portion, separated expanded to form the bulb o f the aorta (bulbus
by the aortic arch. The initial part attaches to aortae). Peripheral to each of the three semi
the fibrous base of the heart, is largely located lunar cusps which form the aortic valve are the
within the pericardium, and is known as the sinuses o f the aorta (sinus aortae) (see discussion
ascending aorta (aorta ascendens). It is about under aortic valve, p. 282). The aggregate of the
2 cm. long before it makes a U-turn dorsocau- sinuses form the bulb of the aorta. The coronary
A o r t ic A rch 289
arteries arise from the aortic bulb (see discussion The right common carotid (a. carotis com
on pp. 283 to 285). munis dextra) diverges from the left and ob
The normal development of the heart and aor liquely crosses the ventrolateral surface of the
tic arches in several domestic animals is dis trachea as it runs toward the head. Throughout
cussed by Krediet (1962), in a thesis on anoma the neck it lies in the angle formed by the longus
lies of the arterial trunks in the thorax. colli or longus capitis dorsally, the trachea ven-
tromedially, and the brachiocephalicus and
sternocephalicus laterally. At the thoracic inlet
the vagosympathetic nerve trunk becomes asso
AORTIC ARCH ciated with the dorsal surface of the artery and
remains bound to it during its course through the
A r t e r ie s of the H ea d , Neck, and T horax neck. The internal jugular vein is also associated
with the common carotid artery in the middle
The blood supply of the head and neck and half of the neck. The fascia which binds these
the thoracic limbs leaves the aorta through two structures together and attaches them rather
great vessels arising from the aortic arch, the loosely to adjoining parts is the carotid sheath. It
brachiocephalic trunk, and the left subclavian is a part of the poorly developed deep cervical
artery. fascia. The common carotid artery terminates at
or near a transverse plane through the body of
Brachiocephalic Trunk the hyoid bone by dividing into internal and ex
ternal carotid arteries. The internal carotid,
The brachiocephalic artery or trunk (truncus much smaller than the external one, leaves the
brachiocephalicus) (Fig. 4-12), the first large medial side of the parent vessel and immediately
artery from the aortic arch, passes obliquely to runs through the deep structures of the head to
the right and cranially across the ventral surface the brain. The external carotid is the main sup
of the trachea. It is about 4 cm. long and 8 mm. ply to either half of the head. The branches of
in diameter. The left common carotid artery is the common carotid arteries are the caudal thy
the first branch to leave the brachiocephalic roid occasionally, the cranial thyroid, and the
trunk. Frequently a small branch leaves the terminal external and internal carotid vessels.
brachiocephalic artery close to the heart to aid The caudal thyroid artery (a. thyroidea cau-
in the supply of the thymus and pericardium. dalis) (Fig. 4-13) is a small vessel which usually
The brachiocephalic artery terminates in the arises from the brachiocephalic between the
right common carotid and the right subclavian origin of the common carotids. It is not constant
arteries. This termination is medial to the first in its origin; it may arise from the brachioce
rib or first intercostal space of the right side. phalic, the left subclavian, or the ascending cer
The common carotid arteries (aa. carotides vical, a branch of the omocervical artery. It
communes) arise from the brachiocephalic artery occasionally comes from the costocervical trunk
about 1 cm. apart. Of 123 dogs examined, the on the right side. The most common origin is in
right and left common carotids arose from the the form of a short trunk from the brachioce
brachiocephalic artery by a common trunk in phalic, giving rise to the right and left caudal
four specimens. The interval between the origins thyroid arteries, which run cranially toward the
of the common carotids varies from 15 to less respective lobes of the thyroid gland. They lie
than 1 mm. When a bicarotid trunk (truncus bi- on the trachea and in contact with the respec
caroticus) is formed, it usually arises from the tive borders of the esophagus. Branches from the
brachiocephalic artery about 2 cm. distal to its caudal thyroid arteries are freely supplied to the
origin from the aorta opposite the first intercostal esophagus, trachea, caudal cervical ganglia, and
space or second rib. nerves in the region of the thoracic inlet. They
The left common carotid (a. carotis communis anastomose with the larger cranial thyroid
sinistra) usually arises opposite the vertebral end arteries.
of the second rib and ventral to the trachea. Its The cranial thyroid artery (a. thyroidea cra
relations are similar to those of the right vessel as nialis) (Fig. 4-13) is a short vessel which arises
it traverses the neck, except that it is on the left from the common carotid opposite the caudal
side and is loosely bound to the esophagus dorso- part of the larynx. It is the largest and the only
medially by the deep cervical fascia. Its branches constantly present branch of the common ca
and termination are similar to those of the right rotid. It has the following branches: thyroid,
vessel, which is described herei pharyngeal, cricothyroid, and muscular. The
290 Chapter 4. T he H ea rt and Ar t e r ie s
Cost o c e r v ic a l a. {V o r t e b r a l a / B r a c h i o c e p h u l i c a.
\
Com c o ro t i d oa. i /?. s u bcl avi an a / ,L. s u b c l a v i a n a.
- Longus colli m
-Descending a o rta
Brachiocephalic us m
Superf. c e r v i c a l a. - - ~ Esophagus
-Pulmonantj a
S u p ra s p in o u s a
S u p ra s c a p u la r a - A o r t i c arch
S c a le n u s m'
S u p r a s p in o tu s m.
Om o c e r y i c a l a '
--R v e n tric le
Ex t th o r a c ic a.
Descending br, Omacervico / a
- In t e r n a l th o r a c ic a.
r i c eps m.
B r a c h i a l a. T en sor fa s c ia e a n t e b r a c h i i m.
BI c eps m. -
tat 3a.
Common c a r o t i d aa.
l / e r t e b r a l a. V e r t e b r a l a.
C ostocervical C ostoce rvica l a.
O m ocervical a
m o c e rv ic a l a.
Ax i 11a ry ar
A x i l l a r y a.
Int- t h o r a c i c a.- -
In te rn a l th o ra c ic a
Ft s u b c lo v ia n a
P recava L. s u b c l a v i a n a.
B r a c h i o c e p h a l i c a. Esophagus
-A o rtic a rc h
R a u r ic le
Pulmon a r t j a
L. a u r i c l e
R v e n t r i c le -
L v e n tr ic le
- D e s c e n d i na a o rta
P o stca va Esophagus
Med. r e t r o p h a r y n g e a l - T h y r o id c a r t i l a g e
ly m p h n o d e
_ - M . c r ic o t h y r o i d e u s
C r i c o i d c a r t i la g e - _
__ -C ric o th y ro id b ra nch
M u s c u la r b ra n c h - -
---- T h y ro id b r a n c h e s
Cran. t h y r o i d a.—
-P a ra th y ro id g la n d
P haryngeal b r -
" T hyro id g l a n d
T h y ro id g la n d '
---- C ra n ia l t h y r o i d a.
— T ra c h e a
---- E s o ph a gu s
R t com m on c a r o t id a -
----- L. com m on c a r o t i d a.
R t.c a u d a l t h y r o i d o.~
- E s o p h a g e a l br.
Rt. v e r t e b r a I a -
Rt. cos t o c e r v i c a l a — ~L .ca u d a l t h y r o i d a.
Rt. int. t h o r a c i c a. -
0 r a c h io c e p h a 1 1c a. —
- ~L. s u b c l a v i a n a.
Fic. 4-13. The relation of the common carotid arteries to the larynx, trachea, and related structures, ventral aspect.
292 Chapter 4. The H e a r t an d A r te r ie s
branches vary in distribution and constancy. longus capitis and longus colli. This usually
Frequently the thyroid branches and the pha comes from the external carotid vessel near its
ryngeal and cricothyroid branches come directly origin as described under the muscular branches
from the common carotid as two separate trunks. of that vessel.
The thyroid branches (rr. thyroidei) are those The external carotid artery (a. carotis ex
which run in a caudal direction to the thyroid terna) (Figs. 4-14, 4-15) is the main continua
lobe. Their number and location vary; usually, tion of the common carotid to the head. It is
however, several branches enter the dorsal and about 4 cm. long and forms a sigmoid flexure as
ventral borders of the lobe from its middle to the it winds its way under the cranial end of the hy
cranial pole and diverge as they ramify on its poglossal nerve, submandibular salivary gland,
lateral and medial surfaces. Thus the blood sup and digastric muscle. It is bounded deeply by
ply to the thyroid lobe may be divided into the muscles of the larynx and pharynx. The fol
dorsal and ventral groups of vessels. One ramus, lowing branches leave the external carotid
usually from the dorsal group, and more often artery: occipital, laryngeal, ascending pharyn
larger than the others, extends from the cranial geal, lingual, facial, great auricular, parotid,
pole of the thyroid lobe caudally past the dorsal terminal superficial temporal, and maxillary
border of the gland. This branch continues cau arteries.
dally in association with the recurrent laryngeal The occipital artery (a. occipitalis) is most fre
nerve and anastomoses with the caudal thyroid quently the first branch of the external carotid.
artery, giving off esophageal and tracheal It may, however, arise in the angle formed by
branches along its course. When the caudal thy the splitting of the common carotid into the ex
roid artery is well developed the cranial vessel is ternal and the internal carotid. In some dogs it
reduced in a reciprocal ratio. In specimens with arises an appreciable distance out on the exter
well-developed caudal thyroid arteries most of nal carotid and therefore it will be regarded as a
the cervical parts of the trachea and esophagus branch of this vessel. It is slightly smaller than
are supplied by them. the internal carotid, measuring about 1.5 mm. in
The pharyngeal branch (r. pharyngeus) leaves diameter. Occasionally a slight bulbous enlarge
the cranial side of the cranial thyroid artery, or it ment can be detected in the vessel at its origin.
may leave in common with one of the thyroid In general, the vessel takes a tortuous course
branches. It is the smallest of the branches of the dorsally, its terminal branches anastomosing
cranial thyroid artery. It runs obliquely dorso- with those of its fellow. This occurs caudal to the
cranially and supplies twigs to the beginning of external occipital protuberance in the dorsal
the esophagus; ventrally, a twig enters the larynx straight muscles of the head. The vessel initially
in company with the recurrent nerve; continu runs dorsocranially, being crossed laterally by
ing forward, the pharyngeal branch supplies the the hypoglossal nerve and medially by the inter
constrictor muscles of the pharynx. nal carotid artery. A larger structure, which
The cricothyroid branch (r. cricothyroideus) is bears important relations to the initial part of the
a freely branching vessel which leaves the cranial vessel, is the medial retropharyngeal lymph
thyroid artery and runs cranioventrally over the node, lying caudal and partly over the vessel.
cricothyroid muscle. Twigs go to the sterno- The boundary, craniolaterally, is the digastric
hyoideus, sternothyroideus, thyrohyoideus, and muscle, and medially the last four cranial nerves,
cricothyroideus. End-twigs go through the crico although the accessory nerve may lie lateral to
thyroid membrane to the mucosa of the caudal it. Usually the first 15 mm. of the occipital artery
compartment of the larynx, where they anasto is free of branches. When it reaches the condy
mose with the laryngeal artery. Right and left loid fossa, several branches arise. The vessel then
vessels anastomose on the cricothyroid mem forms an arc around the caudal surface of the
brane. jugular process and, lying between the two
The muscular branches (rr. musculares) go nuchal lines, runs to the median plane dorsally.
dorsolaterally to both parts of the sternocephal- In this location it is covered by the brachioce
icus and the mastoid part of the brachiocephali phalicus, splenius, obliquus capitis cranialis and
cus. Only small parts of these muscles are sup the semispinalis capitis muscles. The branches of
plied by the muscular branches. The cranial the occipital artery are: occasionally a ramus to
thyroid artery also sends twigs to the capsule of the cranial pole of the medial retropharyngeal
the submandibular salivary gland and subman lymph node, a condyloid artery, a cervical ramus,
dibular and medial retropharyngeal lymph a descending ramus, a caudal meningeal artery,
nodes. A large muscular branch may go to the and occipital branches.
A r t e r ie s of th e H ea d 293
The condyloid artery (a. condylica) may arise caudalis) leaves the occipital as it lies between
from the cervical branch of the occipital or di the nuchal lines of the occipital bone. This is
rectly from the occipital. It enters the petro about 1 cm. from the base of the jugular process.
basilar fissure and, in association with the acces The vessel is about 0.7 mm. in diameter, and the
sory nerve, passes through the petrobasilar canal extracranial part is 5 mm. long. It goes through
and finally becomes dissipated in the dura at the the supramastoid foramen and ramifies in the
ventral end of the pyramid. It supplies twigs to dura of the occipital cranial fossa. Some branches
the middle and inner ear. A branch goes to the supply the tentorium cerebri just dorsal to the
digastricus before the vessel enters the fissure. pyramid.
Davis and Story (1943) describe this vessel as From the digastricus to the mid-dorsal line,
the inferior (caudal) tympanic in the canids on where the occipital artery anastomoses with its
which they worked. fellow, numerous branches arise. Most of these
The cervical branch (r. cervicalis) is the sec are dissipated in the muscles which attach to the
ond branch of the occipital. It usually is a short caudal surface of the skull. Some, however,
trunk which arises medial to the jugular process ramify in the temporal muscle and the post-
and dorsolateral to the last four cranial nerves. auricular musculature, where they anastomose
One branch descends under the wing of the with branches of the great auricular artery.
atlas, where it lies close to the bone and supplies The laryngeal artery (a. laryngea) (Fig. 4-14)
the atlanto-occipital joint capsule. It also sup is usually the second branch of the external
plies parts of the rectus capitis ventralis, obliquus carotid artery, arising ventrally nearly opposite
capitis cranialis, longus capitis, and rectus capitis the occipital artery. Sometimes this vessel, which
lateralis muscles. Some branches end by forming is less than 1 mm. in diameter, arises from the
a feeble ventral anastomosis with the vertebral common carotid at its termination. Near its
artery. Others enter minute foramina on the origin it supplies one or two small twigs to the
ventral surface of the atlas in the region of the sternomastoid muscle. Another branch runs
transverse foramen. The ascending branch runs dorsally and supplies the dorsal portions of the
medial to the last four cranial nerves as they cricopharyngeal, thyropharyngeal, and hyopha-
leave the skull. It sends minute twigs to these ryngeal muscles. The main continuation of the
nerves, as well as to the cranial cervical ganglion vessel runs ventrally with the cranial laryngeal
and the two ganglia on the glossopharyngeal nerve over the surface of the larynx and dis
and the vagus nerve. The main muscular branch appears in the triangle formed by the mm. thyro-
continues cranially past these ganglia and ends hyoideus, thyropharyngeus, and hyopharyngeus.
primarily in the longus capitis and rectus capitis It perforates the thyrohyoid membrane and sup
ventralis. Some twigs go to the mucosa of the plies most of the mucosa and intrinsic muscles of
roof of the pharynx, where they anastomose the larynx. Cranial branches have been found
with the ascending pharyngeal artery. which supply the m. hyoglossus and in which
The descending branch (r. descendens) is the they may anastomose with the lingual artery.
largest branch of the occipital. It is nearly as Caudally and ventrally, twigs ramify in the thy-
large as the continuation of the parent vessel. A rohyoideus where an anastomosis occurs with
branch enters the origin of the digastricus from the cricothyroid branch of the cranial thyroid
the proximal end of the descending branch or artery.
from the occipital artery directly. The descend A muscular branch (r. muscularis), slightly less
ing vessel takes origin about 3 mm. from the than 1 mm. in diameter, frequently leaves the
cervical branch and goes directly under the ob- dorsal surface of the external carotid at its origin.
liquus capitis cranialis to the alar notch of the Usually this origin is in the same transverse plane
atlas. Here it becomes associated with the ven as the origin of the occipital artery. It runs di
tral division of the first cervical nerve and the rectly to the ventral surface of the longus capitis,
vertebral artery and vein. The vertebral artery is on which it arborizes. Its most caudal branches
several times larger than the descending branch also supply the longus colli. In many specimens
of the occipital. An anastomosis between the the branch just described arises from the cranial
two vessels occurs. The descending branch con thyroid artery or from the external carotid di
tinues dorsomedially in the obliquus capitis cau rectly or by a short trunk with the ascending
dalis, the cranial part of which it supplies. It also pharyngeal.
supplies the cranial part of the dorsal straight The ascending pharyngeal artery (a. pharyn-
muscles of the head and the semispinalis capitis. gea ascendens) (Fig. 4-16) is a small, freely
The caudal meningeal artery (a. meningea branching vessel that arises from the external
294 Chapter 4. The H eart and A rte rie s
L a t. a u r i c u l a r , Ant. a u r i c u l a r
, Masse t e r i c
In te rm e d a u ric u la r
yPa r o t i d
M u s c u la r b r.y
sTrans facial
/
Med a u r i c u l a r D orsal p a lp e b ra l
y e n tr a l p a l p e b r a l
O c c ip it a l br. / M a la r
D orsal la b i a l
S u p e rfic ia l
o ris
Gr. a u r i c u l a r
M u s c u l a r br.
Stylo m asto i d
-O n e , u , t n ,
A s c e n d , ng
pharyn geal
Int. c a r o t i d Ant.
Ext. carotL m e n ta l
Cran th y r o id Middle mental
Com. c a ro l i a Post, men t a l
F a c ia l
Thyroid g la nd
D ic f a s tr ic u s m., c u t
Sternothyroid m. S u b lin q u a l
'r fol 'jiomiA/'
S ty lo g lo s s u s m
S t e r n o h u o id e u S rr. i H y o g lo s s u s m.
T h y r a h y o i d e u s rri. I J .Fac i n
R L a ru n je a } LinguaI
Tem p oral b ra n c h e s i M a s s e t e r ic
Middle m e n i n g e a l , Ext o p h th a lm ic
T ra n s v e r s e f a c i a l N /Ext. e th m a id a l
/ D o rs a l p a lp e b ra l
P re -a u ric u ia rx V e n tra l p a lp e b ra l
O c c ip ita l b r v X -
s Antenar deep
y te m poral
M e d ia l \_
a u ric u la r^ '
^ B u c c in a to r
Cr. a u r ic u la r - Unor p a l a t i n e
£xf c a ro tid ' Pter tjC/oid branch
'"A af p te r y g o id c a n a l
S u p e r fic ia l te m p o ra l O rb ita l
Max 11la ry I KA n a s to m o tic
M asse te n c r a m u s 1 ' P t e r y g o id b ra n c h
M a n d i b u l a r ramus Posterior deep t e m p o r a l
T y m p a n ic 1 M a n d ib u la r a lv e o la r
Fic.. 4-15. Arteries of the head in relation to lateral aspect ol the skill!
296 Chapter 4. The H e a r t an d A rte r ie s
carotid in common with or close to the occipital of about 4 cm. in this location the nerve usually
artery. When the relatively large muscular crosses the medial surface of the lingual artery
branch to the longus capitis and longus colli and then, in contact with its dorsal surface, ex
arises here, instead of from the cranial thyroid, tends to the tip of the tongue.) At the root of the
it also may be closely related to the ascending tongue two or more branches are given off which
pharyngeal or arise in common with it. Thus can be traced to the hyoid and pharyngeal mus
from a medial origin the ascending pharyngeal cles and the palatine tonsil. These are (1) the
runs dorsomedially on the pharyngeal mucosa hyoid branches (rr. hyoidei), and (2) the tonsillar
medial to the tympanic bulla. The pharyngeal artery (a. tonsillaris). The tonsillar artery leaves
branch of the vagus is the only nerve to cross its the dorsal surface of the lingual opposite the
medial surface. The artery extends as far for lateral surface of the keratohyoid bone, and runs
ward as the pharyngeal opening of the auditory dorsally rostral to it. In its course dorsocaudally
tube, where its terminal branch anastomoses it perforates the styloglossal muscle to become
with the loop of the internal carotid artery. Its related medially to the pharyngeal mucosa. The
branches are the palatine and pharyngeal. vessel then enters the tonsil near its middle by
The palatine branches (rr. palatini) are a few three or more minute branches. Twigs from the
small branches which leave the initial part of the hyoid branches enter the caudal end of the ton
ascending pharyngeal. They run ventrally in the sil and anastomose with the tonsillar artery from
lateral wall of the pharynx to the soft palate, the lingual. The twigs from the caudally lying
where they supply the extensive palatine glands, hyoid branches may be the major supply of the
and the palatine mucosa and muscles. These palatine tonsil. All other branches of the lingual
branches anastomose with their fellows, as well are destined for the supply of the muscles of the
as with the tonsillar branch of the lingual. tongue. The artery lies near the mid line, near
The pharyngeal branches (rr. pharyngei) are the ventral part of the organ. The lingual vein
distributed to the musculature and mucosa of accompanies the artery only in its anterior third.
the cranial part of the pharynx as well as to the The vein lies in a more ventral and superficial
ventral axial muscles, mainly the longus capitis. position than the artery in the remainder of the
The main ascending pharyngeal artery lies ex organ. According to Prichard and Daniel (1953),
ternal to the pharyngeal musculature directly arteriovenous anastomoses occur in the tongue
against the bulla. It sends many branches to the of the dog. The lingual and sublingual arteries
cranial part of the roof and sides of the pharynx. anastomose in the tongue.
After giving origin to these the artery continues The facial artery (a. facialis), or external max
to the external carotid foramen. Here an unusual illary (Figs. 4-14, 4-16), is about 3 cm. long and
vascular occurrence takes place. The internal 1.5 mm. in diameter. It arises near the angle of
carotid, after having traversed the carotid canal, the jaw, 1 cm. from the lingual artery, and for
forms a loop which fills the external carotid fo the first centimeter is bounded medially by the
ramen. The ascending pharyngeal artery ends styloglossal muscle; it then runs anteriorly super
by anastomosing with the internal carotid loop. ficial to the stylohyoid muscle. The masseter
In the adult domestic cat, where the internal muscle is related to it dorsally and laterally, and
carotid is not patent, the ascending pharyngeal the digastricus lies ventral to it. The artery runs
artery extends through the external carotid fo forward, but its deviation from the horizontal
ramen and contributes directly to the formation depends on the degree of closure of the jaws.
of the arterial circle at the base of the brain. This The facial artery gives rise to a glandular branch
distal part of the ascending pharyngeal in the and to muscular branches, before its first large
domestic cat is morphologically the internal collateral branch, the sublingual artery, arises.
carotid (Davis and Story 1943). The glandular branch (r. glandularis) is the
The lingual artery (a. lingualis) (Figs. 4-14, 4 - largest but not necessarily the first branch to
16, 4-18) is usually the largest collateral branch leave the initial part of the facial artery. It is the
of the external carotid. It leaves the parent trunk main supply to the mandibular and sublingual
just medial to the digastric muscle, runs cranio- salivary glands.
ventrally in company with the hypoglossal nerve, The muscular branches (rr. musculares) are
and enters the tongue medial to the hyoglossal usually two small vessels which supply the adja
muscle, and lateral to the genioglossus. (The cent parts of the digastricus, pterygoideus medi
hypoglossal nerve does not accompany the vessel alis and, occasionally, the styloglossus. The larg
during its initial intramuscular course but runs est of these branches are smaller than the glan
lateral to the hyoglossal muscle. After a course dular branch and arise anterior to it. A twig may
A r t e r ie s of th e H ea d 297
supply a small patch of mucosa in the region of masseter artery, which runs forward. It supplies
the caudal pole of the palatine tonsil. The branch mainly the orbicularis oris and levator nasolabi-
to the mucosa, when present, usually anasto alis.
moses with the ascending pharyngeal. The great auricular artery (a. auricularis
The sublingual artery (a. sublingualis) arises magna) (Figs. 4-14, 4-15, 4-16) arises at the
from the facial medial to the ventral part of the base of the annular cartilage from the dorsocau-
body of the mandible, in the depths of a deep dal surface of the external carotid. It circles
cleft which is bounded laterally by the masseter, around the caudal half of the base of the ear. It
medially by the caudal part of the mylohyoideus, is a medium-sized vessel which at first lies under
and ventrally by the digastricus. The sublingual the parotid salivary gland, then is located more
artery parallels the medial surface of the mandi dorsally under the postauricular group of mus
ble near its ventral border and is distributed cles. For convenient exposure of the origin of
largely to the mylohyoid and the anterior belly the great auricular it is necessary to remove the
of the digastric muscle, although some branches digastric muscle which lies caudoventral to it.
perforate the mylohyoideus and supply the Its branches may vary considerably in origin and
genioglossus and geniohyoideus. It is accompa disposition. They are: (1) stylomastoid, (2) glan
nied by a satellite vein and the mylohyoid nerve. dular, (3) muscular, (4) lateral auricular, (5) inter
It anastomoses with the lingual and ventral labial mediate auricular, (6) deep auricular, (7) medial
arteries. Near the middle of the body of the auricular, and (8) occipital.
mandible the subm ental artery (a. submentalis) The stylom astoid artery (a. stylomastoidea) is
is given off. This runs to the ventral surface of the smallest branch of the great auricular. It
the symphysis of the mandible, supplying this leaves the posteroventral surface of the vessel
region and the incisor teeth. Other branches are and runs directly to the stylomastoid foramen in
distributed to the muscles and mucosa in the company with the facial nerve, which it sup
region of the frenulum of the tongue. plies. Sometimes the vessel is double. It is lo
The ventral labial artery (a. labialis ventralis) cated directly caudal to the external acoustic
arises about 1 cm. from the ventral border of the process.
mandible anterior to the masseter muscle. As it The glandular branches (rr. glandulares) go to
runs anteriorly it lies along the ventral border of the parotid and mandibular salivary glands.
the orbicularis oris. Some fibers of this muscle These may arise not from the great auricular di
may actually cover the artery during its course rectly but from its muscular or lateral auricular
forward. At the posterior mental foramen it branches. The mandibular branches enter the
anastomoses with the posterior mental artery. dorsal surface of the gland; the parotid branches
The ventral labial artery sends twigs across the enter the deep surface of the parotid salivary
ventral border of the mandible which anasto gland.
mose with the sublingual artery. The muscular branches (rr. musculares) are
The angular artery o f the mouth (a. angularis one or two large vessels which supply the cranial
oris) arises from one to several centimeters pe parts of the brachiocephalicus and sternocephal
ripheral to the origin of the ventral labial. It icus. They also supply the skin, platysma, and
takes a rather tortuous course to the commissure subcutaneous fat, and finally anastomose with
of the lips, where usually one branch extends to the omocervical artery. The muscular branch lo
the dorsal and the other to the ventral margin. cated at a deeper level runs under the sterno
The angularis oris supplies in part the buccina cephalicus and brachiocephalicus and supplies
tor, orbicularis oris, and the skin and mucosa of the cranial end of the splenius muscles. Occa
this region. It anastomoses with the dorsal and sionally branches supply the medial retropha
ventral labial arteries, and may anastomose with ryngeal lymph node. The dorsal deep part of the
the posterior mental artery. muscular branch anastomoses with the occipital
The dorsal labial artery (a. labialis dorsalis) is artery.
the termination of the facial artery and ramifies The lateral auricular artery (a. auricularis lat
on the cheek and nose. Small branches may ex eralis) arises from the muscular branch to the
tend dorsally to the orbit and anastomose with splenius or from the intermediate auricular. It is
the terminal branches of the ventral palpebral a large artery which branches as it passes
and malar arteries; others run forward in the or through or in contact with the caudal border of
bicularis oris and anastomose with the lateral the parotid salivary gland. It extends distally on
nasal artery. Fine twigs, following the buccal the caudal surface of the auricular cartilage,
nerves caudally, anastomose with those of the near its lateral border. It usually extends beyond
298 Chapter 4. The H e a rt and A r te r ie s
M id d le rn e n m g e a l\ Subl i ngual
\
P te r yg oi d br anc h Facial
T y m p a n ic ~ Masseteric a
Cr. a u r i c u l a r - - . Preauricular
M u s c u la r b r - — '' G l a n d u l a r br.
Lat a u r i c u l a r " Facial
Palatine branches - "
S p h e n o p a l a t i nc
In f r a o r b i t a l
In f r a o r b i ta l
D or^o1 nasal
M a x i I l or y. ( ,L a t e r a l n a s a l
A n t septal
Minor palatine
M a jo r
I
M a jo r p a l a i m e
-S tylo g lo s su s m.
-T o n s il l o r a
Root o f tongue,
h t s td e
K eratohyo-d bone
B a s ih y o id Done
H yoid b ra n c n
Gen loalossus m, cut
S u b lin g u a l a: 'H y o q lo J s u s m
S tq log loss us m
J ty lo q lo s s u s m
M a n d ib u la r a lv e o la r a s
M a x i l l a r y a. tG eniohyoideus m
1 G cm o a lo ssu s m,
G l a n d u l a r b r s SN
N v
Foci a I a -> '
\
Ton si 11 a r a.
R. l i n y u o l a.
Hyoy I o ss u s rr ^ s
tiy o p h a n ijn y e u s
La r y n c j e a l a v A n t,m e n ta l a.
x liy o c jlo s s u s m.
Fir,. 4 -1 9 . The mandibular alveolar artery and intermamlibular structures. ventrolateral aspect
300 Chapter 4. The H e a rt and A r te r ie s
the cutaneous pouch 1 cm. or more and termi diately enters it. It freely branches in the gland.
nates by anastomosing with a branch of the in Although the periphery of the parotid gland is
termediate auricular artery. supplied by parotid rami from adjacent vessels,
The interm ediate auricular artery (a. auricu such as the great auricular and superficial tem
laris intermedia) is the largest artery to the ear. poral, its main supply in the dog is the parotid
It arises about 1 cm. peripheral to the lateral artery. This vessel sends twigs to the facial nerve
auricular deeply under the postauricular mus and the most dorsal mandibular lymph node,
cles. During its initial course toward the apex of and may supply the skin.
the ear it sends branches to the postauricular The superficial temporal artery (a. temporalis
muscles. After emerging it sends many anasto superficialis) (Figs. 4-14, 4-15) is the smaller of
mosing branches both laterally and medially the terminal branches of the external carotid. Its
over the auricular cartilage. Many twigs pass diameter is approximately 1.5 mm., compared
through the foramina in the auricular cartilage with 4 mm. for the maxillary, the other terminal
to its concave surface. branch. It arises in front of the base of the auric
The deep auricular artery (a. auricularis pro ular cartilage and at first extends dorsally. As i
funda) usually arises independently from the crosses the zygomatic arch it makes a sweepin
great auricular peripheral to the origin of the in curve anteriorly and, about 1 cm. above it, dip
termediate auricular. Occasionally it arises from under the heavy, deep temporal fascia. Durin
the intermediate or medial auricular. It is a small part of its subsequent course toward the eye
vessel which runs distally about 2 cm. and passes actually lies in the temporal muscle. Opposil
through the space between the tragus and the the orbital ligament the superficial temporal pe
anthelix to supply part of the dermis of the car forates the deep temporal fascia and divides ini
tilaginous external acoustic meatus. its two terminal branches, which lie in the supe
The m edial auricular artery (a. auricularis me ficial fascia. The branches of the superficial ten
dialis), about the same size as the lateral auricu poral artery are the (1) masseteric, (2) transvers
lar, arises about 1 cm. from the considerably facial, (3) anterior auricular, (4) temporal, (E
larger intermediate auricular. It crosses the cau dorsal and (6) ventral palpebral arteries.
dal part of the temporal muscle under cover of The masseteric artery (a. masseterica) is a rel
the auricular cartilage. At the scutiform cartilage atively large branch, usually over 1 mm. in diam
it becomes subcutaneous and continues along eter, which arises from the anterior side of th<
the anterior border of the cartilage to within 2 superficial temporal near its origin or from th<
cm. of the apex of the ear. It anastomoses freely maxillary artery directly. Hidden by the parotic
with the intermediate auricular artery and the salivary gland, it runs forward and enters the
anterior auricular branch of the superficial tem deep surface of the masseteric muscle where it
poral. Branches also perforate the cartilage and passes anteroventrally between the muscle anc
extend around its margin to supply the fascia the masseteric fossa. Usually several other fine
and dermis of the concave surface. branches arise from the vessel and supply other
The occipital branch (r. occipitalis) is the main structures. In about half of the specimens they
peripheral continuation of the great auricular come off separately from the superficial tempo
after the auricular branches are given off. It en ral close to the masseteric artery. Some twigs
ters the caudal part of the temporal muscle and enter the parotid salivary gland and parotid
at first nearly parallels the dorsal nuchal line. It lymph node. Others run forward on the face
supplies a large caudal part of the temporal mus with the dorsal and ventral buccal branches of
cle and finally anastomoses with the posterior the facial nerve and anastomose with arterial
deep temporal artery. Branches from this or a branches of the dorsal labial; still other branches
separate vessel from the great auricular also sup supply the skin and occasionally the temporo
ply parts of the postauricular muscles and anas mandibular joint capsule.
tomose with the ascending cervical branch of The transverse fa c ia l artery (a. transversa
the omocervical artery. faciei) is no larger than the nutrient branches
The parotid artery (a. parotis) is a small vessel which accompany the buccal nerves. It usually
which arises, 5 to 15 mm. distal to the origin of arises distal to the masseteric from the anterior
the great auricular, from the dorsal surface of border of the superficial temporal. It emerges
the external carotid artery as this artery crosses from under the parotid salivary gland, usually
the ventral end of the annular cartilage; it may after the artery has divided. One branch follows
arise from the great auricular artery. Thus, the the zygomatic branch of the facial nerve, and
vessel arises under the parotid gland and imme the other runs parallel and ventral to the zygo
A r t e r ie s of th e H ead 301
matic arch in company with the auriculotempo convenience in describing its branches, it may
ral branch of the trigeminal nerve. be divided into three parts: the mandibular por
The anterior auricular branch (r. auricularis tion, the pterygoid portion, and the pterygopala
anterior) arises distal to the transverse facial on tine portion. The mandibular portion extends to
the opposite or caudal side of the superficial the alar canal. The pterygoid portion lies in the
temporal. It is larger than the transverse facial, canal, and the pterygopalatine portion extends
but less than 1 mm. in diameter. It runs between from the alar canal across the pterygopalatine
the upper anterior part of the parotid gland and fossa. No branches arise from the vessel as it
the temporal muscle. It supplies both of these passes through the alar canal.
and finally ends in the preauricular muscles near The first part, or m andibular portion, o f the
the tragus. maxillary artery includes that part of the artery
The tem poral branches (rr. temporales) arise from the point where the superficial temporal
from the distal half of the superficial temporal. leaves the external carotid to the alar canal. It
These are variable in number, size, and origin. begins at the base of the ear, where the vessel
Usually two to five branches leave the dorsal sur reaches its most dorsal level and is covered by
face of the vessel and are distributed to the sub the parotid salivary gland. It continues the arch
stance of the temporal muscle. From the ventral formed by the external carotid forward and
surface of the vessel an average of two dissect- downward to the caudal border of the mandible,
able rami are present. These also supply the tem where it is bounded laterally by the masseter
poral muscle. Some branches run medial to the muscle. On reaching the mandible the artery
zygomatic arch and then ventral to it, to supply changes its course and runs medially, lying
the masseter. The larger temporal branches against the caudal part of the temporomandibu
anastomose with the deep temporal arteries of lar joint capsule as it does so. It actually follows
the maxillary. the ventral border of the retroglenoid process
The dorsal palpebral artery (a. palpebrae dor closely, and since this border is convex the ar
salis), about 1 mm. in diameter, is about twice as tery also makes a ventral arch, lying, as it makes
large as the ventral palpebral artery. It arises op the arch, on the pterygoid muscles. Before en
posite the orbital ligament and, by a tortuous tering the alar canal the vessel is embraced by
course at the junction of the upper eyelid and the mandibular division of the trigeminal nerve
frontal bone, extends toward the medial canthus dorsally, and the chorda tympani ventrally. The
of the eye. It freely branches along its course, first part ends by making a bend forward and
sending twigs to the various structures which entering the alar canal. The following vessels
form the upper eyelid. Twigs also supply the leave the first part of the maxillary artery: man
muscles, fascia, and the skin covering the tem dibular branch, mandibular alveolar, posterior
poral muscle and the subcutaneous part of the deep temporal, tympanic, pterygoid branch, and
frontal bone. It forms an anastomosis with the middle meningeal.
small arteries that leave the dorsal part of the The mandibular branch (r. mandibularis) is
orbit and with those that perforate the skull the main supply to the caudal part of the tempo
through small foramina in the region of the romandibular joint capsule. Sometimes two or
frontonasomaxillary suture. three branches are present, instead of one. The
The ventral palpebral artery (a. palpebrae branch or branches leave the dorsal surface of
ventralis) sends branches to the caudal half of the maxillary 5 to 15 mm. distal to the origin of
the lower eyelid. Several branches pass ven- the superficial temporal. When more than a sin
trally across the zygomatic arch and masseter. gle vessel is present, they are small and thread
Here in the superficial fascia these branches like.
anastomose with the transverse facial and malar The m andibular alveolar artery (a. alveolaris
arteries. The palpebral arteries are the terminal mandibularis) (Fig. 4-19) measures slightly over
branches of the superficial temporal artery. 1 mm. in diameter and enters the mandibular
The (internal) maxillary artery (a. maxillaris) canal after a course of about 1 cm. It arises from
(Figs. 4-15, 4-16, 4-17, 4-20, 4-22) gives off the ventral surface of the first part of the maxil
many branches which supply the deep struc lary artery. Sometimes a trunk is formed from
tures of the head lying outside of the braincase. which the mandibular alveolar and posterior
It is the larger of the two terminal branches of deep temporal arise in common. After entering
the external carotid and, in a medium-sized dog, the mandibular canal the alveolar artery of the
measures about 4 mm. in diameter. It is the mandible closely follows the ventral border of
main continuation of the external carotid. For the bone. It runs from the mandibular foramen to
302 Chapter 4. The H e a rt and A r te r ie s
To n a s o la c r im a t d u c t
M id d le s e p t a l
A n a s to m o s is with max 111ary s m u s
v e n tr a l p a lp e b r a l a.
To m a x illo tu rb in a te
A n a s to m o s i s with K l a t e r a l n a s a l aa.
trans. fo c i a I a.
D o rs al n a s al
M a la r
To o r b i t a l g l a
I n f r a o r b i ta l
Ma x i I lari
M in o r p a la t in e
Ta s o f t p a l a t e
Sphenopalatine
M a jo r p a la ti Ant. septal branches
Post d o r s a l alveolar To m a xilla ry sinus ethmoturbmate
A c c e s s o ry p a l a t i n e ' M id d le dorsal alveolar
F ig. 4-20. Scheme of the terminal brunches of the maxillary artery, lateral aspect.
Dors, la b io l -
A n g u la r is a r i s -
Ant. mentol
Fa cia l
the middle mental foramen. During its course in masseteric nerve through the mandibular notch
the mandible it sends many small twigs through to the masseter muscle. This is the masseteric
the apical foramina to the roots of the teeth branch (r. massetericus). It anastomoses with the
(Boling 1942) and others to the bone itself. The masseteric artery of the superficial temporal,
mandibular nerve is dorsolateral in the mandib which is the main supply to the masseter, as it
ular canal. The artery is in the middle, and the runs on its deep surface. The posterior -deep
vein is ventromedial to the artery. Usually a con temporal artery is accompanied by a satellite
siderable amount of fat surrounds these struc nerve and vein or veins.
tures. Three vessels continue anteriorly from the The tym panic artery (a. tympanica) is a small
mandibular alveolar to supply the anterior part inconstant branch of the maxillary artery. It may
of the lower jaw. These are the posterior, mid arise from the posterior deep temporal artery. It
dle, and anterior mental arteries. usually leaves the maxillary medial to the tempo
The posterior m ental artery (a. mentalis pos romandibular joint and enters one of the small
terior), with its satellite nerve and vein, leaves foramina located in a depression medial to the
the posterior mental foramen and runs to the joint. It courses through the temporal bone into
lower lip. It is much smaller than the middle the middle ear. Davis and Story (1943) describe
mental artery, with which it anastomoses. It also a similar vessel for the cat under the name of
anastomoses with the ventral labial artery. tympanica anterior.
The middle m ental artery (a. mentalis media) Branches leave the ventral surface of the max
is the largest of the three mental vessels and is illary posterior to its entrance into the alar canal
the main blood supply to the anterior part of and arborize in the medial and lateral pterygoid
the lower jaw. It leaves the middle mental fo muscles. Only small posterior portions of the
ramen, which is located in the ventral half of the pterygoid muscles are supplied by this source.
mandible, ventral to the first two cheek teeth. Twigs also supply the origins of the tensor and
With its accompanying vein and nerve it sup levator veli palatini, the pterygopharyngeus, the
plies the skin, tactile hair follicles, and other soft palatopharyngeus, and the mucosa of the nasal
structures. It forms an anastomosis with the an pharynx.
terior and posterior mental arteries. It is the The middle meningeal artery (a. meningea
main continuation of the alveolar artery of the media) (Fig. 4-15) leaves the dorsal surface of
mandible. the maxillary before this vessel enters the alar
The anterior m ental artery (a. mentalis ante canal. It is about 1 mm. in diameter and runs
rior) is the smallest of the three mental arteries. through the oval foramen, which is closely ad
It leaves the mandibular alveolar less than 1 cm. jacent to the maxillary artery. A notch, or still
caudal to the middle mental foramen and, with more rarely a foramen (foramen spinosum), is
its satellite vein and nerve, runs in the delicate formed in the anterior wall of the oval foramen
incisivomandibular canal, which closely follows for the passage of the vessel. Within the cranial
the ventral border of the body of the mandible cavity the middle meningeal artery gives off the
to the anterior mental foramen. It anastomoses ramus anastomoticus which runs medially and is
with its fellow of the opposite side, as well as about equal in size to the parent artery. The ra
with the middle mental artery. mus anastomoticus joins the anastomotic artery
The posterior deep temporal artery (a. tempo of the external ophthalmic. After giving off the
ralis profunda posterior) arises from the ventral ramus anastomoticus, the middle meningeal ar
surface of the maxillary just distal to or in com tery follows the vascular groove on the cerebral
mon with the mandibular alveolar. It immedi surface of the skull. It runs in company with two
ately crosses the lingual, mylohyoid, and alveo satellite veins along the lateral border of the su
lar branches of the trigeminal nerve, as well as ture between the petrous and squamous parts of
the lateral pterygoid muscle. It enters the tem the temporal bone. It then passes almost directly
poral muscle and extensively arborizes in it. It dorsally across the middle part of the brain case
also sends rami which accompany the mylohy and bifurcates into anterior and posterior
oid and lingual nerves. Most of the branches, branches. At its termination along the mid-dor-
however, are confined to that part of the tem sal line it anastomoses with its fellow of the op
poral muscle lying medial to the coronoid proc posite side. The middle meningeal artery is the
ess. It forms anastomoses with the anterior deep largest of the meningeal arteries. Its branches
temporal, the occipital branches of the great leave the parent vessel at right angles and run
auricular, and the temporal branches of the su both anteriorly and posteriorly. They supply the
perficial temporal. One branch passes with the dura and adjacent portions of the skull.
304 Chapter 4. The H e a r t an d A r te r ie s
The ramus anastom oticus enters the cavern Usually there are dorsal and ventral muscular
ous sinus and makes two to four loops before branches which supply the muscles of the eye
joining the anastomotic artery from the external ball. These arteries arise independently from the
ophthalmic lateral to the hypophysis. anterior surface of the external ethmoidal artery,
The second part, or pterygoid portion, o f the but occasionally they arise by means of a com
maxillary artery is about 1 cm. long, lies in the mon trunk from the external ethmoid or directly
alar canal, and gives off no branches. from the orbital artery. The external ethmoidal
The third part, or pterygopalatine portion, artery, after passing through the ethmoidal fora
lies on the lateral side of the lateral pterygoid men, reaches the dura. In the dura, between the
muscle and crosses it obliquely. The following cribriform plate and the olfactory bulb, it di-
vessels leave the pterygopalatine portion of the videTinto a dorsal and a ventral branch. These
maxillary artery: orbital, artery of the pterygoid branches anastomose anteriorly and form an
canal, pterygoid branches, anterior deep tempo arterial circle on the lateral wall of the ethmoidal
ral, buccinator, minor palatine, and infraorbital fossa. Many small branches leave this arterial
arteries, and a trunk which gives rise to the ma circle and reunite, so that an ethmoidal rete is
jor palatine and sphenopalatine arteries. formed. The internal ethmoidal arteries, from
The orbital artery (a. orbitalis) (Figs. 4-15, 4 - the anterior cerebral arteries, run in the falx
22, 4-23) is the short vessel (it is wider than it is cerebri to the cribriform plate, where they anas
long) that gives rise to the vessels supplying the tomose and aid in forming the ethmoidal rete.
orbit and anastomosing with the vessels inside Many branches pass through the cribriform plate
the skull. Tandler (1899) originated the term from the rete to supply the mucosa of the eth
which Davis and Story (1943) and Jewell (1952) moturbinates and the nasal septum. Those to the
adopted in their works. According to these au nasal septum are the posterior septal arteries (aa.
thors the orbital artery has no homologue in septales posteriores). Another branch, the an
man, although Ellenberger and Baum (1943) call terior meningeal artery (a. meningea anterior),
it the external ophthalmic artery in the rumi runs dorsally in the dura at the caudal margin of
nants and the horse. The orbital artery arises the cribriform plate, passes through the inner
from the dorsal surface of the maxillary immedi table of the frontal bone, and enters the muco-
ately after it leaves the alar canal. The orbital periosteum on the floor of the major compart
artery is bounded medially by the maxillary ment of the frontal sinus. After running caudally
nerve, laterally by the zygomatic and lacrimal in the frontal sinus it passes through the inner
nerves. The zygomatic and lacrimal nerves, table of the frontal bone 5 to 10 mm. from the
as well as the terminal part of the orbital mid-sagittal plane and arborizes in the dura
artery, are encased within the periorbita. The ventral to the caudal part of the frontal sinus. It
orbital artery typically divides into the exter forms a delicate anastomosis with the middle
nal ethmoidal and external ophthalmic ar meningeal artery. There is considerable varia
teries. tion in the size and distribution of the branches
The external ethm oidal artery (a. ethmoidea of the external ethmoidal artery.
externa) (Figs. 4-22, 4-24) is the anterior branch If a common trunk of origin is formed for the
of the orbital artery. It is homologous with the muscular branches, it is usually short. Each of its
anterior and posterior ethmoidal arteries of man resultant branches dips between adjacent recti
(Davis and Story 1943). It makes an initial curve muscles to the fat which lies between these and
anterodorsally across the lateral surface of the the retractor bulbi. Many branches are dispersed
ocular muscles, where it runs through the plexus to the recti and oblique muscles of the eye and
formed by the ophthalmic vein in this location. to the eyeball itself. The zygomatic and lacrimal
It sometimes gives off branches to the dorsal ob arteries, which are small, arise from the dorsal
lique muscle and to the frontal bone. It then muscular branch.
makes one or two more bends and enters the The ventral muscular branch (r. muscularis
larger, more dorsally located ethmoidal foramen ventralis) (Fig. 4-23) extends toward the globe
in company with its small satellite vein. The en of the eye between the ventral and lateral recti,
trance of the artery into the ethmoidal foramen although some branches pass to the medial side.
is unusual in that the vessel enters it from above The muscles it supplies are primarily the lateral
and in front and not from the side from which and ventral recti and the ventral portions of the
the vessel approaches it. Also peculiar is the fact retractor bulbi. It also supplies the medial rectus,
that a separate, smaller and more ventral eth the gland of the third eyelid, and the conjunctiva
moidal foramen conducts the ethmoidal nerve. of the lower eyelid near the fornix. One small
A r t e r ie s of the H ea d 305
arterial branch runs with a branch of the oculo leave the external ophthalmic artery. About 15
motor nerve to the ventral oblique muscle. It mm. before reaching the eyeball, this artery
anastomoses with the dorsal muscular branch forms one or two flexures in the fat on the dorsal
and the ciliary arteries. surface of the optic nerve. It runs to the medial
The dorsal muscular branch (r. muscularis side of the optic nerve, where it anastomoses with
dorsalis) of the external ethmoidal artery arises the smaller internal ophthalmic artery. From the
in common with or 1 cm. from, the ventral mus union of the external and internal ophthalmic
cular branch, which it exceeds slightly in size. It arteries, two to four ciliary arteries (aa. ciliares)
crosses the proximal third of the lateral rectus arise and run to the eyeball. On reaching the
obliquely and passes between the lateral and sclera which surrounds the optic nerve, the
dorsal recti toward the globe of the eye. In its vessels break up into several branches which ex
course it sends branches to the lateral and dorsal tend through the cribriform area and ramify in
recti, dorsal oblique, retractor bulbi, and levator the choroid part of the vascular coat as the cho
palpebrae muscles. On reaching the globe of the roid arteries (aa. choroideae). Other branches do
eye it gives off one small branch which extends not perforate the sclera in the cribriform area but
dorsally over the eyeball and ends in the bulbar continue, closely applied to the sclera, toward
conjunctiva under the upper eyelid. In its course the cornea, and are called the episcleral branches
it supplies the terminal part of the levator palpe (rr. episclerales). As pointed out by Tandler
brae muscle and a portion of the lacrimal gland. (1899), and confirmed by Jewell (1952), the
This part is comparable to the supraorbital artery central artery o f the retina (a. centralis retinae)
of man, whereas the dorsal muscular branch rep arises from the arc formed by the anastomoses
resents the superior muscular set of vessels in of the two ophthalmic arteries which pass into
man. As the dorsal muscular branch crosses the the optic nerve and run anteriorly to the cribri
lateral rectus it divides into lacrimal and zygo form area of the sclera. Here the vessel branches
matic branches, which follow the respective repeatedly and emerges around the periphery
nerves. The arterial twigs supplying the muscles of the optic papilla as nine or more arterioles
of the orbit are peculiar in that they run centrif- which radiate into the retina. According to
ugally. The main arteries run deeply in the mus Catcott (1952), the venules which lie in the
cular cone and issue their fine branches pe retina are three or four in number and con
ripherally. verge to the center of the optic papilla. Great
The lacrimal artery (a. lacrimalis), larger than variation exists among dogs and between the
the zygomatic artery, accompanies its satellite eyes of the same dog. The central artery of the
nerve and supplies the lacrimal gland. It passes retina is the main supply to the retina.
deep to the orbital ligament and terminates in The anastomotic artery (a. anastomotica)
the conjunctiva and skin of the upper eyelid. (Fig. 4-22) leaves the external ophthalmic, the
The threadlike zygomatic artery (a. zygomat- orbital, or even the maxillary artery, according
ica) follows the zygomatic nerve to the lacrimal to Jewell (1952), close to the orbital fissure which
gland, the skin, the conjunctiva near the lateral it traverses. It sends minute twigs to the dura
canthus of the eye, and the adjacent lower eye and to the nerves which pass through the orbital
lid. The lacrimal and zygomatic arteries may fissure. Sometimes the vessel is double through
anastomose with each other. The lacrimal occa out part or all of its course. It enters the cavern
sionally joins the dorsal palpebral; the zygomatic ous sinus and receives the ramus anastomoticus
usually unites with the ventral palpebral. from the middle meningeal artery. It continues
The external ophthalm ic artery (a. ophthal- posteriorly as a single tortuous vessel and unites
mica externa) (Fig. 4-22) is the caudal, smaller with the internal carotid at a transverse plane
branch of the orbital. The splitting of the orbital which passes through the dorsum sellae. Thus it
into the external (Simoidal and the external is possible for blood to pass from the maxillary
ophthalmic arteries is variable. Rarely it may artery to the internal carotid, or vice versa, by
occur at their origins from the maxillary artery the orbital and anastomotic arteries.
so that anterior and posterior trunks are formed. The pterygoid branch (r. pterygoideus) of the
In such instances the posterior trunk gives rise to maxillary artery is a freely branching muscular
the ramus anastomoticus. Peripheral to the twig to the medial and lateral pterygoids. It is
origin of this vessel it extends between the about 0.5 mm. in diameter and arises opposite
medial and dorsal recti to the fat and retractor the origin of the anterior deep temporal artery.
bulbi muscle. The origin of the retractor bulbi Its origin is about 2 mm. peripheral to the origin
muscle receives most of its blood from rami which of the orbital artery, but it usually arises from
l i a r y ao.
*Ext. e th m oi dal a.
/
/A n t. m e n in g e a l br.
D o r s a l muse
- - E x t o p h t h a l m i c a.
Ext. e t h m o i d a l
V entral m u s c u la r - - I n t . o p h t h a l m i c a.
y - - In t . e th m o id a l a.
A n a s ta m a tic - Ant. c e r e b r a l a.
M a x i I l a r y a-
----- M i d d l e c e r e b r a l a.
O rb ita l fis s u r e ~ - Int. c a r o t i d a.
~~Post. c o m m u n i c a t i n g O.
A n a s to m o tic ram u s ~ - P o s t, c e r e b r a l a.
M a x i l l a r y a. " - A n t c e r e b e l l a r a.
in a l a r c a n a l
M i d d l e m e n in g e a l a . ' / " ~Bast la r a
F o r a m e n o v a le '
Fir:. 4-22. Arteries of tlie orbit and base of the cranium, dorsal aspect.
,M re c tu s d o r s a l i s
M le v a to r palpebrae
Dorsal m u s c u la r br.
M. r e t r a c t o r bulbi
lExf. e th m o id a l a.
L a c r im a l g l a n d /Ext. o p h th a lm ic a.
i /
I /Int. o phthalm ic a.
M. r e e f us l a t e r a l is -
, A n t c e re b r a l a.
M .obliquus v e n t r a l i s - M iddle c e r e b r a l a.
M a la ra .
Middle meningeal a
i I l a r y a.
' A n a s to m o tic a
B r to m. r e c t u s m e d i o l i s
1O r b ita l a.
V e n t r a l m u s c u la r br.'
iAnt. deep te m p ora l a■
Fit.. 4-23. Arteries of the orbit and extrinsic ocular muscles, lateral aspect.
A r t e r ie s of th e H ea d 307
A n t m e nin g ea l
Cut ds of criib r i f a r m p la te
L ext. ethmoidal a.
V e n t ra l br. -
Falx c e r e b r i ( c u t ) -
In t e thm o idal a . '
Vent. br. r ext. ethmoidal a .' Lat. n a s a l a.
V om er' / \R .ce ntral in c is a r
/ 1
P a la tin e ' ' In c is iv e
i
Middle septal a. Isphenopalatine) 1 A n t septal bn (major palatine)
M a x illa r y a.
M in o r p a l a t in e a.
Sphenopalatine ay
M a jo r p a l a t in e a ' Ant. se pta l b r
I 1
M id d l e se p ta l a./ 'Major p a la t in e a.
Loc a tio n a l m a x i l l a r y s in u s ' 1Periosteum
I
Ato m a x illa r y sinus r ethmaturbinate 1 1A.to m axilloturbinate
Fit.. 4-25. A dissection showing arteries of the lateral nasal wall.
308 Chapter 4. The H e a rt and A r te r ie s
the opposite side. It may arise from the medial major palatine arteries (Figs. 4-17, 4-25) arises
side of the orbital artery. Several branches sup from the maxillary anterior to the origin of the
ply both the lateral and medial pterygoid mus minor palatine artery. It usually has a single, but
cles. Occasionally a twig can be traced through sometimes a double, muscular ramus to the an
the muscle into the pterygoid canal. The ptery terior portion of the medial pterygoid muscle.
goid branch anastomoses with the muscular The muscular ramus may arise from the maxil
branch of the buccinator artery, which supplies lary or from one of the terminal branches of the
part of the medial pterygoid. common trunk.
The anterior deep temporal artery (a. tem The m ajor palatine artery (a. palatina major)
poralis profunda anterior) (Fig. 4-15) is a vessel arises from the common trunk as one of its
less than 1 mm. in diameter which arises close to terminal branches. The vessel, which is slightly
the orbital artery. It may be double. From the more than 1 mm. in diameter, passes through
dorsal surface of the maxillary it runs dorsally the posterior palatine foramen and the palatine
between the temporal muscle and the caudal canal with a delicate vein and relatively large
part of the frontal bone. The small anterior deep satellite nerve. Within the palatine canal the
temporal artery enters the temporal muscle near nerve and artery divide so that two or more sets
the middle of its anterior border and arborizes in of major palatine arteries and nerves emerge on
the muscle. Accompanied by two satellite veins, the hard palate. The main channel draining the
it forms an anastomosis in the temporal muscle area of the hard palate does not follow the major
with the superficial and caudal deep temporal palatine artery through the palatine canal but
arteries. runs caudally in the soft tissue of the hard palate
The buccinator artery (a. buccinatoria) (Fig. as a spongy, poorly developed venous plexus.
4-15) arises from the ventrolateral surface of the The plexus continues in the soft palate, where it
maxillary about 1 cm. distal to the origin of the lies dorsal to the palatine glands. It empties into
anterior deep temporal. It is nearly 1 mm. in di the maxillary vein caudal to the temporomandib
ameter as it leaves the maxillary at an acute ular joint, ventrolateral to the tympanic bulla.
angle and runs toward the cheek. Usually near Some of the nerve and artery branches pass
its origin a small branch is given off to the medial through the accessory palatine foramina located
pterygoid muscle. It soon becomes related to the caudal to the major palatine foramen. The
buccinator nerve which accompanies it to the arteries anastomose with each other and the most
cheek. A tiny twig is given off to the ventral por caudal branch anastomoses with the minor pala
tion of the orbital gland, and larger twigs are tine. The most anterior branch is the main con
distributed to the masseter, temporal, and buc tinuation of the major palatine artery. The pala
cinator muscles. The vessel finally terminates in tine groove on the surface of the hard palate, in
the region of the soft palate and the pterygo which the vessels lie, is situated midway between
mandibular fold. the alveoli and the mid line. Anastomoses be
The minor palatine artery (a. palatina minor) tween the right and left palatine vessels occur
(Figs. 4-17, 4-25) arises from the ventral surface throughout their course. The major palatine
of the maxillary or one of its terminal branches artery and nerve usually leave the palatine
dorsal to the last upper cheek tooth. It is less groove midway between the palatine fissure and
than 0.5 mm. in diameter and passes ventrally the major palatine foramen. They extend through
through a notch in the caudal part of the maxilla. the palatine venous plexus in their anterior
It is distributed to the adjacent soft and hard course so that they lie closely under the oral
palate. The branch to the soft palate runs nearly mucosa. The groove anterior to the plane in
the whole length of this part and lies close to the which the artery and nerve leave it contains a
mid plane. It supplies the palatine glands, mus portion of the palatine venous plexus. The major
culature, and mucosa. Fine twigs anastomose palatine artery supplies the mucosa of the oral
with the ascending pharyngeal and the major surface of the hard palate, the periosteum, and
palatine arteries. Occasionally a branch of the the bone which forms the alveoli. A small branch
minor palatine artery sends a twig to the orbital passes through the palatine fissure and anasto
gland. moses with a branch of the sphenopalatine
The infraorbital artery and a common trunk artery, which supplies the mucosa on the nasal
which gives rise to the sphenopalatine and major side of the hard palate. A small artery extends
palatine arteries are the terminal branches of the anterolaterally, passes through the interdental
maxillary. They are nearly equal in size. space between the canine and corner incisor
The common trunk o f the sphenopalatine and teeth, and anastomoses with the lateral nasal
A r t e r ie s of the H ead 309
artery. The anterior septal branches (rr. septi goes to the dorsal part of this bone and has an
anteriores) (Fig. 4-24) from the major palatine extensive anastomosis with a vessel which runs
artery run dorsomedially and supply that part of anteriorly in endoturbinate I from the ethmoidal
the septum caudal to the area supplied by septal rete.
branches of the lateral nasal and anterior to the The branch which goes to the maxilloturbinate
area supplied by the middle septal artery from is short, medially inclined, and variable in origin.
the sphenopalatine. By an extensive, fine arterial It may come from either the dorsal or the ven
plexus they anastomose with adjacent vessels. tral branch previously described. It goes to the
The major palatine artery continues forward, caudal part of the maxilloturbinate crest and di
branching profusely, and in back of the incisor vides into five or six small arteries which arbor
teeth turns toward the mid line and anastomoses ize on the primary scrolls into which the bone is
with its fellow. At the anastomosis a small vessel divided. It anastomoses anteriorly with a branch
runs dorsally through the incisive foramen and of the lateral nasal artery, which curves around
joins with the right and left lateral nasals as these the dorsal part of the nostril and extends pos
anastomose with each other at the mid plane. teriorly on the ridge of tissue which is continu
This anastomotic branch is small as it passes ous with the maxilloturbinate crest. In addition
dorsally through the interincisive suture. to the ventral, dorsal, and maxilloturbinate parts
The sphenopalatine artery (a. sphenopalatina) of the posterior lateral nasal arteries, smaller
(Fig. 4-25), which is the other terminal branch of branches supply the mucosa and bone of the
the common trunk, is over 2 mm. in diameter maxillary sinus and the anterior parts of the eth-
and leaves the pterygopalatine fossa by passing moturbinates and a large part of the middle of
through the sphenopalatine foramen with its the nasal septum. The twigs to the maxillary
satellite nerve and vein. On reaching the naso sinus arise from the posterior side of the dorsal
pharyngeal duct the artery runs anteroventrally branch of the posterior lateral nasal as this vessel
under the mucoperiosteum and on the dorsal runs in the mucoperiosteum which forms the
surface of the palatomaxillary suture to a point anteroventral and anterolateral parts of this cav
ventral to the opening into the maxillary sinus. ity. A twig to the caudal part of the maxillary
Here the sphenopalatine artery swings dorsoan- sinus may leave the sphenopalatine shortly after
teriorly for a few millimeters and divides into a it enters the nasopharyngeal duct. Many other
dorsal and a ventral branch, and a branch which branches supply the mucoperiosteum of the
goes to the maxilloturbinate. All of the terminal floor and sides of the ventral nasal meatus. The
branches of the sphenopalatine artery are col m iddle septal artery (a. septi media) is the first
lectively known as the posterior lateral nasal ar branch of the sphenopalatine artery after it
teries (aa. nasales posteriores laterales). leaves the sphenopalatine foramen. It runs be
As the main sphenopalatine vessel makes its tween the mucoperiosteum and the plate of
dorsal bend, the ventral vessel continues forward bone separating the nasopharyngeal duct and
to supply the mucoperiosteum of the side and the nasal fundus, to the middle part of the nasal
floor of the nasal fossa and the adjacent middle septum. Anteriorly it anastomoses with the ante
portion of the nasal septum. A small artery leaves rior septal branches from the major palatine and
the dorsal surface of this vessel and, curving dor- posteriorly it anastomoses with the posterior sep
socaudally, runs toward the eye on the nasolacri tal branches from the ethmoidal rete. All the
mal duct. This twig supplies blood to the anterior branches of the sphenopalatine and the middle
part of the duct and anastomoses with the twig of septal arteries form voluminous arterial plexuses
the malar artery, which supplies its caudal part. in the mucoperiosteum which they supply. Nu
Beyond the origin of the small artery to the naso merous anastomoses also occur between adja
lacrimal duct, the ventral vessel continues for cent vessels.
ward and slightly medially; its terminal twigs The infraorbital artery (a. infraorbitalis) (Figs.
anastomose with a branch of the major palatine, 4-17, 4-20, 4-21) is the main continuation of
which ascends through the palatine fissure. the maxillary across the medial pterygoid mus
The dorsal branch arises near the opening into cle. Accompanied by the maxillary division of
the maxillary sinus aligned with a transverse the trigeminal nerve, it leaves the pterygopala
plane passing between the third and fourth tine fossa, gives off the posterior dorsal alveolar
upper premolar teeth. This vessel runs dorso- artery, and passes through the maxillary foramen
anteriorly and bifurcates: one branch supplies to enter the infraorbital canal. It gives off a
the ventral part of the nasal turbinate and the branch to the orbital gland, posterior dorsal alve
mucoperiosteum lateral to it; the other branch olar, malar, middle dorsal alveolar, and anterior
310 Chapter 4. The H e a rt and A r te r ie s
Int. e t h m o i d a I a. O p t i c n.
I n t o p h t h a l m i c a. - _ _ Ant. c e r e b r a l a.
- M i d d l e c e r e b r a l a.
Ant. i n t e r c a r o t i d a.
A n a s t o m o t i c ra m u s of
'-Int. c a r o t i d a. w i t h i n
m i d d l e men i n q e a l a. ■gpf venous ca v e rn o u s s in u s
A n a s t o m o t i c a. ~
( f r o m ext. o p h t h a l m i c > 4 M m __ H y p o p h y s i s
- - S u p e r f i c ia l la y e r of d u ra
P o s t h y p o p h y s e a l aa - P a s t i n t e r c a r o t i d a.
I nt. c a r o t i d a Deep l a y e r o f d u r a
O pti nt. o p h t h a l m i c a.
A n t i n t e r c a r o t i d a. Ant. c e r e b r a l a.
--M id d le c e r e b r a l a.
Infundibular r e c e s s - Int.: c a r o t i d u.
C u t in fu n d ib u la r s t a l k ''''
M a m m illa r y body
A n t. c e r e b e l l a r a . '
Abducens Oculomotor n.
O p h t h a lm ic b r of n V s Post, c e r e b r a l a.
A n astom otic a
^ A n t. c e r e b e l l a r a
(from ext.ophtho I mi c ) "
M a x illa r y br: of a V ~ ^ M id d le meningeal o.
A n a s to m o tic ra m u ~ 'i te a r n.
o f middle m e n in g e a l a.
Abducens n
M a n d ib u la r b r of n .V - ~ _• T r i g e m in a l n
Middle m e n in g e al F a c ia l n.
I n t c a r o t id a. _ - A c o u s t i c n.
A coustic, a. — Accessory n.
Fic 4-28 Dorsal aspect of the base of the skull showing arteries and nerves. The dura is partially removed on the left side, open
ing the cavernous sinus.
312 Chapter 4. The H e a rt and A r te r ie s
dorsal alveolar. It terminates by dividing into the branches and anastomoses with branches of the
lateral and dorsal nasal arteries. These terminal dorsal labial. It first crosses under the maxillo-
arteries arise either before or after the vessel has nasolabialis muscle and then runs among the fi
passed through the infraorbital foramen (Chris bers of the orbicularis oris. The vessel branches
tensen and Toussaint 1957). profusely and supplies the upper lip and snout
The posterior dorsal alveolar artery (a. alveo- as well as the follicles of the vibrissae, or tactile
laris dorsalis posterior) is a small vessel which hairs. It furnishes blood to the anterior part of
may arise from the minor palatine or either of the upper lip and the adjacent part of the nose.
the terminal branches of the maxillary. It usually At the philtrum the vessel anastomoses with its
arises from the ventral surface of the infraorbital fellow and sends a relatively large branch dor
before the latter enters the infraorbital canal. sally between the nostrils, and another branch
The posterior dorsal alveolar divides and runs di posteriorly in the mucosa of the parietal carti
rectly to the alveolar canals of the last two molar lage.
teeth. These are minute arterial branches ac The dorsal nasal artery (a. dorsalis nasi) is less
companied by satellite nerves and veins. than 0.5 mm. in diameter and travels anterodor-
The m alar artery (a. malaris) arises from the sally across the lateral surface of the nose to its
dorsal surface of the infraorbital after this vessel dorsal surface. It runs under and supplies the
enters the infraorbital canal. It passes dorsocau- levator nasolabialis muscle, then it continues to
dally into the orbital fossa. Near its origin a small supply the structures of the dorsal surface of
branch is given off, which supplies the ventral the anterior half of the muzzle. It anastomoses
oblique muscle and passes along its deep surface with its fellow of the opposite side, as well as
to anastomose with the ventral muscular branch with the lateral nasal, ventral nasal, and pos
of the orbital or external ethmoidal artery. The terior lateral nasal arteries, and their middle
main trunk runs to the medial canthus of the eye septal branches.
superficial to the periorbita. During its course it The internal carotid artery (a. carotis interna)
gives off a delicate branch which enters the nasal (Figs. 4-16, 4-22, 4-29) arises with the external
fossa in company with the nasolacrimal duct. carotid as the smaller of the two terminal
The terminal twigs go mainly to the lower eye branches of the common carotid. Other arteries,
lid, where they anastomose with the ventral pal which arise in close association with this vessel,
pebral and the transverse facial artery. are the occipital and ascending pharyngeal. The
The m iddle dorsal alveolar branches (rr. alve- termination of the common carotid is directly
olares maxillares media) leave the ventral surface medial to the medial retropharyngeal lymph
of the infraorbital artery as it runs through the node which is bound to the artery and adjacent
infraorbital canal. They run short distances with structures by the fascia which forms the carotid
their satellite nerves and veins and enter the al sheath. At a still more lateral level is the sterno-
veolar canals of the three roots of the shearing, cephalic muscle. The internal carotid artery at
or last premolar, tooth. first runs dorsoanteriorly across the lateral sur
The anterior dorsal alveolar arteries (aa. alveo- face of the pharynx. At its origin, from the dorsal
lares dorsales anteriores) consist of one relatively surface of the parent artery, is a bulbous enlarge
large branch, which enters the incisive canal, ment, the carotid sinus (sinus caroticus), which
and two or more smaller branches, which en is about 3 mm. in diameter and 4 mm. long. The
ter the maxilla anterior to the infraorbital fora vessel then narrows to approximately 1 mm. The
men. The smaller twigs supply the alveoli and internal carotid gives off no branches before en
possibly the roots of the second and third pre tering the petrobasilar fissure. Just before enter
molar teeth. The main branch arches over the ing this depression it crosses the lateral surface
root of the canine tooth and terminates in the of the cranial cervical sympathetic ganglion and
roots of the incisors. Thus this artery which is the medial surface of the digastric muscle. The
accompanied by its satellite vein and nerve has artery enters the posterior carotid foramen in
an intraosseous course throughout its length. It the petrobasilar fissure and traverses the carotid
supplies the first premolar, the canine, and in canal. On leaving the internal carotid foramen,
cisor teeth of the same side. which is the anterior opening of the carotid
The lateral nasal artery (a. lateralis nasi) (Figs. canal, it passes ventrally through the external
4-17, 4-21) is the larger of the two terminal carotid foramen, forms a loop, and re-enters the
branches of the infraorbital. It measures slightly cranial cavity through the same foramen. Fre
more than 1 mm. in diameter at its origin at the quently a small twig from the ascending pharyn
infraorbital foramen. It runs forward into the geal artery anastomoses with the loop formed by
muzzle with many large infraorbital nerve the internal carotid. On re-entering the cranial
A r t e r ie s of th e H ead 313
cavity the internal carotid runs at first obliquely arteries and provides alternate routes by which
toward the dorsum sellae, then directly anterior blood can reach the brain.
to the optic chiasm. The vessel first perforates Several anterior hypophyseal arteries (aa. hy
one layer of the dura mater and runs a short dis pophyseos anteriores) leave the posterior com
tance in the blood-filled cavernous sinus which municating artery and run over the tuber cinere-
separates the dura into two layers. It then per um to the stalk of the hypophysis. These, with
forates the second layer of dura and the arach their fellows, supply the major portion of the
noid, and comes to lie in the subarachnoid space. gland. The pars nervosa, however, is supplied by
On entering this space it trifurcates as the mid the posterior hypophyseal artery, a branch of the
dle cerebral, anterior cerebral, and posterior posterior intercarotid artery.
communicating arteries. A small anterior inter The middle cerebral artery (a. cerebralis
carotid artery arises from the trifurcation. While media) (Figs. 4-30, 4-31) is the largest vessel
in the cavernous sinus the internal carotid artery which supplies the brain. It leaves the internal
forms an anastomosis with the anastomotic artery carotid as a terminal branch about 1 mm. from
of the external ophthalmic. the origin of the posterior communicating. It lies
The posterior intercarotid artery (a. intercarot- at first on the anterior perforated substance,
ica posterior) (Fig. 4-26) is a small vessel which where it gives rise to the choroid artery (a. cho-
leaves the first part of the internal carotid as it roidea). This enters the lateral ventricle at the
enters the cavernous sinus. The vessel runs ob apex of the piriform gyrus, circles around the
liquely toward the mid line and joins with its cerebral peduncle with the hippocampus, and
fellow, posterior to the hypophysis. It is closely supplies the vessels of the choroid plexus of the
applied to the dura of the cavernous and inter- lateral ventricle. The middle cerebral then
cavernous sinuses. It gives off a branch which crosses in front of the piriform lobe and divides
perforates the dura surrounding the hypophysis into at least two large branches, which supply
and supplies the posterior lobe. This is the pos the whole cortex of the lateral surface of the
terior hypophyseal artery (a. hypophyseos pos cerebral hemisphere. The vessels follow the sulci
terior). Occasionally the posterior intercarotid in some places, and run over the gyri in others.
artery arises from the anastomotic artery. Terminal cortical branches (rami corticales), in
The posterior comm unicating artery (a. com- the form of minute twigs, enter the cortex and
municans posterior) (Figs. 4-27, 4-28) leaves the richly supply it. The central branches (rami
caudal surface of the internal carotid after it centrales) leave the middle cerebral near its
perforates the dura and arachnoid and enters the origin in the form of several branches which sup
subarachnoid space. It forms the lateral third of ply the basal nuclei and adjacent tracts. The
the arterial circle. Caudally it anastomoses with middle cerebral artery anastomoses with the an
the posterior cerebral artery. It is readily identi terior and posterior cerebral arteries.
fied by the fact that the third cranial nerve The anterior cerebral artery (a. cerebralis an
crosses its dorsal surface. terior) (Fig. 4-32) arises lateral to the optic
The arterial circle of the brain (circulus arteri chiasm and runs dorsal to the optic nerve in an
osus cerebri) (Figs. 4-27, 4-28) is an elongated anteromedial direction. On reaching the longi
arterial ring on the ventral surface of the brain, tudinal fissure it unites with its fellow. This side-
formed by the right and left internal carotids and to-side union of right and left anterior cerebral
the basilar artery. From the arterial circle, on arteries is usually about 2 mm. long, after which
each side, arise three vessels which supply the the two vessels separate. In some specimens
cerebrum. These are the anterior, middle, and there is an arterial bridge rather than a broad
posterior cerebral arteries (Fig. 4-33). The an union connecting the right and left vessels.
terior cerebellar arteries from the circulus arteri When an arterial bridge is present, it is called
osus cerebri, and the posterior cerebellar arteries the anterior communicating artery (a. communi-
from the basilar artery supply the cerebellum; cans anterior). The anterior cerebral artery runs
pontine and medullary branches of the basilar dorsally to the genu of the corpus callosum, turns
supply the pons and medulla oblongata. All of backward along the corpus callosum, and anas
these vessels form anastomoses with adjacent tomoses with the posterior cerebral which
vessels on the surface of the brain. A rich capil comes into the longitudinal fissure from behind.
lary network is found in the cortex, whereas the Numerous tortuous and freely branching vessels
white matter of the brain has a less abundant leave the dorsal and anterior surfaces of the an
supply. The arterial circle ensures the mainte terior cerebral artery. These extend dorsally to
nance of constant blood pressure in the terminal the lateral sulcus, where they anastomose with
314 Chapter 4. The H e a rt and A r te r ie s
D o r s a l br.
Post meningeola. in
tronsverse canol
(from o c c ip ita l a.)
V e n t r a l br.^
Hypoglossal n. (XII)
\Acoustic a.
Int. e thm oid al a
Ant. c e r e b r a l a.1 s \T r ig e m in a l n. (V)
Abducens n. (VI)
O p tic n./Ill 1
1 \ \lnt. ca ro tid a. in cavernous sinus
In t.o p h th a lm ic a.i
, ' Oculom otor n. (Ill)
M id d l e c e re b ra l a.1
Post, com m unicating a.
Fic. 4-29. \ parasagittal section of the cranium, showing internal arteries and nerves.
A r t e r ie s of the B r a in 315
Int. e t h m o i d a l a.
Int. o p h th a lm ic a
, O p tic n.
Ant. c e r e b r a l a.^
„ - M i d d l e c e r e b r a l a.
Int. c a r o t i d a.
Post, c o m m u n ic a tin g a.
Post, c e r e b r a l a.~
— Oculomotor n.
T r o c h le a r n - -
---- P o s t c e re b ra l a.
Ant. c e r e b e l l a r a .— --P o n tin e br
B a s i l a r a. - -
----- T r iy e m in o l n.
Abducens n -
- — F a c i a l n.
A c o u s t i c n .-'
"A c o u s tic a
G lo s s o p h a ry n g e a l n."
Post, c e r e b e l l a r a.
V o y u s n.' ,
Accessory n
C e r e b r o s p in a l a " I n t e r v e r t e b r a l fo ra m en o f a tla s
/
C e r v i c a l n.I,vent. r o o t ' 'Transverse foramen of a t l a s
2nd in te r v e r te b r a l fo ra m e n '
Ramus s p in a lis II
Ram us s p i n a l i s I I I '
V e ntra l s p in a l a
V e r t e b r a l a.
Fic. 4-30. Arteries of the brain and cervical spinal cord, ventral aspect.
316 Chapter 4. The H eart and A r te r ie s
the middle cerebral. The area of their distribu well developed, lies ventrolateral to the thoracic
tion to the dorsal surface of the hemisphere is inlet. The right subclavian artery arises medial
largely anterior to the lateral sulcus. The anterior to the first right intercostal space and is about 2
cerebral artery, like the middle vessel, not only cm. long. The left subclavian artery arises medial
supplies the cortex with its cortical branches, to the left third intercostal space and is about 6
but also sends branches into the medullary sub cm. long. Since the four arteries which arise
stance. Farther ventrally, toward the olfactory from each subclavian have similar origins and
bulbs, they are confined to the longitudinal distributions only a single description of them
fissure. will be given. All arise medial to the first rib or
The internal ophthalmic artery (a. ophthal- first intercostal space.
mica interna) (Fig. 4-22) is homologous with the The vertebral artery (a. vertebralis) (Figs. 4 -
ophthalmic of man. It is less than 0.5 mm. in di 30, 4-34) is the first branch of the subclavian. It
ameter, and leaves the anterior cerebral artery. arises from the dorsal surface on the ventro
It follows the dorsal surface of the optic nerve lateral side of the trachea. As it ascends to the
through the optic canal, and may be double. transverse foramen of the sixth cervical verte
As the artery travels anterolaterally with the bra, it crosses the trachea obliquely on the right
optic nerve, it passes from the dorsal surface to side (the trachea and esophagus on the left side),
the medial surface of the nerve. At a location ap and the longus colli and the lateral surface of the
proximately 15 mm. posterior to the bulbus transverse process of the sixth cervical vertebra
oculi, the internal ophthalmic artery anastomoses on each side. Near the thoracic inlet, on each
with the external ophthalmic artery. From the side, a small muscular branch supplies the peri
anastomosis of the two vessels three or four tracheal fascia and, running forward, terminates
ciliary arteries and the central artery of the in the caudal part of the longus capitis. The
retina arise. Their distribution is discussed in vertebral artery, before entering the transverse
connection with the description of the external canal at the sixth cervical vertebra and at every
ophthalmic artery. intervertebral space cranial to this, sends dorsal
The internal ethm oidal artery (a. ethmoidalis and ventral muscular branches (rami muscu-
interna) (Fig. 4-24) is a small artery which arises lares) into the adjacent musculature. The dorsal
from the ventral part of the anterior cerebral branches are distributed to the scalenus, inter
artery and runs toward the cribriform plate. It transversarius colli, serratus ventralis cervicalis,
lies near the attached portion of the falx cerebri, and omotransversarius. The ventral branch sup
where it parallels the medial olfactory stria. plies mainly the longus capitis and longus colli,
Upon reaching the most anterior portion of the although some twigs go to the brachiocephalicus
ethmoidal fossa it anastomoses with the ventral and sternocephalicus. Arising as a rule sepa
branch of the external ethmoidal artery, forming rately, but occasionally from a short common
a rete. Most of the branches of the internal eth trunk, the dorsal and ventral branches follow the
moidal artery pass through the cribriform plate dorsal and ventral divisions of the corresponding
to supply the ethmoturbinates and the nasal spinal nerve. They are accompanied by satellite
septum. Branches on the nasal septum anasto veins. According to Whisnant et al. (1956), four
mose with the middle septal branch of the sphe or five anastomoses occur between the muscular
nopalatine as well as with the anterior septal branches of each vertebral artery and the costo-
branch of the major palatine arteries. cervical artery. These workers indicate that a
secondary anastomosis exists between the verte
Subclavian Artery bral and omocervical arteries. These combined
anastomoses are large enough to sustain life in
The subclavian artery (a. subclavia) (Figs. 4 - the majority of dogs when both the vertebral and
35, 4-36) is the intrathoracic portion of the par common carotid arteries are ligated bilaterally at
ent vessel to each pectoral limb. It arises on the the base of the neck.
left side from the arch of the aorta and on the From the medial surface of the vertebral
right side as a terminal branch of the brachio artery, usually opposite the muscular branches,
cephalic artery. It is continued at the cranial arise the first seven cervical .spinal branches
border of the first rib on each side by the axillary (rami spinales). These enter the spinal canal at
artery. The name subclavian implies that the each of the first seven cervical intervertebral
vessel lies under the clavicle. This is not the case foramina. Within the spinal canal each divides
in quadrupeds, because the thorax is laterally into a small dorsal and a slightly larger ventral
compressed so that the clavicle, even when it is branch. The ventral branches are all united
S u b c l a v ia n Artery 317
through the medium of the ventral spinal artery it sends at least one large branch caudally into
(a. spinalis ventralis) (Fig. 4-30). This is an un the thoracic part of the serratus ventralis and
paired vessel which lies at the ventral median mainly two or three smaller branches cranially
fissure and extends along the length of the spinal into the cervical part of the serratus ventralis.
cord. It sends segmental twigs into the ventral The transversa colli at the proximal end of the
median fissure to the gray matter of the spinal first rib inclines dorsocaudally, crosses the lateral
cord. Other branches, lying in the pia, partially surface of the first costotransverse joint, and
encircle the spinal cord and supply the ventral arborizes extensively in the dorsal part of the
and lateral white matter. The dorsal branches of thoracic portion of the serratus ventralis. It gives
the spinal arteries follow the dorsal nerve root to origin to the eighth cervical spinal branch as it
the spinal cord, where they are dissipated with passes the intervertebral foramen. In some speci
out a continuous dorsolateral trunk being mens this branch arises from that part of the
formed. The largest spinal ramus is usually the vessel which supplies the thoracic part of the
third cervical, but occasionally the fourth cervi serratus ventralis. During its course it obliquely
cal spinal branch equals it in size. The vertebral crosses the deeply lying deep cervical artery.
artery divides unequally into a large dorsal and a The deep cervical artery (a. cervicalis pro
small ventral branch (Fig. 4-34). The ventral funda) is the dorsocranially extending terminal
branch anastomoses with the small cervical branch of the costocervical trunk. It leaves the
branch of the occipital artery ventral to the wing thorax at the proximal end of the first intercostal
of the atlas. The larger dorsal branch, at the space. A medium-sized vessel usually leaves the
transverse foramen of the atlas, anastomoses parent artery here and, extending dorsally, ar
with the descending branch of the occipital. The borizes mainly in the semispinalis capitis in the
branch of the vertebral which enters the spinal region of the withers. The main part of the deep
canal by passing through the intervertebral fora cervical runs craniomedially to the median
men of the atlas becomes the cerebrospinal ar plane, supplying along its course the deep struc
tery (a. cerebrospinalis) (Fig. 4-30). It perforates tures of the neck, particularly the semispinalis
the dura and arachnoid and divides into the cra capitis, multifidus cervicis, longissimus capitis,
nially running cerebral branch (ramus cerebralis) spinalis et semispinalis thoracis et cervicis, and
and the caudally running spinal branch (ramus the terminal fasciculus of the thoracic portion of
spinalis). Right and left cerebral branches anas- the longissimus. It anastomoses with the dorsal
tamose to form the large basilar artery (a. basi- muscular branches of the vertebral artery and, in
laris) which runs along the ventral surface of the the cranial part of the neck, with the descend
brain stem and is the largest source of blood to ing branch of the occipital artery. It gives origin
the brain via the circulus arteriosus cerebri. to the first thoracic spinal branch.
The costocervical trunk (truncus costocervi- The supreme intercostal artery (a. intercostalis
calis) (Figs. 4-35, 4-36) arises from the sub suprema) leaves the costocervical trunk in the
clavian artery 5 to 10 mm. peripheral to the proximal end of the first intercostal space. It ex
origin of the vertebral artery. Since it courses tends caudally to the third and occasionally the
dorsally and the vertebral courses cranially, the fourth intercostal space, where it anastomoses
costocervical on the left side crosses first the with the dorsal (aortic) intercostal artery of that
lateral surface of the vertebral artery, then the space. It passes through the costotransverse fo
esophagus; on the right it crosses the trachea. ramen, the space formed by the neck of the rib
On either side the vessel crosses the longus laterally and the vertebra medially. Usually the
capitis muscle and terminates as it enters the vessel is smallest in the third intercostal space
first intercostal space by dividing into the small, and therefore the anastomosis may be regarded
caudally running a. intercostalis supreina and to be at this site, but this is variable. Caudal to
the larger, dorsally running a. cervicalis pro each of the ribs it crosses, it sends a small dorsal
funda. On either side the vessel lies largely intercostal vessel ventrally, which anastomoses
medial to the first rib and has only one collateral with the intercostal vessels from the first two or
branch. three intercostal spaces and with the ventral
The transverse artery o f the neck (a. transversa intercostal arteries of the internal thoracic in the
colli) arises from the cranial surface of the costo third and/or fourth intercostal spaces. The small
cervical trunk, at an acute angle, at about the second and third, and occasionally the fourth,
middle of the medial surface of the first rib. It thoracic spinal branches arise from the supreme
runs mainly dorsally and leaves the thoracic intercostal.
cavity in front of the first rib. From its initial part In addition to the three main branches of the
318 Chapter 4, The H e a rt and A r te r ie s
Int. e t h m o i d a l a - ' N C e r e b ro s p in a l a.
In t. o p h t h a l m i c a. NB a s i l a r o
M id d le c e re b ra l a ' s Post, c e r e b e l l a r a .
,T h a l a m u s
/
/ R ig h t p o s t c e re b ra l o
C o rp u s c a l lo s u m y
R i g h t a n t. c e r e b r a l ak
L e f t a n t c e r e b r a l a -^
z C e re b ro s p in a l
/
L. i n t. e t h m o i d a l a . ' '
B a s i l a r a.
L int. o p h t h a l m i c a /
v Post, c e r e b e l l a r a
L. a n t c e r e b r a l a \ \
i i , A c o u s t i c a.
L . m i d d l e c e r e b r a l a .1 1
i VL. a n t. c e r e b e l l a r a.
L.int. c a r o t i d o . 1 sL . p o s t. c e r e b r a l a. (c u t)
' L p a s t . c o m m u n i c a t i n g a.
Fic 4-32- Arteries of the cerebellum and medial surface ot cerebrum.
V ertebra l Artery 319
Ramus s p in o h s II
C e re b ro s p in a l a
M u s c u la r b r .
-V e rte b ra l a
Ramus s p i n a l i s V III
V e r t e b r a l a. „
Int. c a r o t i d a.
C o s t a c e r v i c a l a. 1C o m m o n c a r o t i d a
Braehiocephah
costocervical trunk, smaller vessels exist. A con the scapula, on which it ramifies. It supplies the
stant branch, the first intercostal artery (a. inter supraspinatus and sends one large and several
costalis prima) leaves the costocervical at its small nutrient arteries into the bone. Passing
origin and, extending under the pleura covering across the distal end of the spine of the scapula,
the first two or three intercostal spaces and the it sends branches to the infraspinatus, teres
intervening ribs, supplies the principal inter minor, and the shoulder joint. Near the caudal
costal vessels of these spaces. The dorsal portions border of the scapula, it anastomoses with the
of its intercostal branches anastomose with the circumflex scapular artery. The medial branch of
smaller dorsal intercostal arteries from the su the suprascapular passes between the subscapu
preme intercostal. The ventral portions anasto lar muscle and the medial surface of the neck of
mose with the ventral intercostal arteries from the scapula. It supplies both of these as well as a
the internal thoracic. part of the shoulder joint. It ends in an anasto
Small branches leave the costocervical trunk mosis with the circumflex scapular artery.
near its origin and supply the adjacent muscula The ascending cervical artery (a. cervicalis
ture. Usually a small ramus accompanies the ascendens) leaves the omocervical peripheral to
common carotid artery a short distance up the the origin of the suprascapular. Frequently the
neck. The costocervical trunk and its branches artery is double. The profusely branching
are accompanied by satellite veins. ascending cervical is considerably smaller than
The omocervical artery (a. omocervicalis) the suprascapular. It courses cranially, medial to
(Figs. 4-36, 4-37) is more nearly homologous the brachiocephalicus and lateral to the scalenus.
with the thyrocervical trunk of man than is It supplies the sternocephalicus, the cervical
any other artery. It arises from the cranial sur portions of the brachiocephalicus, rhomboideus,
face of the subclavian medial to the first rib and omotransversarius, scalenus, and the prescapu
opposite the origin of the internal thoracic artery. lar lymph nodes. In the cranial half of the neck
It is a long meandering artery which lies in the its terminal branches are distributed chiefly to
angle between the shoulder and the neck. It lies the omotransversarius and brachiocephalicus.
dorsal to the pectoral, brachiocephalic, and omo- Some branches anastomose with the cervical
transverse muscles, and ventral to the brachial branch of the great auricular artery.
plexus. It has five named branches in addition to The supraspinous artery (a. supraspinatus) is
several small muscular twigs to the muscles given off the omocervical 1 cm. or less peripheral
which lie adjacent to it. to the origin of the ascending cervical, and is
The descending branch (ramus descendens) usually larger than the latter. The supraspinous
arises from the omocervical artery about 3 cm. artery circles around the cranial border of the
from its origin. Occasionally the descending supraspinatus and goes into its lateral surface,
branch arises from the internal thoracic artery being largely distributed by ramifying proximally
instead of from the omocervical. It runs disto- through it. A few terminal branches, however,
laterally on the brachium in the groove between reach as far as the infraspinatus. In some speci
the pectoral muscles, which bound it caudally, mens a small branch extends distally over the
and the brachiocephalicus, which bounds it tendon of the supraspinatus and the major
cranially and partly covers it. It ends in the distal tubercle of the humerus, and anastomoses with
third of the brachium in either the superficial the descending branch. Under the brachial part
pectoral, brachiocephalicus, or biceps brachii. of the brachiocephalicus deep branches anasto
Peripheral to the origin of the descending mose with the suprascapular artery.
branch, the omocervical sends one or more The superficial cervical artery (a. cervicalis
branches to the muscles which lie ventral to the superficialis) is the terminal part of the omo
trachea. cervical. It is a continuation of the parent artery
The suprascapular artery (a. suprascapularis) after the supraspinous artery has been given off.
leaves the caudal side of the omocervical about It runs in the space between the shoulder and
2 cm. distal to the origin of the descending neck, caudal to the prescapular lymph nodes,
branch. Accompanied by the suprascapular which it supplies. It reaches the ventrocranial
nerve, it goes through the triangular space border of the cervical part of the trapezius. At
bounded by the subscapularis, supraspinatus, this point it divides into an ascending and a de
and pectoralis profundus. On reaching the neck scending branch.
of the scapula it divides into a large lateral and a The ascending branch usually becomes super
small medial branch. The lateral branch passes ficial, sending many branches dorsocranially into
under the supraspinatus to the lateral surface of the superficial fascia and the cutaneous muscles
T horax 321
which cover the cleidocervicalis. This branch precardial mediastinum from the internal tho
varies greatly in development; when it is fully racic arteries. Those to the cardiac and post-
developed it may anastomose with the cervical cardial mediastinum perforate the origin of the
branch of the great auricular artery in the cranial overlying transverse thoracic muscle and extend
third of the neck. It may remain relatively deep, vertically to either the pericardium or the dia
supplying the superficial muscles of the neck. phragm. Coming from the various phrenic arter
The descending branch passes under the cervi ies are mediastinal branches which extend for
cal part of the trapezius to which it sends many ward into the ventral part of the mediastinum.
branches. It terminates in the muscle near the Other twigs from the phrenic arteries ramify in
cranial angle of the scapula. The branches of the the plica venae cavae.
omocervical are accompanied by satellite veins. The perforating branches (rami perforantes)
The internal thoracic artery (a. thoracica in are straight, short, ventrally directed branches
terna) (Figs. 4-36, 4-38) leaves the caudoventral which leave the ventral surface of the internal
surface of the subclavian opposite the origin of thoracic artery. One is present in each inter
the omocervical. It runs caudoventrally in a nar chondral space except the first and the last
row, lateral pleural plica from the precardial (eighth), and occasionally the seventh. These lie
mediastinum to the craniomedial border of the close to the lateral surfaces of the sternebrae and
transversus thoracis. Lying parallel to the ster give off sternal branches (rami sternales) to them.
num, it passes under the transversus thoracis and The perforating branches also supply the internal
runs caudally above the sternal ends of the costal intercostal and pectoral musculature adjacent to
cartilages and the intervening interchondral them. They are continued subcutaneously near
spaces. It ends just inside the costal arch at the the sternum as the ventral cutaneous branches
thoracic outlet by dividing into the small muscu (rami cutanei ventrales) along with their satellite
lophrenic and the large cranial epigastric artery. veins and comparable nerves. The twigs which
The pericardiacophrenic artery (a. pericardia- supply the medial portions of the thoracic mam
cophrenica) (Fig. 4-39) is a small vessel which mary glands are called mammary branches (rami
leaves the caudal side of the internal thoracic mammarii). These, present only when the glands
artery near its origin and runs with the phrenic are developed, come from the fourth, fifth, and
nerve to the pericardium. In addition to supply sixth vessels.
ing the precardial, intrathoracic part of the The ventral intercostal arteries (aa. inter
phrenic nerve, the left vessel may send one or costales ventrales) (Fig. 4-38) usually are double
more branches to the precardial mediastinum for each of the interchondral spaces, starting
and the thymus, when this is well developed. At with the second and ending with the eighth.
the level of the heart on both sides the peri Starting with the caudal artery of the eighth
cardiacophrenic artery anastomoses with the space, all remaining arteries come from the mus
branch from the musculophrenic artery which culophrenic artery. Those from the ventral sur
courses cranially on the nerve from the dia face of the internal thoracic arise singly, so that
phragm. Twigs leave the vessel to supply the a small artery lies on each side of the costal carti
pericardium. A ventral bronchial branch may lages, except the first, which has a delicate single
course to the root of the left lung. artery. The artery caudal to the cartilage is
The main thymic branches (rami thymici) slightly larger than the one cranial to it, and is
usually leave the precardial parts of the internal accompanied by the intercostal nerve, in addi
thoracic artery as it passes through the thymus. tion to the satellite vein. These double arteries
Usually a single thymic branch supplies each anastomose with each other across the ribs. The
lobe, but more than one vessel may be present. ventral intercostal artery lying caudal to the rib
The bronchial branches (rami bronchiales) anastomoses with the dorsal intercostal artery.
leave the left pericardiacophrenic artery or they The ventral intercostal artery lying cranial to the
may come from the internal thoracic arteries di rib anastomoses with the collateral branch of the
rectly (Berry, Brailsford, and Daly 1931). They dorsal intercostal. These vessels lie, for the most
go to the roots of the lungs and furnish the minor part, in the endothoracic fascia closely under the
blood supply to the bronchi, bronchial lymph pleura. Occasionally some fibers from the inter
nodes, and connective tissue. They are fre nal intercostal muscles cover them. They supply
quently absent. the ventral part of the costal pleura, the adjacent
The mediastinal branches (rami mediastinales) intercostal musculature, and the costal cartilages.
supply the ventral part of the mediastinum. Usu The musculophrenic artery (a. musculophren-
ally two to four branches run directly into the ica) (Fig. 4-38) is the smaller, lateral, terminal
322 Chapter 4. The H e a rt and A r te r ie s
S p in a l far, In te rco s ta l o
T r a n s v e r s e c o l 11 a. t- iD o r s a l br.
S uprem e in te r c o s t a l a L. b ro n c h o e s o p h a g e V F b r.
D eep c e r v i c a l o v
A o rta
C a s to c e rv ic a l tru n k
V e r t e b r a l a. Esophagus
iPostcava
L .c o m m o n c a r o t i d v
L . c a u d a I th y ro id a-*
F i r s t in te rc o s ta l a -
P h re n ic
O m o c e rv ic a l
A x i I l a r u a. '
X
L. s u b c l a v i a n a '
Ext. t h o r a c i c a.
\ . 1
P e r i c a r d ia c o p h r e rue a.1
B ra c h io c e p h a
Thym
L . m t t h o r a c i c o-
P u lm o n a ry a .1 f
P e r f o r a t i n g br.
H e a rt
M e d ia s tin a l b r
P h re n ic a *
D ia p h ra g m '
S p in a l br.t i i Transverse c o ll i a.
i i
i _______I Supreme in te rc o s ta l a.
Dorsol br.\ \ i i i
i ,Deep c e r v ic a l a.
I n t e r c o s t a l a.s I I /
zC ostocervical tr u n k
, V e r te b ra l a.
Esophageal br
\
A ly g o s Us
f i t . common carotid
A o r ta -
Rt. caud. th y ra id a
Esophagus
- - F ir s t intercostal a.
— Phrenic n.
" - R t s u b d a v io n a.
" ■'Omocervical a.
a.
ardiacophrenic a.
th o ra c ic a.
ranch to thymus
Brachiocephal ic a.
Thymus
P e rforating b r
1H ea rt
i i ' \ ’
i i y 'int. th o ra c ic a
1' 'i 'I . ' P o s tc a v a
I |
i i Phrenic a.
I i i
i i i D iap hrag m
i
1C ra n ia l e p i g a s t r i c a.
1Musculophrenic a.
Fic. 4-36. Arteries of the right thorax.
324 Chapter 4. The H e a r t an d A r te r ie s
branch of the internal thoracic. It arises under The deep cranial epigastric artery (a. epigas
the caudal part of the transverse thoracic muscle trica cranialis profunda) (Fig. 4-38) runs initially
opposite the eighth interchondral space close to on the deep surface of the rectus abdominis
the sternum. It runs caudodorsolaterally, in the where it lies about 1 cm. from and parallel to the
angle formed by the diaphragm and the lateral linea alba. At a transverse level through the um
thoracic wall, where it lies in a small amount of bilicus many of its branches enter the rectus ab
fat covered by the pleura. After it has traveled dominis and shortly thereafter anastomose with
about one-fourth of the length of the costal arch the cranially running terminal branches of the
it perforates the diaphragm and comes to lie caudal deep abdominal artery. It is the primary
under the peritoneum. It ascends on the inner blood supply to the middle portion of the rectus
surface of the costal arch by following the mar abdominis. It is accompanied by its laterally
gins of the interlocked digitations of attachments lying satellite vein.
of the diaphragm and the transverse abdominal The ventral abdominal wall thus has two
muscle. Along its course it sends the ventral arterial channels on each side of the median
intercostal arteries (aa. intercostales ventrales) plane which connect the thoracic circulation
dorsally in the caudal part of the eighth inter with that of the pelvic limbs. One of these is
chondral space, and two each for spaces 9 and superficial and the other is deep. The deep
10. The single terminal branch of the musculo epigastric vessels are always well developed, but
phrenic anastomoses with the eleventh dorsal the superficial ones reach their maximum size
intercostal artery caudal to the diaphragm. These only during the height of lactation. The deep
ventral intercostal arteries form feeble anasto vessels anastomose feebly with each other across
moses with the ventral parts of the eighth, ninth, the linea alba.
and tenth dorsal intercostal arteries. Numerous
small branches leave the musculophrenic to sup
ply the muscular periphery of the diaphragm. A r t e r ie s o f t h e T h o r a c ic L im b
to the large caudodorsally running subscapular nutrient artery into the humerus, continues ob
artery. It runs caudally in the axillary fat and liquely distocranially on the brachial muscle and
crosses the lateral surface of the axillary lymph anastomoses with the proximal collateral radial
node, which it supplies. It also supplies an area artery above the flexor angle of the elbow joint.
of the latissimus dorsi ventral to the area sup During its course it supplies branches to the bra
plied by the thoracodorsal artery. Other chialis and heads of the triceps which lie along
branches supply the deep pectoral and cutane its course. This vessel is accompanied by its
ous trunci muscles. Its lateral mammary small satellite vein and the radial nerve.
branches (rami mammarii laterales) supply the The main part of the caudal circumflex hu
dorsolateral portions of the cranial and caudal meral artery arborizes extensively in the triceps
thoracic mammary glands when these are fully as ascending and descending branches. The cau
developed. The lateral thoracic artery is accom dal part of the shoulder joint capsule, infraspina
panied by a satellite vein and nerve. tus, teres minor, and coracobrachialis also
The subscapular artery (a. subscapularis) receive twigs from this vessel. Some of the prox
(Figs. 4-40, 4-41, 4-42) may be larger than the imal branches anastomose with the small cir
continuation of the axillary in the true arm. This cumflex scapular artery, and some of the de
great size is explained by the fact that the vessel scending branches anastomose with the deep
supplies a greater muscular mass in the shoulder brachial artery. The anastomosis between the
and arm than is present in the remainder of the posterior and anterior circumflex humeral arter
limb. It arises from the caudal surface of the ax ies in man is classical; in dogs, there is only a
illary usually just peripheral to the origin of the small union between the homologous arteries.
lateral thoracic artery. It runs obliquely in a dor- The main stem of the caudal circumflex leaves
socaudal direction between the subscapularis the triceps mass and enters the deep face of the
and teres major, and becomes subcutaneous near deltoideus. Other branches extend between the
the caudal angle of the scapula. It has the follow deltoideus laterally and the long and lateral
ing branches: thoracodorsal, caudal circumflex heads of the triceps medially, to appear subcu-
humeral, circumflex scapular, muscular, and cu taneously near the middle of the lateral surface
taneous. of the brachium. Some of these branches extend
The thoracodorsal artery (a. thoracodorsalis) is proximally to anastomose with the descending
a large artery which leaves the caudal surface of branch of the omocervical; others run distally
the subscapular less than 1 cm. from its origin. It and anastomose with the proximal collateral ra
runs caudally, usually supplying a part of the dial artery. The caudal circumflex humeral ar
teres major as it crosses the medial surface of tery is accompanied through the fleshy part of
the distal end of the muscle, and terminates in the brachium by its satellite vein and the axillary
the latissimus dorsi and skin. A satellite vein and nerve.
nerve accompany the artery. The circumflex scapular artery (a. circumflexa
The caudal circumflex hum eral artery (a. cir- scapulae) is a small vessel which leaves the cra
cumflexa humeri caudalis) leaves the lateral sur nial surface of the subscapular artery and, ex
face of the subscapular artery at about the same tending obliquely dorsocranially between the
level as the thoracodorsal and plunges immedi subscapularis medially and the long head of
ately between the head of the humerus and the the triceps laterally, reaches the caudal border
teres major. It is the principal source of blood to of the scapula near its middle. Here it divides
all four heads of the large triceps muscle. The into a medial and a lateral branch. The medial
collateral radial artery o f the humerus leaves the branch ramifies in the periosteal part of the sub
distal surface of the caudal circumflex humeral scapularis, and the lateral branch arborizes in a
about 1 cm. from its origin and takes a direct similar manner in the infraspinatus. Minute twigs
course distally, lateral to the terminal ends of enter the bone from both medial and lateral
the teres major and latissimus dorsi, and the me parts.
dial head of the triceps. It lies medial to the ac Distal to the origin of the circumflex scapular
cessory head and caudal to the brachial muscle. artery there are muscular branches (rami muscu-
It may terminate, as the nutrient artery o f the lares) to the proximal ends of the teres major,
humerus, by entering the nutrient foramen near subscapularis, infraspinatus, deltoideus, and
the middle of the caudal surface of the bone. In latissimus dorsi. A large patch of skin covering
most specimens, as Miller (1952) has pointed the region over and caudal to the caudal angle of
out, the collateral radial artery, after sending the the scapula is supplied by the terminal cutane-
326 Chapter 4. T he H e a rt and A rte rie s
b r a e h i o c e p h a l i cus
,' M . s t e r n o c e p h a l i c u s
P r e s c a p u l a r Inn.
Y A s c e n d in g
c e r v i c a l b ra n c h e s
C o m m o n c a r o t i d a.
M tra p e z iu s '
S ^ u j j - - M u s c u l a r ram u s
S u p e r f c e r v i c a l br: '
---- /ji - -Omocervical v.
M. s u p r a s p i n a tu S ' - E x t j u g u l a r v.
M o m o tra n s v e r s a r iu s '
~ - O m o c e r v i c a l a.
S u p r o s p m o u s br.
^ M p e c t o r a l i s prof.
S u pro scap ul a r b r ' " D e s c e n d 1n g br.
M. d e l t c i d e u s ' ' " ^ M. p e c t o r a l i s s u p e r f
P r o x i m a l b r,'
M b r a c h i o c e p h a l 1c u s
D is ta l br-~
Pectoral limb —
L common c a r o t id - M anubrium
O m o c e r v ic a l
, - R common c a r o t i d
A x illa ry -
V e rte b ra l„ -Ft. s u b c l a v i a n
C o s to c e rv ic a l tru n k - _
F i r s t i n t e r c o s t a l ----- - - B ra c h io c e p h a lic
L. s u b c l a v i a n - -
-R. int. t h o r o c i c a. r v.
L int. t h o r a c i c a.vv.-
D e s c e n d in g a o r t a -
-----C o l l a t e r a l br.
M e d i a s t i n a l c a - '- ' - - D o r s a l in t e r c o s t a l
S te rn u m - - - - M . i n t e r c o s t a l i s int.
M. trans. thoracis -
- - V e ntra l
i n t e r c o s t a l aa
M u scu lo p h re n ic
- C r a n i a l superf.
D ia p h ra g m -- epi g a s t r i c
- C r a n ia l deep
epi g a s t r i c
M tra n sversus - -
a b d o m i n is
- M re c tu s
a b d o m in is
Fic. 4-38. The internal thoracic arteries, dorsal aspect, in relation to sternum.
328 Chapter 4. The H e a r t an d A r te r ie s
oils branches (rami cutanei). The subscapular ar branch enters the medial head of the triceps,
tery and its branches are accompanied by satel and a larger one, after a course of over 1 cm.,
lite veins. is distributed to the long head. Within the tri
The cranial circumflex humeral artery (a. cir- ceps the relatively small deep brachial anasto
cumflexa humeri cranialis) usually arises from moses with the caudal circumflex humeral ar
the medial surface of the axillary proximal to the tery proximally and with the collateral ulnar
origin of the subscapular. It may arise from the distally. It is accompanied by a satellite vein.
axillary, distal to the origin of the subscapular, or The radial nerve enters the triceps cranial to
from the subscapular artery itself. It is a small the artery.
vessel which curves around the neck of the hu The bicipital artery (a. bicipitalis) is fre
merus under the tendon of origin of the biceps quently called the muscular ramus to the biceps.
brachii after crossing the insertion of the cora- It may be double and usually, when it is double,
cobrachialis. A relatively large branch supplies one branch arises from the proximal collateral
the proximal end of the biceps, and smaller twigs radial artery. The bicipital artery, when it is sin
go to the coracobrachialis and to the conjoined gle, usually arises from the medial surface of the
teres major and latissimus dorsi. A branch ex brachial at the junction of the middle and distal
tends proximally to supply the cranial part of the thirds of the arm. The bicipital artery anastomo
joint capsule. In the region of the major tubercle ses with the muscular branch to the biceps from
of the humerus the two circumflex humeral ves the cranial circumflex humeral artery. It is ac
sels join with each other as well as with the su companied by a satellite vein. It runs distally
prascapular above and with the descending and enters the distal end of the biceps brachii. It
branch of the omocervical below. Occasionally may arise from the proximal collateral radial ar
the supraspinous also joins in this anastomosis. tery.
The collateral ulnar artery (a. collaterals ul
Brachial Artery naris) (Figs. 4-40, 4-43) arises from the caudal
surface of the brachial in the distal third of the
The brachial artery (a. brachialis) (Figs. 4-40, arm. Usually the first branch runs proximocau-
4-41) is a continuation of the axillary. It begins dally to enter the medial surface of the triceps.
at the distal border of the conjoined tendons of This may be paired with one branch leaving the
the teres major and latissimus dorsi. It termi brachial directly. Another branch runs distally
nates in the forearm by bifurcating into the ra toward the palmar side of the forearm, with the
dial and ulnar arteries. (Some authors name the ulnar nerve below. Usually a strong branch
antebrachial part of the parent vessel the median arises from this vessel proximal to the elbow
artery.) The brachial artery in the brachium lies joint and, running under the medial head of the
caudal to the musculocutaneous nerve and bi triceps and anconeus, plunges into the olecranon
ceps muscle, medial to the medial head of the fossa. It supplies primarily the fat and the pouch
triceps and humerus, and cranial to the median of the elbow joint capsule which are located
nerve and axillary vein. The deep pectoral mus here. The branch which continues distally arbor
cle and a nerve connecting the musculocutane izes in the proximal parts of the flexor muscles of
ous and median nerves form the medial bound the antebrachium. An anastomosis exists with a
ary of the brachial artery in the brachium. It proximally extending branch from the accessory
crosses the distal half of the humerus obliquely interosseous artery between the ulnar and hu
to reach the medial surface of the elbow joint, meral heads of the deep digital flexor. Slightly
then it passes under the pronator teres muscle caudal to the branch which runs with the ulnar
and, at the junction of the proximal and middle nerve is a superficial branch which runs distally
thirds of the forearm, it terminates in the small across the medial surface of the elbow joint and
radial and the large ulnar artery. The collateral continues in the subcutaneous tissue of the proxi
branches of the brachial artery as it lies in the mal half of the palmar surface of the antebra
arm are: the deep brachial, bicipital, collateral chium. It supplies the skin here and anastomoses
ulnar, proximal collateral radial, and the distal with a proximally extending subcutaneous twig
collateral radial. from the palmar interosseous artery. It is accom
The deep brachial artery (a. brachialis pro panied by a vein and the caudal (palmar) cutane
funda) leaves the caudal side of the brachial in ous antebrachial nerve.
the proximal third of the arm. Occasionally the The proximal collateral radial artery (a. col-
artery is double. The deep brachial enters the lateralis radialis proximalis) leaves the cranial
medial and long heads of the triceps; a smaller surface of the brachial about 3 cm. proximal to
T h o r a c ic L im b 329
the elbow joint and extends obliquely distocra- most of which supply this muscle. Occasionally
niad to its flexor surface. After it crosses the ten a branch runs proximally in company with the
don of the biceps it gives off a cutaneous branch radial nerve and anastomoses with that part of
to the skin of the medial surface of the antebra the nutrient artery of the humerus which de
chium. In the region of the cephalic vein, under scends beyond the foramen. Besides the
which it runs, it gives off the medial and lateral branches which go to the extensor carpi radialis
branches. there are twigs which go to the supinator, com
The m edial branch (ramus medialis) extends mon digital extensor, and brachialis. Anastomo
from the flexor surface of the elbow joint to the ses with the dorsal interosseous and proximal
medial part of the forepaw. In its course down collateral radial sometimes occur.
the antebrachium it lies on the extensor carpi ra The median artery (antebrachial part of the
dialis, where it is bounded laterally by the large brachial artery) (Figs. 4-43, 4-44) extends from
antebrachial part of the cephalic vein and medi the flexor angle of the elbow joint to its terminal
ally by the small medial branch of the superficial radial and ulnar arteries which are located un
radial nerve. At the carpus it anastomoses with der the radial origin of the flexor carpi radialis in
the dorsal branch of the radial artery before con the middle third of the antebrachium. It is ac
tinuing to the dorsomedial part of the metacar companied by the median nerve which lies cau-
pus. The resultant branches of this anastomosis domedial to it and a relatively small satellite
run on the dorsal carpal ligament and there form vein. At the elbow joint several small branches
the medial part of the poorly defined dorsal rete leave the brachial artery. A few twigs go to the
o f the carpus (rete carpi dorsale). The lateral adjacent part of the elbow joint capsule. Other
part of this rete is formed by the distal dorsal in larger branches are distributed primarily to the
terosseous artery. The deep dorsal metacarpal flexor carpi radialis and pronator teres. The re
arteries, which the dorsal rete of the carpus con current ulnar, common interosseous, and palmar
tributes to the forepaw, are described under the antebrachial arteries are the branches of the an
heading: “Arteries of the Forepaw.” tebrachial part of the brachial which have re
The lateral branch (ramus lateralis) of the ceived definite names.
proximal collateral radial artery is the main con The recurrent ulnar artery (a. recurrens ul
tinuation of this vessel down the front of the naris) is a small vessel which leaves the caudal
forearm. It runs transversely, laterally across the side of the median at the proximal end of the
distal end of the biceps brachii and, upon emerg radius and extends from the caudal border of
ing from under the cephalic vein, bends distally the pronator teres into the flexor group of mus
and accompanies this vein on its lateral side cles. The artery first runs under the flexor carpi
throughout the antebrachium. A small ascend radialis near its origin and contributes to its sup
ing branch, which arises as it turns distally, anas ply. It continues caudally through the humeral
tomoses with the descending branch of the omo head of the deep digital flexor, which it supplies,
cervical artery. The lateral branch is somewhat and terminates mainly in the superficial digital
larger than the medial branch, but, like it, it is flexor. Two traceable anastomoses are present;
small, long, and sparsely branched. Throughout one of these is with the collateral ulnar artery, as
its antebrachial course it is flanked medially by the recurrent ulnar runs proximally over the me
the large cephalic vein of the antebrachium and dial epicondyle of the humerus, and the other is
laterally by the lateral branch of the superficial with the palmar antebrachial on the deep sur
radial nerve. On the proximal part of the meta face of the superficial digital flexor. Davis (1941)
carpus the artery trifurcates. The three resultant describes two recurrent ulnar arteries in the dog.
arteries are described under the heading: “Arter The common interosseous artery (a. interossea
ies of the Forepaw.” communis) (Fig. 4-45) is the largest branch of
The distal collateral radial artery (a. collater- the median artery. It is about 3 mm. in diam
alis radialis distalis) (Fig. 4-44) escapes casual eter and 1 cm. long, as it runs from the lateral
observation. It arises from the lateral surface of surface of the brachial to the interosseous space
the brachial, about 1 cm. before that vessel runs at the proximal end of the pronator quadratus.
under the pronator teres. The distal collateral This places the artery about 1cm. distal to the
radial, which is about as large as the proximal elbow joint. Before entering the interosseous
vessel, runs under the distal end of the biceps space it sends one or more muscular twigs cra
brachii and brachialis. On reaching the extensor nially into the pronator teres and gives off the
carpi radialis it breaks up into many branches, large caudally running accessory interosseous
(Text continued on page 333.)
330 Chapter 4. The H eart an d A r te r ie s
iEsophagus
L . s u b c la v ia n a.x
I
\ ^ L. bronchoesophageaI a.
B ra c h io c e p h a lic t r u n k s x
Branches to thym us - _
P h r e n ic n.—
P e r ic a r d i a c o p h r e n ic a
B r a n c h to thymus '
L . i n t t h o r a c i c a.
Thymus
H e a r t'’
F it. 4-39. Arterial supply of the thymus gland in a young dog, left lateral aspect.
T h o r a c ic L im b 331
M s u p ra s p in a tu s
- M. s u b s c a p u la r is
S u p ra s c a p u la r a .- -
-M. te re s m a j o r
-M. l a t is s i m u s d a r s i
Ext. t h o r a c i c a -
- S u b s c a p u la r a
A x i l l a r y a.-
- T h o ra c o d o r s a l a
Lat. t h o r a c i c a - -Caud. c i r c u m f l e x hum eral o.
-Accessory a x il l a r y lymph node
To m .p e c to r a lis p r o f -
- A x i l l a r y lymph node
Cron circu m fl. h u m e ra l a -
M. p e c t o r a l i s p r o f . - - - M u s c u la r b r
Deep b r a c h i a l o.- -
M b ic e p s b r a c h i i -----
- ~M. t r i c e p s
B r o c h i a l a : ------
- -M. tensor fa s c ia e a n te b ra c h ii
M. p e c t o r a l is su pe rf. -------
B i d p i ta l a - - - - C o ll a t e r a l u l n a r a.
----- Dist. c o l l a t e r a l r a d i a l a
P r o x .c o lla t e r a I r a d i a l a., m e d i a l b r —
M. pronator te re s
Common in te ro s s e o u s a- -
M p ro n a to r t e r e s - -
------ Palmar a n teb ra c h ia l a.
artery. Within the interosseous space it termi of the lateral part of the dorsal carpal rete. A re
nates by dividing into the palmar interosseous current branch extends proximally and anasto
and proximal dorsal interosseous arteries. moses with the accessory interosseous artery on
The accessory interosseous artery (a. interos- the deep digital flexor.
sea accessoria) (ulnar of Davis 1941) can be ex The proximal dorsal interosseous artery (a. in
posed by separating the humeral from the ulnar terossea proximalis dorsalis) continues in the di
head of the deep digital flexor. The large ulnar rection of the common interosseous after the
nerve, which lies closely applied to the lateral palmar interosseous arises. It is a small vessel
border of the deep digital flexor, largely covers which emerges from the interosseous space,
the artery. On leaving the common interosseous about 2 cm. distal to the lateral epicondyle of
artery the accessory interosseous courses ob the humerus. It enters the deep surface of the
liquely distocaudally across the medial surface proximal extremities of the extensor carpi ulnaris
of the ulna and enters the deep digital flexor. A and the lateral and common digital extensors.
recurrent branch extends proximally between Small twigs also go to the pronator quadratus,
the radial and ulnar heads of the deep digital supinator, and the flexor muscle adjacent to the
flexor and anastomoses with the collateral ulnar proximal third of the ulna. The caudolateral part
artery. The bulk of the artery continues distally, of the elbow joint receives twigs from under the
however, in association with the humeral head caudal border of the extensor carpi ulnaris.
of the deep digital flexor but closely applied to Anastomoses exist between the proximal dorsal
the deep surface of the flexor carpi ulnaris. Near interosseous artery and the collateral ulnar and
the carpus the accessory interosseous and the palmar interosseous arteries.
palmar antebrachial anastomose. The small The palm ar antebrachial artery (a. antebra-
trunk vessel which results passes distally into the chialis palmaris) (Fig. 4-43) arises from the pal
carpal canal, where it usually anastomoses with mar surface of the brachial about 1 cm. distal to
the palmar interosseous artery. The accessory in the origin of the common interosseous. It is a
terosseous supplies largely the ulnar and hu branched vessel, about 1 mm. in diameter,
meral heads of the deep digital flexor and the which runs distocaudally under the flexor carpi
corresponding heads of the flexor carpi ulnaris. radialis into the deep digital flexor. It supplies
The palm ar interosseous artery (a. interossea the flexor carpi radialis, superficial and deep
palmaris) lies between the apposed surfaces of digital flexors, and the flexor carpi ulnaris. It
the radius and ulna. The pronator quadratus anastomoses prominently with the recurrent ul
muscle lies on the palmarolateral side of the ar nar under the superficial digital flexor, and with
tery. In its course down the forearm the artery the accessory interosseous under the humeral
supplies many small branches to adjacent struc head of the flexor carpi ulnaris in the distal
tures. The pronator quadratus and the ulnar and fourth of the antebrachium. Frequently the
radial heads of the deep digital flexor receive small common trunk formed by this anastomosis
twigs on the caudal side of the forearm. The ab joins the palmar antebrachial in the carpal canal.
ductor pollicis longus, common digital extensor, It traverses the antebrachium in such a way that
lateral digital extensor, and extensor pollicis a series of branches leave the vessel, as it lies in
longus et indicis proprius receive branches on the deep digital flexor, and terminate in the su
the dorsolateral side of the forearm. In the prox perficial digital flexor. These appear in a linear
imal half of the forearm it forms a feeble anasto series of about eight vessels lying 1 to 2 cm.
mosis with the proximal dorsal interosseous ar apart. The palmar antebrachial artery is accom
tery. At the junction of the proximal and middle panied by a branch of the median nerve and a
thirds of the radius the nutrient artery o f the ra satellite vein.
dius (a. nutriciae radii) extends distally into the The ulnar artery (a. ulnaris) (Fig. 4-43) is the
bone. At a similar location on the ulna the nutri principal source of blood supply to the forepaw.
ent artery o f the ulna (a. nutriciae ulnae) extends It arises as the larger terminal branch of the me
proximally into the ulna. At the base of the sty dian artery near the beginning of the middle
loid process of the ulna the palmar interosseous third of the antebrachium. It lies on the medial
artery bifurcates. One of these arteries, the dis borders of the radial and humeral heads of the
tal dorsal interosseous artery (a. interossea dor deep digital flexor under the heavy antebrachial
salis distalis) leaves the dorsal side of the inter fascia and tendon of the flexor carpi radialis. Its
osseous space proximal to the carpus. From distal antebrachial part obliquely crosses the hu
under the abductor pollicis longus it runs to the meral head of the deep digital flexor, which it
dorsal carpal ligament and aids in the formation grooves. Usually a small twig to the medial sur-
(Text continued on page 337.)
334 Chapter 4. The H e a rt and A r te r ie s
- M. trap eziu s
M te re s m a j o r - -
M. d e lto id e u s
M. s u b s c a p u la ris Ci r c u m fle x s c a p u la r a.
S u b s c a p u la r a
M. i n fro s p m a tu s
A x illa ry a
M. t r i c e p s c a p u t lonqum
Lat. t h o r a c i c a.
T h o r a c a d o r s a I a.
M. omotransversarius
M. tr ic e p s , c a p u t access
M. +eres m a j o r -
Deep b r a c h i a l a.
IV tric e p s , c a pu t medr - M. t r i c e p s , c a p u t l a t
B r a c h i a l a. - - C o lla te ra l r a d ia l a.
N u t r i e n t a. o f humerus -
- -M. b r a c h i a l is
M.biceps b r a c h i i
M. t r i c e p s -
M anconeus
M u sculocutaneous n.- - - -B ra c h ia l a
- -M t r i c e p s , c a p u t med.
M.biceps b r a c h i i -----
- -M. t e n s o r f a s c a n te b ra c h n
B i c i p i t a l o --------
- -U l n a r n.
M f le x d i g i t o r u m prof., ca p u t r a d i c l e
Accessory interosseous Q
P a lm a r a n te b r a c h ia l a.
M p ro n a to r q u a d ra tu s-
M . f l e x d ig it o r u m superf.
UI no -
U l n a r n,
R a d i a I a.-----
M. fle x, d i g i t o r u m p r o f .7 -
c a p u t h um e ra le
U l n a r a.
R a d i u s ------
--------Trans, p a l m a r c a rp a l hg.
face of the carpus is its only branch in the an collateral radial artery, and by a direct continua
tebrachium. As it passes through the carpal tion of the medial branch of the proximal collat
canal with the tendon of the deep digital flexor eral radial artery into the metacarpus from the
it lies lateral to the median nerve. It emerges carpus. These small vessels first lie on, then be
from the carpal canal in the palmar groove of tween, the tendons of the common digital exten
the deep flexor tendon. A small branch to the sor as both diverge unequally. The distal
carpal pad arises from the ulnar just distal to the portions of the main arteries sink into the distal
carpal canal. Lying between the superficial and portions of the intermetacarpal spaces and join
deep flexor tendons in the proximal part of the the deep dorsal metacarpal arteries. The first
metacarpus, the vessel anastomoses with a small artery anastomoses with the corresponding su
branch of the palmar interosseous to form the perficial palmar metacarpal artery.
superficial palm ar arch (arcus palmaris superfi The deep dorsal metacarpal arteries II, III,
cialis). This arch is not apparent without close and IV (aa. metacarpeae dorsales profundae)
inspection, as the ulnar artery dominates in the arise from the distal part of the dorsal rete of the
formation of the arch as well as in the supply of carpus. They are the smallest of all the metacar
blood to the paw. Essentially the ulnar artery pal arteries as they lie in the dorsal grooves be
terminates as three principal palmar metacarpal tween adjacent metacarpal bones, Proximally
arteries. they send branches to the deep palmar arch, and
The radial artery (a. radialis) (Fig. 4-43), from distally communicating branches are sent to the
its origin just proximal to the middle of the fore deep palmar metacarpal arteries. They terminate
arm, runs distally under the aponeurotic origin by anastomosing with corresponding arteries of
of the flexor carpi radialis. It closely follows the the superficial series to form the common dorsal
palmaromedial border of the radius in the fore digital arteries.
arm. At the carpus it divides into palmar and The dorsal common digital arteries II, III,
dorsal branches. The dorsal branch (ramus car- and IV (aa. digitales dorsales communes) are
peus dorsalis) supplies the dorsal part of the car each about 1 cm. long as they terminate oppo
pal joint capsule. It contributes to the formation site the metacarpophalangeal joints. Anasto
of the dorsal rete of the carpus by sending a motic twigs leave them to go to the correspond
branch to anastomose with the medial branch of ing palmar common digital arteries. The dorsal
the proximal collateral radial artery. The palmar common digital arteries, upon reaching the skin
branch (ramus carpeus palmaris) runs toward which ensheathes the digits, divide into the lat
the first digit by passing in the superficial part of eral and m edial dorsal proper digital arteries II,
the transverse carpal ligament. It is a small vessel III, and IV (aa. digitales propriae dorsales later-
which anastomoses with the larger palmar inter ales et mediales II, III, et IV). These go to the
osseous artery in the interosseous musculature of dorsal parts of the contiguous sides of adjacent
the proximal part of the metacarpus. The deep digits and, as branched cutaneous vessels, extend
palmar arch (arcus palmaris profundus) is to the claws. The superficial and deep dorsal
formed by this anastomosis. metacarpal arteries, common dorsal digital ar
teries, and the proper dorsal digital arteries are
accompanied by satellite veins and the dorsal se
ries by companion nerves. In the digits these
Arteries of the Forepaw structures are not closely related.
The palmar set of arteries of the forepaw, like
The arteries of the forepaw may be divided the dorsal set, is divided into a superficial and a
into a dorsal and a palmar set, each of which is deep series of arteries, but, unlike the dorsal set,
further divided into a superficial and a deep se these vessels arise from the superficial and deep
ries. In the metacarpus they are known as meta palmar arches. The ulnar artery dominates in the
carpal arteries; in the digits they are called the formation of the superficial series so completely
digital arteries. All are small and of minor im that the contribution of the palmar interosseous
portance, except for the superficial series of the artery, which anastomoses with it to form the
palmar set, which are the main source of blood superficial palm ar arterial arch (arcus palmaris
supply to the digits and the footpads. superficialis), is minor. From this arch arise the
The superficial dorsal metacarpal arteries I, relatively large superficial palmar metacarpal
II, III, and IV (aa. metacarpeae dorsales superfi- arteries.
ciales) (Figs. 4-46, 4-48) are formed by the tri The superficial palmar metacarpal artery I (a.
furcation of the lateral branch of the proximal metacarpea palmaris superficialis I) (Figs. 4-47,
338 Chapter 4. The H e a rt and A r te r ie s
F ic . 4-45. Arteries of the right antebrachium, <audolateral aspect .T h e shaft of the uliia is removed.)
T h o r a c ic A orta 339
4-48) runs about 1 cm. before entering the space digits. In fact, those that go to the palmar sur
between the first digit and the metacarpus. It is faces of the opposite, or abaxial, sides are mainly
the first branch from the metacarpal part of the small cutaneous branches. However, in the pal
ulnar artery or the superficial palmar arterial mar vascular canals of the third phalanges the
arch. It anastomoses with the superficial dorsal medial and lateral proper digital arteries anasto
metacarpal artery I to form the palmar common mose. Branches given off from these terminal
digital artery I (a. digitalis communis palmaris I). arches nourish the corium of the claws and the
The main part of the ulnar artery or the medial third phalanges.
limb of the superficial palmar arterial arch gives
rise to the superficial palmar metacarpal arter THORACIC AORTA
ies II, III, and IV (aa. metacarpeae palmares
superficiales II, III, et IV). These run distally The thoracic aorta (aorta thoracica) (Figs. 4 -
under the superficial flexor tendons where, proxi 49, 4-50) continues from the aortic arch, oppo
mal to the large metacarpal footpad, each sends site the fourth thoracic vertebra, and extends to
a branch into the proximal part of the pad. At the caudal border of the second lumbar vertebra.
the distal ends of the intermetacarpal spaces the There it enters the abdominal cavity by passing
arteries anastomose with the comparable deep between the obliquely placed crura of the dia
palmar metacarpal arteries to form the palmar phragm, to become the abdominal aorta. It is ac
common digital arteries. The superficial palmar companied by the azygos vein, thoracic duct,
metacarpal arteries are accompanied by small and the cisterna chyli as it passes through the
sensory nerve branches from the median nerve. aortic hiatus of the diaphragm. At its beginning,
The satellite veins accompany the arteries only the bulk of the thoracic aorta lies to the left of
in the distal half of the metacarpus. the median plane, being displaced to this posi
The deep palmar metacarpal arteries II, III, tion by the horizontally running esophagus,
and IV (aa. metacarpeae palmares profundae) which crosses it on the right. It lies in the medi
arise from the deep palm ar arch (arcus palmaris astinal septum and inclines slightly to the right
profundus), which is formed by the palmar and dorsally as it runs to the diaphragm. Cau
branch of the radial artery anastomosing with dally it is separated from the bodies of the
the terminal part of the palmar interosseous thoracic vertebrae by a small amount of fat. The
artery. This arch lies distal to the palmar carpal thoracic duct lies in this fat as it follows the dor
ligament, where it is covered by the interosseous sal surface of the aorta forward to the mid-
muscles. The deep palmar metacarpal arteries thoracic region.
run distally between these muscles, which they The branches of the thoracic aorta may be di
supply, and anastomose with the corresponding vided into visceral and parietal. The visceral
superficial palmar metacarpal arteries. These branches are the bronchial and esophageal. The
unions take place more than 1 cm. proximal to parietal branches include the intercostal, last
the metacarpophalangeal joints. The deep vessels thoracic, and the first two lumbar arteries.
send the several nutrient arteries into the proxi
mal third of the palmar surfaces of the four main
metacarpal bones. They are accompanied by Visceral Branches
satellite veins and companion nerves.
The palmar common digital arteries II, III, The bronchial and intrathoracic esophageal
and IV (aa. digitales palmares communes) are branches vary in number and origin. The chief
formed by the anastomoses of the superficial and nutritional blood supply to the lungs and related
deep palmar metacarpal arteries. Each of these structures at the hila of the lungs are the right
three pairs of arteries sends a prominent branch and left bronchial branches (rami bronchiales) of
into the center of each of the three parts into the bronchoesophageal artery (a. broncho-
which the distal portion of the large metacarpal esophagica). This vessel also sends ascending and
footpad is divided. After running about 1 cm. descending esophageal branches (rami eso-
the palmar common digital arteries divide into phagei) to most of the intrathoracic portion of
the lateral and m edial palmar proper digital the esophagus. The bronchoesophageal artery
arteries (aa. digitales propriae palmares laterales usually arises from the right fifth intercostal
et mediales), which supply the adjacent palmar artery close to the aorta. Variations described by
sides of contiguous digits. The proper vessels Berry et al. (1931) include: a branch from the
lying on the digits which are closest to the axis right fifth or sixth intercostal artery to the right
through the paw are the chief arteries to the lung, direct branches from the first part of the
340 Chapter 4. The H e a rt and A r te r ie s
- -Prox. c o /la te ra l D is t d o r s a l -
r a d ia l a , med. b r in te r o s s e o u s o
-R a d ia l a.,
d o r s a l br.
Prox. c o l la t e r a l
ra d ia l a, la t b r
- -D o rs a l c a r p a l re te
D eep d o r s a l<■
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m e ta c a rp a l a a
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-P a lm a r com m on d ig ita l a. I l l
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L o t d o rs a l
p r o p e r d ig i ta l a IV Lot. d o r s a l -
- Lot. p a lm a r p r o p e r d ig i t a l a.IV
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- P a lm a r - -P a lm a r
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p a lm a r a r c h - -
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descending aorta, and a single twig from the left arise close together, and occasionally the first
sixth intercostal artery to the left lung. The eso pair come from a common trunk. In the mid-
phageal branches come primarily from the bron thoracic region the paired vessels may be sepa
choesophageal artery in the form of ascend rated by as much as 4 mm. at their origins, and
ing and descending vessels. The descending this spacing continues throughout the remainder
branches anastomose with several long eso of the thorax. Right and left vessels are not sym
phageal branches that arise from two or more of metrical in their origins; the right vessels usually
the right intercostal arteries caudal to the origin arise caudal to the left. From the mid-thoracic
of the bronchoesophageal. These anastomose region caudally the dorsal intercostal arteries run
with each other and the last esophageal branch directly laterally across the bodies of the verte
anastomoses with the esophageal branch of the brae. Each dorsal intercostal artery gives off a
left gastric artery, which ascends through the dorsal branch.
diaphragm on the esophagus. The ascending The dorsal branches (rami dorsales) arise from
branches anastomose with the esophageal the intercostal arteries lateral to the bodies of
branches from the first aortic intercostal artery the vertebrae. They pass directly dorsally in the
of the left or right sides. This in turn anasto medial portion of the m. longissimus, obliquely
moses with the esophageal branch from the cau across the spines of the vertebrae, and end in the
dal thyroid. There is a small but definite arterial epaxial muscles or the skin as the small dorsal
passage from the caudal thyroid to the left gastric cutaneous branches (rami cutanei dorsales).
artery in or on the wall of the esophagus. Each has a spinal branch (ramus spinalis), which
Most bronchial and esophageal arteries have crosses the dorsal surface of the spinal nerve to
satellite veins. Usually the bronchoesophageal reach its caudal border. It follows this border
vein empties into the azygos at the level of the centrally to the nerve roots, after which it is dis
seventh thoracic vertebra. According to Berry tributed like the spinal branches of the vertebral
et al. (1931) only a capillary anastomosis exists artery. After the dorsal branches arise, the inter
between the bronchial and pulmonary circula costal arteries extend laterally dorsal to the
tions. pleura, sympathetic trunks, azygos vein (on the
Three or four small arteries leave the dorsal right side) and hemiazygos vein (at the last two
part of the thoracic aorta and run ventrally over or three intercostal spaces on the left side). The
its sides. Two or more of these are distributed to intercostal arteries supply segmental branches to
the dorsal part of the mediastinal septum as the overlying epaxial musculature. Some of these
m ediastinal branches (rami mediastinales). In branches become cutaneous along the lateral
the caudal part of the thorax one or more of border of the iliocostalis. They supply the dorsal
these vessels may arise from the ventral surface part of the latissimus dorsi and cutaneus trunci,
of the aorta. Other branches arise from the which they perforate to reach a paramedian
bronchoesophageal, aorta, or intercostal vessels band of skin. On their way to the skin they are
and run to the fibrous pericardium as the peri accompanied by the dorsal lateral cutaneous
cardial branches (rami pericardiaci). nerve of their segment. Sometimes a single
artery bifurcates so that each of its branches ac
Parietal Branches companies intercostal nerves which arise from
adjacent segments.
The intercostal arteries (aa. intercostales) Hughes and Dransfield (1959) have found that
(Figs. 4-51, 4-52) are 12 in number on each side. in the regions of the body where the skin is
The first three or four are branches of the su covered by hair, there are superficial, middle,
preme intercostal artery, and the last eight or and deep arterial and venous plexuses. The deep
nine are branches of the aorta. The artery which plexus may be arranged in more than one layer.
follows the caudal border of the last (thirteenth) The lateral cutaneous branches (rami cutanei
rib is the subcostal artery. All of the aortic inter laterales) form a second series of cutaneous
costal arteries arise from the dorsal surface of the branches (Marthen 1939). These start usually at
aorta and are similar in distribution. The fifth the third segment and perforate the external
right intercostal artery may serve as a common intercostal muscle and the thoracic part of the
trunk for the fourth, occasionally the third, and, serratus ventralis when it is present. The distal
in rare instances, for the second intercostal lateral cutaneous branches appear along the
artery, according to Reichert (1924). The first ventral border of the latissimus dorsi. They con
left aortic intercostal artery is rarely farther for tinue laterally through the cutaneus trunci and
ward than the fourth. Right and left arteries subcutaneous fascia to terminate in long ventral
T h o r a c ic Aorta 343
-Deep p a l m a r a r c h
U ln a r a
Deep d o r s a l m e t a c a r p a l a
D ee p p a l m ar m e t a c a r p a l a.
Superf. d o r s a l m e ta c a rp a l a - —
S u p e rf p a lm a r m e ta c o rp a l a II
S u p e r f p a l m a r m e t a c a rp a l a III
Dorsal common d i g i t a l a - -
P a lm a r common d i g i t a l a.
L a td o rs a l
p ro p e r d i g i t a l a III
L a t p a l m a r p r o p e r d i g i t a l a. Ill
Med. d o r s a l
proper d i g i t a l a IV
Med. p a l m a r p r o p e r d i g i t a l a IV
P h a la n x I V
F ig. 4-48. Arteries of the fourth digit of the right forepaw. medial aspect.
344 Chapter 4. The H e a rt and A r te r ie s
S e v e n th r i b >
' ✓, F i f t h i n t e r c o s t a l a.
and short dorsal branches. In the cranial part of lumbar vertebrae. They are distributed like other
the series the dorsal and ventral branches usu typical lumbar arteries. The first lumbar artery
ally arise independently from the intercostal anastomoses with the subcostal and the second
arteries. They are accompanied by satellite veins lumbar arteries and may effect a union with the
and the lateral cutaneous nerves. The nerves are cranial abdominal branch of the phrenico
constantly present, although the vessels may be abdominal.
missing in some of the segments. The compa
rable second nerve is the intercostobrachial
nerve, which goes largely to the skin of the ABDOMINAL AORTA
brachium.
The collateral branches (rami collaterals) The abdominal aorta (aorta abdominalis)
arise from the intercostal arteries near the begin (Figs. 4-49, 4-53) is that portion of the descend
ning of the ventral half of the thorax. They are ing aorta which lies in the abdomen. The dia
the last of four to eight oblique branches which phragm which separates the thoracic from the
run ventrocranially across the medial surfaces of abdominal cavity is perforated so obliquely by
each of the last 9 or 10 ribs to reach the inter the aorta that the first visceral branch of the ab
costal spaces cranial to them. They descend to dominal aorta is at a more cranial plane than the
anastomose with the ventral intercostal arteries origin of the second lumbar arteries. The abdom
which ascend in front of the costal cartilages. inal aorta terminates opposite the seventh lum
They supply the ventral halves of the intercostal bar vertebra by bifurcating into right and left in
muscles which lie immediately cranial to the ternal iliac and middle sacral arteries. Cranially,
ribs. Usually the last three dorsal intercostal it lies in the median plane between the crura of
arteries (intercostal arteries 10, 11, and 12) ter the diaphragm; caudally it is displaced slightly to
minate medial to the costal arch and ventro- the left by the postcava. It lies in the furrow
cranial to the intercostal spaces of the same formed by the right and left iliopsoas muscles.
number. After circling the costal margin of the The branches of the abdominal aorta supply
diaphragm each artery gives a branch to the dia both visceral and parietal structures.
phragm and another to the musculature of the
lateral abdominal wall. The phrenic components
of these vessels anastomose with the phrenic U n p a ir e d V is c e r a l B r a n c h e s of
the nutritional blood to the gland. When three right and left gastric arteries which lie on the
branches are present the first branch, the right lesser curvature. Long, freely branching epiploic
branch (ramus dexter), goes to the right portion branches (rami epiploici) leave the opposite or
of the liver, the caudate and right lateral lobes. omental side of both gastroepiploic arteries and
The m iddle branch (ramus medius) goes to the ramify mainly in the parietal leaf of the greater
right medial lobe, dorsal part of the quadrate, omentum. Gravenstein (1938) demonstrated
and part of the left medial lobe; this vessel may that there is a strong epiploic branch which
be replaced by two or more arteries. The left leaves the right gastroepiploic artery near its
branch (ramus sinister), shortest of the three, origin and runs caudally in the parietal leaf of
supplies the large left lateral lobe, the quadrate the greater omentum near its right border. A
lobe, and part of the left medial lobe. From this second vessel with a similar origin essentially
branch arises the cystic artery (a. cystica). This parallels the first, but runs in the visceral layer
vessel leaves the left branch, about 1 cm. before of the greater omentum near its right border.
it enters the liver, and ramifies by two or more The comparable vessels near the left border of
branches primarily on that surface of the gall the greater omentum are continuations of the
bladder which is attached to the liver. After giv splenic artery near the apex of the spleen. There
ing off the proper hepatic arteries, the common are many other smaller epiploic branches which
hepatic artery terminates as the small right gas arise at short intervals from the gastroepiploic
tric and the much larger gastroduodenal artery. arteries as they lie in the omentum just periph
The right gastric artery (a. gastrica dextra) eral to the greater curvature of the stomach.
leaves the common hepatic artery at nearly a These vessels anastomose freely with each other.
right angle and, running in the lesser omentum The fat of the omentum is deposited around the
at the pylorus, continues to the lesser curvature epiploic vessels. The right and left gastroepiploic
of the stomach. It frequently arises from one of arteries anastomose with each other opposite the
the proper hepatic arteries, in which case the beginning of the pyloric antrum. The long left
common hepatic artery becomes the gastroduo gastroepiploic artery on its way to the stomach
denal after the last proper hepatic arises. The supplies most of the epiploic vessels to the
right gastric artery sends branches to both the greater omentum.
parietal and the visceral surface of the pylorus, The cranial pancreaticoduodenal artery (a.
pyloric antrum, and the lesser omentum. It pancreaticoduodenalis cranialis) is the slightly
anastomoses with the much larger left gastric ar larger terminal branch of the gastroduodenal. It
tery close to the pylorus on the pyloric antrum continues the parent artery in the mesoduode-
as it runs toward the cardia in the lesser omen num and enters the pancreas at the junction of
tum. its right and left limbs. Shortly before entering
The gastroduodenal artery (a. gastroduode- the right limb of the pancreas it sends a small
nalis) runs across the first part of the left limb branch to the left limb which, after extending
of the pancreas to the medial surface of the duo a short distance, anastomoses with the pancre
denum, where it terminates. It may issue deli atic branch of the splenic, which is the chief
cate twigs to the pylorus, and constantly sends supply to this part of the gland. The principal
one or more larger branches into the left limb part of the cranial pancreaticoduodenal artery
of the pancreas. It terminates as the right gas continues caudally in the right limb of the pan
troepiploic and the cranial pancreaticoduodenal creas, sending pancreatic branches (rami pan-
artery. creatici) to the right limb of the pancreas and
The right gastroepiploic artery (a. gastroepi- du oden al branches (rami duodenales) through
ploica dextra) leaves the pancreas from the the gland to the duodenum. In the caudal half
medial surface of the duodenum and enters of the right limb of the pancreas it anastomoses
the greater omentum. It lies about 1 cm. from the with one or more pancreatic branches coming
greater curvature of the stomach as it runs from the caudal pancreaticoduodenal. The main
in the greater omentum toward the cardia. It part of the artery usually leaves the caudal third
sends branches to the stomach at intervals of of the right limb of the pancreas, traverses a part
about 5 mm. Most of these gastric branches (rami of the mesoduodenum, and comes to lie on the
gastrici) divide on reaching the greater curva mesenteric border of the descending duodenum.
ture of the stomach into a branch which goes to Near the caudal flexure it anastomoses with the
the parietal surface and one which goes to the main duodenal branch of the caudal pancreati
visceral surface. These anastomose in the mus coduodenal artery.
culature of the organ with gastric branches of the The splenic artery (a. lienalis) (Fig. 4 -5 8 ) is
A b d o m in a l A o r t a 347
/ M. t r a p e z i u s
Mm. s p i n a l i s ? s e m i s p i n a l i s
th o ra c is ? c e r v ic is
M. i n t e r s p i n a l i s - - /M . lo n g is s im u s d o r s i
Mm. r o t a t o r e s r m u l t i f i d u s d o rs i le v a to r c o s ta e
,M . i I i o c o s t a l i s d o r s i
D o r s a l br.
la tis s im u s d o rsi
S p in a l b r a n c h e s s e rra tu s d o rs a lis
Prox. l a t c u ta n e o u s br.
In te rc o s to l a r te r ie s
A o rta ■M. i n t e r c o s t a l i s ext.
-M , i n t e r c o s t a l i s in t.
--6 th rib
- -M . s e r r a t u s vent.
- D i s t a l lat. c u t a n e o u s br.
-P le u ra
o b l iq u u s a b d o m i n i s ext.
I n t e r c o s t a l br. nsversus th a r a c is
I nt. t h o r a c i c a.- b d o m in is
S te r n u m ~
prof.
Vent. c u ta n e o u s br.-
F ig . 4-51. Scheme of the arteries of the thoracic wall in cross section, caudal aspect.
348 Chapter 4. The H e a rt and A r te r ie s
1. Superficial cervical branch of omocervical 8. Distal lateral cutaneous branches of inter- 12. Caudal superficial epigastric
2. Cranial circumflex humeral costals 13. Medial genicular
3. Caudal circumflex humeral 9. Proximal lateral cutaneous branches of in- 14. Cutaneous branch of caudal femoral
4. Proximal collateral radial tercostals 15. Perineal
5. Lateral thoracic 10. Ventral cutaneous branches of internal 16. Deep circumflex iliac
6. Cutaneous branch of thoracodorsal thoracic 17. Tuber coxae
7. Cutaneous branch of subscapular 11. Cranial superficial epigastric 18. Cutaneous branches of superficial lateral
coccygeal
A b d o m in a l A o r t a 349
that branch of the celiac which runs to the left gastroepiploic artery. The vascular pattern in
and lies in a groove of the left limb of the pan the visceral leaf of the greater omentum is simi
creas near its free end. It is usually over 2 mm. lar to that in the parietal.
in diameter and gives off 3 to 5 long primary The left gastric artery (a. gastrica sinistra)
branches as it courses in the greater omentum arises from the cranial surface of the celiac as its
toward the apical or ventral third of the spleen. smallest terminal branch. It may be double.
The first branch usually is the pancreatic branch When single, it may form a common trunk with
(ramus pancreaticus), which leaves the right side the splenic. It runs cranially in the lesser omen
of the vessel at its origin and enters the apex of tum to the fundus of the stomach adjacent to the
the left limb of the pancreas. It is the main sup cardia. It sends long subserous branches to both
ply to this limb of the gland and anastomoses surfaces of the organ with the heavier branches
with the smaller pancreatic branch from the cra going to the parietal surface. An anastomosis be
nial pancreaticoduodenal artery. Usually a single tween right and left gastric arteries takes place
vessel to the pancreas is present at this site, but in the lesser omentum. In addition to supplying
occasionally this is augmented by as many as the fundus small epiploic branches go to the
three other small twigs. Arising from the cranial lesser omentum and one or more esophageal
surface of the splenic artery are two long vessels branches (rami esophagei) run through the
which run toward the proximal half of the esophageal hiatus to supply the caudal part of
spleen. They do not enter the gland directly, but the esophagus. These branches anastomose with
pass at right angles to the long hilus of the the esophageal rami from the thoracic aorta.
spleen, where they send splenic branches (rami The celiac trunk is surrounded by the heavy
lienales) to it. They then continue in the gastro- celiac plexus of nerves and the two celiac ganglia
splenic ligament to the greater curvature of the which are intimately connected with both the
stomach, where they supply the small gastric ar adrenal and cranial mesenteric nerve plexuses.
teries (aa. gastricae) to the fundus which anasto Lymph vessels from the stomach, spleen, pan
mose with the much larger gastric branches from creas, and a portion of the duodenum run dor
the left gastric artery. The main part of the sally in association with the celiac trunk to the
splenic artery approaches the hilus of the spleen cisterna chyli. The hepatic, splenic, and left gas
near its middle, where it sends many branched tric arteries and their branches have accompany
splenic twigs into the distal half of the gland. ing lymphatics and satellite veins which are rad
This main trunk, now called the left gastroepi icles of the portal vein.
ploic artery (a. gastroepiploica sinistra), contin The cranial mesenteric artery (a. mesenterica
ues in the gastrosplenic ligament to the greater cranialis) (Figs. 4-59, 4-60) is the largest visceral
curvature of the stomach. It runs near this cur branch of the aorta. This unpaired artery arises
vature toward the pylorus, where it anastomoses from the ventral surface of the abdominal aorta
with the small right gastroepiploic artery. Other about 5 mm. caudal to the origin of the celiac ar
sizeable branches leave it and run toward the tery opposite the first or second lumbar vertebra.
cardia, where the terminal vessel anastomoses It is about 5 mm. in diameter and is surrounded
with an adjacent, more proximal branch of the by the cranial mesenteric plexus of autonomic
splenic. The gastric arteries (aa. gastricae) in this nerves. It passes ventrocaudally through the dor
region run considerable distances in the sub- sal mesentery and acts as an axis around which
serosa before entering the muscularis. They the whole small and large intestine rotates dur
anastomose with the gastric branches of the left ing development. As it extends into the intestinal
gastric artery which come from the lesser cur mass it is loosely bounded by the duodenum on
vature, and finally the terminal part of the ves the right and caudally. This is followed distally
sel joins the small right gastroepiploic artery by the colon which hooks around the artery in
near the beginning of the pyloric antrum. All pri such a way that the ascending colon lies to its
mary divisions of the splenic, as they run right, the transverse colon in front, and the de
through the greater omentum and the gastro scending colon to the left. The cranial mesen
splenic ligament, give off om en tal (or epiploic) teric artery is most closely associated with its
branches (rami epiploici) to the omentum. These autonomic plexuses of nerves which wrap the
are disposed like the comparable branches from artery and its subdivisions. Peripheral to the
the right gastroepiploic. According to Graven- plexus are the long mesenteric lymph nodes and
stein (1938), these branches swirl to the right in the portal vein. The first two branches of the
the streaks of fat deposited around them and cranial mesenteric artery arise from opposite
anastomose with similar branches from the right sides of the artery, about 2 cm. from its origin.
350 Chapter 4. The H e a rt and A r te r ie s
One of these, the common colic, runs cranially colic junction. It supplies small twigs to the junc
in the transverse mesocolon, while the other, the tion, sends an ascending ramus proximally along
caudal pancreaticoduodenal, from a caudal ori the mesenteric border of the ileum to anasto
gin, runs to the right and cranially. The remain mose with the last intestinal (ileal) artery, and
ing part of the artery gradually diminishes in size sends a branch distally to anastomose on the
as some 14 jejunal arteries arise from its caudal right colon with the colic branch. Variations in
side. Two ileal arteries terminate the cranial the vascularity of this portion of the digestive
mesenteric. tube are extremely common.
The common colic artery (a. colica communis) The ileocecal artery (a. ileocecalis) runs across
is about 1.5 mm. in diameter and 2 cm. long. It the dorsal surface of the ileocolic junction. It
gives origin in succession to the middle colic, ac leaves the subserosa and disappears in the areo
cessory middle colic, and right colic, and con lar tissue uniting the cecum to the ileum. It
tinues as the ileocecocolic. sends ileal branches (rami ileales) to the ileum
The m iddle colic artery (a. colica media) and cecal branches (rami cecales) to the cecum.
arises from the first few millimeters of the left The main part of the vessel, after emerging from
side of the common colic or from the cranial between the ileum and cecum, continues prox
mesenteric directly. It runs cranially in the imally in the ileocecal fold as the ramus ileacus
transverse mesocolon, where it usually makes antimesenterialis. According to Thamm (1941),
one spiral turn. It bifurcates about 2 cm. from this vessel extends along the ileum and anasto
the left colic flexure. One branch runs distally moses with the last jejunal artery in the muscu
in the descending mesocolon and, after supply lature of the small intestine. The common colic
ing about half of the descending or left, colon, artery and its branches are accompanied by
anastomoses with the left colic artery, a branch plexuses of autonomic nerves which take the
of the caudal mesenteric. The other branch names of the vessels they follow, and all except
swings in the opposite direction and forms an the common colic have satellite veins and lym
arcade with the smaller accessory middle colic phatics.
artery. The caudal pancreaticoduodenal artery (a.
The accessory m iddle colic artery (a. colica pancreaticoduodenalis caudalis) (Fig. 4-55)
media accessoria) supplies the proximal portion arises from the cranial mesenteric opposite the
of the transverse colon. It forms terminal arcades origin of the common colic and runs to the right
with the middle and right colic arteries, from in the mesentery to the descending portion of
which vasa recti arise which supply all the trans the duodenum near the caudal flexure. On pass
verse colon and a part of the right colon also. ing the tip of the right limb of the pancreas it
This vessel may be absent or double. sends from one to three pancreatic branches
The right colic artery (a. colica dextra) is a (rami pancreatici) into the gland, which anasto
small vessel which arises as the last branch of the mose in the caudal third of this limb with sim
common colic. It runs in the right mesocolon to ilar branches from the cranial pancreaticoduo
ward the right colic flexure, giving off branches denal. The duodenal branch (ramus duodenalis),
to the distal part of the right colon and adjacent as it runs obliquely to the mesenteric border of
part of the transverse colon. It forms a definite the descending portion of the duodenum, sup
arcade with the accessory middle colic and a plies a small branch to the duodenum which, in
weaker anastomosis with the colic branch of the the region of the caudal flexure, anastomoses
ileocecocolic. with the first jejunal artery. The main branch
The ileocecocolic artery (a. ileocecocolica), anastomoses on the duodenum with the larger
which, as the name implies, goes to the ileum, duodenal branch of the cranial pancreaticoduo
cecum, and colon, is the continuation of the denal artery.
common colic artery after the right colic arises, The jejunal arteries (aa. jejunales) (Fig. 4-59)
and is about four times larger than the right are 12 to 15 in number. Some of these are larger
colic. Before the main artery disappears between than others, and branch after they have traveled
the ileum and cecum, the colic branch (ramus only a few millimeters. They arise from the cau
colicus) arises and goes to the proximal part of dal or convex side of the cranial mesenteric ar
the right colon. Within the wall of the colon the tery. The proximal 8 to 10 vessels are covered on
colic branch anastomoses with the right colic ar each side by the two large jejunal lymph nodes.
tery. The accessory cecal artery (r. caecalis acces Usually the first five to seven arteries arise
sorius of Thamm 1941) arises near the colic closely together; rarely are they separated by
branch and runs to the ventral side of the ileo more than 3 mm. Following this cluster of ves
A b d o m in a l A o r ta 351
sels which go to the proximal part of the jejunum the whole small intestine. The cranial mesen
the artery is free of branches for about 1 cm. The teric artery and its branches are accompanied
cranial mesenteric then gives rise to the by the cranial mesenteric plexus of nerves, the
branches that go to the distal part of the jeju intestinal lymph trunk, and the cranial mesen
num, and finally to the two ileal arteries. The teric vein.
ileal arteries branch on approaching the intestine The caudal mesenteric artery (a. mesenterica
and join to form primary and secondary arcades, caudalis) (Fig. 4-60) arises from the ventral sur
which lie directly adjacent to the intestinal wall. face of the aorta about 4 cm. cranial to the ter
According to Noer (1943) tertiary arcades are mination of the aorta or opposite the caudal part
rudimentary and, when present, consist of com of the fifth lumbar vertebra. It runs caudoven-
munications between the branchings of the trally in the left mesocolon to the mesenteric
secondary arcades. There is little variation of the border of the left colon, where it divides into the
vascular pattern throughout the jejunum, ileum, proximally running left colic and the distally
and colon. running cranial rectal arteries. The caudal mes
The vasa recti leave the terminal arcades and enteric artery is accompanied by the caudal
go directly to the intestine. They are short and mesenteric plexus, which begins about 1 cm.
irregular. Numerous lateral twigs unite with sim from the origin of the artery. No veins or lymph
ilar adjacent vessels. Most bifurcate on entering vessels accompany the main caudal mesenteric
the wall of the intestine; these branches circle artery.
the gut on opposite sides and typically anasto The left colic artery (a. colica sinistra) is about
mose with each other on the antimesenteric 1.2 mm. in diameter. It follows the mesenteric
border. Eisberg (1924) states that the vasa recti border of the left colon anteriorly and anastomo
pierce the muscular coats in the mesenteric ses with the middle colic at the junction of the
quarters of the small intestine and the antimes proximal and distal halves of the left colon. No
enteric quarters of the large intestine. Eisberg arcade is formed; numerous vasa recti run from
also states that there is a well-defined arterial it to the left colon. It is accompanied by the left
anastomosis along the mesenteric border in ad colic vein and autonomic nerves.
dition to those between the vasa recti. These The cranial rectal artery (a. rectalis cranialis),
anastomoses are small and few in the duodenum. formerly known as the cranial hemorrhoidal, de
They are moderate in number in the jejunum scends along the mesenteric border of the left
and occur in large numbers in the ileum and colon and rectum. It supplies many branches
colon. Within the gut there are two arterial net to the rectum and may anastomose with the
works. The subserous one is well developed as middle rectal and caudal rectal arteries.
compared with man and is derived from the
P a ir e d V is c e r a l B ran ch es of A b d o m in a l
mural, or wall, network. The mural network lies
A orta
largely in the submucosa and is formed by direct
and plexiform anastomoses (Noer 1943). The renal arteries (aa. renales) (Fig. 4-53)
Branches are supplied at irregular intervals arise asymmetrically from the lateral surfaces of
from the proximal halves of the jejunal arteries the abdominal aorta. The right renal artery
to the jejunal lymph nodes, one of which lies on arises about 2 cm. cranial to the left renal artery
each side of the jejunal mesentery. in conformity to the more cranial position of the
The cranial mesenteric artery terminates in right kidney. This places it about 4 cm. caudal to
the proximal and distal ileal arteries (aa. ileacae the origin of the cranial mesenteric artery. In a
Thamm 1941). The proximal ileal artery forms a large dog, it is 5 cm. long and 4 mm. in diameter.
terminal arcade with the last jejunal artery and The left renal artery is about the same diameter
issues vasa recti to the proximal part of the ileum as the right, but is only about 3 cm. long. The
and the distal part of the jejunum. The distal fact that the postcava lies to the right of the
ileal artery bifurcates. The proximal branch aorta accounts for the greater length of the right
forms an arcade with the proximal ileal artery, renal artery, which crosses its dorsal surface.
whereas the distal branch runs about 6 cm. along Each renal artery supplies two or three branches
the mesenteric border of the ileum and at the to the caudal pole of the suprarenal gland, and a
ileocolic junction or proximal to it anastomoses small cranial ureteral branch to the ureter, al
with the accessory cecal artery. The first jeju though this vessel may arise from the aorta di
nal artery anastomoses with the duodenal part rectly. About 1 cm. outside the renal hilus each
of the caudal pancreaticoduodenal and all je divides typically into a dorsal and a ventral
junal and ileal arteries anastomose with each branch. According to Christensen (1952) each of
other. Thus a vascular channel is formed along these further divides into two, four, or even
352 Chapter 4. The H eart and A rte rie s
L. g a s t r i c
Common hepatic
C eliac -
C om m on c o l i c ---- ;
Cran. m e s e n t e n i c -
Sup ra re n a l aa
R. r e n a l -
Psoas m in o r- ■
Psoas m a jo r -
- Q u a d r a t u s lumborum
Coud. m esenteric-
I- -U m b ilica l
-Y -S e ve n th lu m ba r
Esophageal
Proper hepatic
Gastroduodenal 4 -
%
C ra n p a n c re a tic o d u o d e n a l- P
Ascend.nj colon
Fit.. 4-54. Diagram of the visceral branches of the aorta with their principal anastomoses, ventral aspect
354 Chapter 4. The H e a rt and A rte rie s
seven interlobar arteries, or they may not divide unilaterally, in addition to the sources already
at all before entering the hilus of the kidney. cited. Flint (1900) gives a classical description of
At the corticomedullary junction numerous the suprarenal gland and details its blood supply
relatively straight arcuate arteries continue the as shown in Table 5.
interlobar arteries in this zone. Only in mature Flint’s (1900) phrenic and accessory phrenic
or elderly specimens are the arcuate arteries arteries represent the phrenic part of the phren-
arched toward the cortex. These interanasto- icoabdominal of modern terminology, whereas
mose and give rise to the numerous inter his lumbar artery is the abdominal part. When
lobar arteries of the cortex. These in turn they reach the adrenal, Flint divides the vessels
give rise to the afferent vessels which are con into three groups: capsular, cortical, and medul
tinued as the glomerular arterioles. The renal lary. The capsular vessels are represented by a
arteries may be double in 20 per cent of dogs, rather poorly defined system of arterioles which
particularly on the left side. Fitzgerald (1940) arise from all the arteries going to the gland.
reported an incidence of 5 per cent, whereas These form a plexus from which most of its
Christensen (1952) found 29 double renal arter branches, of arteriole size, enter the cortex. The
ies in 117 specimens. Reis and Tepe (1956) ex arterioles divide and form capillaries which fol
amined 500 dogs and reported that 99.4 per cent low the reticular septa between the coiled col
had a single right renal artery, whereas 12.8 per umns of cortical cells which they nourish. The
cent had double left renal arteries, and in 0.4 per arteries of the medulla, according to Flint, num
cent the left renal arteries were triple. The renal ber about 50. They come from the capsular
artery and its larger branches are accompanied plexus and run at right angles to it through the
by satellite veins and plexuses of autonomic cortex to the corticomedullary junction where
nerves. Both the arteries and the veins have most turn and run short distances in this area as
companion lymphatics, according to Peirce they give off finer vessels which supply the
(1944). The left renal vein, like the right renal ar medulla. While traversing the cortex they neither
tery, is longer than its fellow and empties into divide nor supply any branches to it.
the postcava more caudally than does the op The arteries which supply the testes and ova
posite vessel. According to Kazzaz and Shank- ries arise from the ventral third of the circumfer
lin (1951), the dog has a system of stellate veins ence of the aorta in the mid-lumbar region or op
on the surface of its kidney. Those on the lateral posite the fibrocartilage between the fourth and
side drain into the interlobar veins; those on the fifth lumbar vertebrae. The right artery usually
medial side go directly into the renal vein. Ac arises several millimeters cranial to the left, in
cording to these authors, the cat has radially ar conformity with the more cranial location of the
ranged veins on the surface of its kidney. right gonad.
The suprarenal (adrenal) arteries (aa. supra- The testicular artery (a. testicularis) (Fig. 4 -
renales) in the dog usually arise from the phreni- 54) is also called the internal spermatic artery. It
coabdominal, the aorta, and the renal artery. is a small vessel, much longer and straighter than
Quite commonly the cranial pole of the adrenal its homologue, the ovarian artery. It runs later
gland receives a branch from the cranial mesen ally or craniolaterally across the sublumbar mus
teric artery and occasionally from the celiac also. cles and the ventral surface of the ureter. Each
Ljubomudrov (1939) states that there are 20 to makes a sweeping bend caudally to the deep
30 or more fine arteries that supply each gland, inguinal ring, and lies in a special plica of peri
and that some of these may arise from the artery toneum, which may be 3 to 4 cm. wide. At the
to the adipose capsule and from a lumbar artery, deep inguinal ring it becomes a constituent of
NO. OF
SOURCE D I S T R IB U T IO N
BRANCHES
the spermatic cord and runs to the testis with its dorsal branches of the spinal nerves as they enter
companion vein and nerve plexuses in the free the epaxial muscles caudal to the craniolaterally
border of the mesorchium. It is the only artery extending transverse processes. Here each artery
supplying the testis and epididymis. divides into a spinal branch (ramus spinalis),
The ovarian artery (a. ovarica), the homologue which runs with the nerve into the spinal canal
of the testicillar artery, varies in size and tortu (see discussion of vertebral artery for descrip
osity, depending on the age and past genital tion), and a dorsal branch (ramus dorsalis), which
activity of the bitch. The ovarian artery (Preuss runs dorsocaudally in the longissimus past the
1959) is also known as the utero-ovarian artery lateral surface of the cranially inclined articular
(Sisson and Grossman 1953, Barone and Pavaux process of the vertebra behind it. The latter
1962), or the internal spermatic artery (Dell- gives off many branches to the epaxial muscles
briigge 1940, Nickel, Schummer, and Seiferle in its course to the subcutaneous fat and skin
1960). The vessel arises from the ventral surface near the mid line. Small branches also leave the
of the aorta at the mid-lumbar level. Like its dorsal branch near the origin of the spinal
homologue, it lies ventral to the postcava. Each branch and run laterally on the ventral surfaces
gives a minute branch to the capsule of the kid of the transverse processes which they supply.
ney. Other twigs may go to the periaortic fat, The adjacent internal oblique and transversus
peritoneum, and the adventitia of the postcava. abdominis muscles receive delicate terminal
Medial to the ovary the vessel bifurcates or tri- twigs from these. The seventh lumbar arteries
furcates. The resultant branches are very tortu differ from the others in that they may arise as a
ous, even in immature females, as they course in common trunk from the terminal part of the
the broad ligament on their way to the ovary. aorta or from the middle sacral artery. Each
Small branches from these supply the peritoneum vessel as it courses laterally runs dorsal to the
and fat of the ovarian bursa, and the uterine lumbosacral nerve plexus and divides into dorsal
tube. One or more of these branches continue and caudal branches at the iliosacral joint. The
caudally to the ovarian end of the uterine horn dorsal branch enters the epaxial muscles by pass
and anastomose with the uterine artery, which ing through the cranial angle formed by the
runs cranially in the broad ligament. The ovarian ilium and the vertebral column. The caudal
artery is accompanied by a plexus of sympathetic branch runs into the pelvis in company with the
nerves, a satellite vein, and lymph vessels. sympathetic trunk. It supplies this, and at the
ganglion impar continues into the ventral sacro
P a r ie t a l B r a n c h e s of A b d o m in a l A o r t a coccygeal muscles. The caudal branch may also
send a branch laterally to the pelvic surface of
The paired lumbar arteries (aa. lumbales) are the wing of the ilium. The lumbar arteries are
seven in number. The first two pairs arise from accompanied by satellite veins.
the thoracic aorta, and the last five pairs come The paired phrenicoabdominal artery (a.
from the dorsal surface of the abdominal aorta. phrenicoabdominalis) (Fig. 4-53) is the common
Those in the cranial part of the series are 4 mm. trunk for the phrenic arteries and the cranial ab
apart at their origins, and usually the right dominal artery, according to Marthen (1939). It
arteries arise 3 to 6 mm. caudal to those on the arises from the lateral surface of the aorta be
left. The vessels are progressively closer at their tween the cranial mesenteric and renal arteries.
origins as they are traced caudally, and the last The right artery occasionally arises from the cor
lumbar arteries may arise from a common trunk. responding renal vessel, and the phrenic and ab
Each pair of vessels runs caudolaterally across dominal parts may arise separately. The artery
the ventrolateral surface of the body of the lum runs caudolaterally, parallel to the ribs, and
bar vertebra of the same serial number. The first crosses the ventral surface of the psoas muscles
three or four pairs arise opposite the disc cranial and the dorsal surface of the adrenal gland. (The
to the vertebra of the comparable number; the corresponding vein grooves the ventral surface
last three or four pairs arise opposite the body of of the adrenal gland.) As it enters the fat lying
the comparable vertebra. For this reason the last lateral to the sublumbar muscles it sends a small
several arteries are less oblique as they run on phrenic branch forward to a part of the dia
the sides of the vertebral bodies, covered in their phragm and a branch into the psoas muscle.
courses by the sublumbar muscles which they The main phrenic artery (a. phrenica) arises
supply. Near their origins each vessel gives off a within 1 cm. of the origin of the phrenicoab
small, short nutrient branch which enters the dominal and runs forward to ramify on the ven-
body of the vertebra. The arteries parallel the trocaudolateral surface of the crus of the dia-
(Text continued on page 359.)
356 Chapter 4. The H e a rt an d A rte rie s
Gastroduodenal a.\
Rt. gastroepiploic a. . C e lia c el
s ' S p l e n i c a.
Cranial
p a n cre atico
d u o d e n a l a. „ L .gastroepiploica.
' A o rta
Caud. pancreaticaduodenal a.
F ig . 4-55. Celiac and cranial arteries, mesenteric, ventral aspect. (Stomach reflected cranially.)
A b d o m in a l A o r t a 357
Di aphragm
Esophageal branches
P r o p e r hepat
C a s t r o d u o d e n a l a.
Common b i l e d u c t -
Portal
Duodenum-
Panereas
Epi pl oic br a nch es
Rt. g a s t r o e p i p l o i c a.
Common h ep a ti
R t medial
lobe
ToG.B., rt. med. r q u a d r a t e l o b e s
To c y s t i c d u c t , quad r ot e v
R t l at . l o b e I. m e d l a l l ob es
F ig . 4-58. Blood supply of the spleen. (The cranial border is reflected laterally.)
A b d o m in a l A o r t a 359
phragm. In its subperitoneal course along the perforates the abdominal wall between the lum
medial border of the dorsal extension of the ten bar and inguinal portions of the internal oblique
dinous center of the diaphragm it usually sends and the cutaneus trunci to reach subcutaneous
two branches ventrolaterally which redivide structures. It is stellate in form as it subdivides
laterally as they cross the tendinous part of the into diverging branches which arborize ventrally
diaphragm and enter its muscular periphery. The and caudally. One or two strong branches spray
branches in general follow the course of the mus out over the rump and loin to anastomose with
cle fibers peripherally and anastomose within their fellows at the mid-dorsal line. The caudal
the muscle with the phrenic branches of the branches extend about halfway to the caudal
tenth, eleventh, and twelfth intercostal arteries. border of the thigh before they anastomose with
The cranial abdom inal artery (a. abdominalis cutaneous twigs of the caudal femoral artery
cranialis) continues the direction of the parent which emerge from the substance of the ham
artery into the abdominal wall after the phrenic string muscles. The ventral branches are longer
vessels arise. It runs medial to the narrow apo than the others as they run downward and back
neurosis of origin of the transversus abdominis, ward over the thigh and stifle joint to the crus.
then perforates the muscle to ramify extensively The most cranial of the ventral branches ends in
between it and the internal abdominal oblique. the fold of the flank. The cranial branches are
Usually the vessel divides after perforating the smallest as they radiate in the subcutaneous fat
transversus abdominis. The cranial branch runs of the caudal parts of the loin and abdomen.
toward the costal arch; the caudal branch di The deep circumflex iliac artery is accom
verges from it and supplies the middle zone of panied by a satellite vein which lies caudal to it.
the lateral abdominal wall. The cranial ab At the lateral border of the psoas major it is
dominal artery anastomoses cranially with the joined by the lateral cutaneous femoral nerve.
phrenic vessels, ventrally with the cranial and
caudal deep epigastric arteries, and caudally A r t e r ie s of the P e l v ic L im b
the deep surface of the internal oblique and di branch enters the laterally lying iliopsoas. Its
vides typically into dorsal and ventral branches. principal intra-abdominal branch is the short
On reaching the caudal border of the trans pudendoepigastric trunk. After leaving the ab
versus abdominis they arborize on the medial domen the deep femoral passes between the
surface of the internal oblique. Branches perfo quadriceps femoris and the medially lying
rate this muscle to reach the external oblique. pectineus. Before reaching the large adductor
The ventral branch lies about 2 cm. from, and muscle the artery divides into the caudally run
parallel to, the lateral border of the rectus ab ning obturator branch and the slightly larger
dominis. It anastomoses with the caudal deep caudodistally running medial circumflex femoral
epigastric artery. The dorsal branch passes dor artery.
sally and anastomoses with the deep branch of The pudendoepigastric trunk (truncus puden-
the deep circumflex iliac artery. The cremasteric doepigastricus) (Figs. 4-49, 4-65) is short,
artery (a. cremasterica) is a small twig which averaging slightly over 2 mm. in length. It may
runs toward the inguinal canal and supplies the extend to the abdominal inguinal ring before
fat lying around the abdominal inguinal ring. It bifurcating terminally. Rarely, it is absent, in
enters the inguinal canal and supplies the cre- which case the terminal deep caudal epigastric
master muscle. According to Joranson et al. and the external pudendal arteries arise sepa
(1929) it, along with the artery of the ductus def rately from the deep femoral.
erens, continues to the testis. Blood supplied The deep caudal epigastric artery (a. epigas-
through these vessels is not sufficient to prevent trica caudalis profunda) runs cranially to lie on
atrophy of the organ when the testicular artery the dorsal surface of the rectus abdominis di
is interrupted. rectly under the peritoneum. When it reaches
The larger arteries of the thigh of the dog pose the caudal border of the sheath of the rectus ab
a real problem as regards their homology with dominis it runs under this. It grooves the dorsal
the similarly named arteries in man. In the dog, surface of the muscle as it parallels the linea alba
two large arteries leave the femoral to supply the in its branched, cranial course. At about the
thigh. The first of these is the cranial femoral, junction of the cranial and middle thirds of the
which supplies the quadriceps, and the second is abdominal part of the rectus abdominis it anas
the caudal femoral, which supplies the distal tomoses with the cranial deep epigastric artery
parts of the “hamstring” muscles. Man has in the substance of the muscle. It sends three or
neither of these vessels. The lateral circumflex more branches laterally, which anastomose with
femoral of man is the main supply to the quad the terminal abdominal branches of the deep
riceps, whereas in the dog it seems most feasible circumflex iliac artery in the middle third of the
to regard a branch of the cranial femoral as the abdomen. Small medial branches supply the
homologous artery. The lateral circumflex fem relatively avascular linea alba. It is accompanied
oral artery of Ellenberger and Baum (1943), by a small satellite vein.
Sisson and Grossman (1953), and Bradley and The external pudendal artery (a. pudenda ex
Grahame (1959), for the dog, is comparable to terna) (Fig. 4-61) arises as the ventral terminal
the superficial circumflex iliac of man and should branch of the pudendoepigastric trunk. It leaves
be so named in the dog. The caudal femoral of the abdominal cavity through the caudal part of
the dog supplies much of the area supplied by the inguinal canal. After emerging through the
the perforating branches of the deep femoral of superficial inguinal ring it continues caudoven-
man. trally to the cranial border of the gracilis. It then
The deep femoral artery (a. profunda femoris) arches cranially and becomes related to the
(Fig. 4-65) is about 2 mm. in diameter and 3 cm. dorsal surface of the superficial inguinal lymph
long, about half of it lying within the abdomen node. Here in the fat-laden superficial abdomi
and half outside. It arises from the caudomedial nal fascia, it gives origin to the caudal superficial
surface of the external iliac at an angle of about epigastric artery which continues along the
45 degrees, and runs obliquely distocaudally mammary row. At this place the external puden
over the medial surface of the external iliac vein dal starts the second arc of the sigmoid flexure it
to leave the abdominal cavity by passing through describes and, running ventral to the superficial
the caudal part of the vascular lacuna. It sends a inguinal lymph node, enters the fat ventral to
small branch over the pelvic surface of the pubis the subpubic tendon. After emerging caudally
to the medial coccygeal muscle, where it anasto from between the legs, it terminates in the vulva
moses with the obturator branch which ascends as the cranial labial branch (ramus labialis
through the obturator foramen. Another small cranialis). In the male, the comparable vessel
A b d o m in a l A o r t a 361
Cran. p a n c r e a t i c o d u o d e n a K j
C o l i c br R i g h t colic
I leocecocolic . M i d d l e c ol i c
Common c o l i c
—A o r t a
- - C r a n . m esenteric
CaudaI
pancreat i co-
duo de na l
— 1m ~ - L e f t col ic
I l e a l aa —
— Vasa r e c t i
M i d d l e col i c
Ri gh t col ic
Col ic br _ Cel i a c
" _ p h re n ic
Caud. p a n c r e a t i c o M
__ J^ H t —Rt. cron, abdominal
duodenal
— ^ i - L . p h r e n i c o a bd om inal
-1
- Renal a a.
-Aorta
A n t i mesenteric - T e s ti c u la r aa.
i l ea l b r
I l e a l ao.
- c a ud mesenteric
~Cranial r ec ta l
F ig . 4-60. Branches of the cranial and caudal mesenteric arteries, ventral aspect.
A b d o m in a l A o r t a 36 3
may be extremely small or absent. Rarely it is supply the caudal part of the hip joint capsule
large. It terminates as the cranial scrotal branch and the neck of the femur. It ends in the semi
(ramus scrotalis cranialis). When the cranial membranosus. Small branches also supply the
scrotal branch is absent, the external pudendal quadratus femoris, pectineus, external obturator,
continues along the prepuce as the caudal super and the semimembranosus. The main terminal
ficial epigastric artery. The external pudendal part of the vessel enters the semimembranosus
artery is accompanied by a laterally lying satel about 3 cm. from the tuber ischii and anasto
lite vein. moses with the caudal gluteal artery. A deeper
The caudal superficial epigastric artery (a. portion of the vessel descends in the adductor
epigastrica superficialis caudalis) (Fig. 4-62) is near the femur and anastomoses with the medial
small in the male as it runs forward to supply the circumflex femoral artery.
prepuce, superficial inguinal lymph node, fascia, The m edial circumflex fem oral artery (a. cir-
fat, and skin. It usually ends before it reaches the cumflexa femoris medialis) (Figs. 4-63, 4-65) is
umbilicus. In the female it may be the largest the terminal branch of the deep femoral. It is
artery of the abdominal wall. Its size is in direct slightly larger than the obturator branch, and as
proportion to the state of development of the it obliquely crosses the surface of the vastus me
mammary glands it supplies. From its origin, on dialis it sends a twig to it. It arborizes extensively
the convex side of the second arc of the external in the deep part of the adductor. After reaching
pudendal artery, it runs forward deep to the the caudal surface of the femur distal to the tro
inguinal mammary gland and medial to its nip chanter major it gives rise to the nutrient artery
ple. It supplies many branches to this potentially o f the fem u r (a. nutricia femori), which enters
pendulous gland. As it advances cranially it the nutrient foramen at the junction of the prox
enters the caudal abdominal mammary gland imal and middle thirds of the caudal surface of
and divides into several branches, some of which the bone. A small branch runs laterally, crosses
become subcutaneous around the nipple. Many the facies aspera, and anastomoses with the lat
of these branches anastomose with like branches eral circumflex femoral artery in the quadriceps.
from the cranial superficial epigastric artery be The main vessel continues distocaudally through
tween the cranial and caudal mammary glands. the adductor and terminates in the middle third
The vessel takes on special significance because of the semimembranosus by anastomosing with
of the high incidence of mammary tumors in the the caudal gluteal artery.
dog. It is accompanied by a satellite vein and
lymphatics which drain into the superficial in Femoral Artery
guinal lymph node.
The obturator branch (ramus obturatorius) The femoral artery (a. femoralis) (Figs. 4-63,
(Figs. 4-63, 4-65), a branch of the deep femoral, 4-64, 4-65) continues the external iliac artery
after running a few millimeters caudally, ascends from the vascular lacuna through the thigh. It in
through the cranial part of the obturator fora turn is continued back of the stifle joint by the
men, lateral to the obturator nerve. This vessel popliteal artery. Throughout the proximal half
takes the place of the obturator artery, a branch of the thigh it lies cranial to its satellite vein and
of the internal iliac as found in man and horse. It either caudal or medial to the saphenous nerve.
furnishes branches to both the medial and lateral Here, it lies superficially in the fem oral triangle
coccygeal and the external and internal obtura (trigonum femorale), which is the favored site
tor muscles. On the pelvic surface of the pubis it for taking the pulse of the dog. The femoral vein
anastomoses with the small branch from the is large and securely placed in the femoral trian
deep femoral which supplies the cranial portion gle so that venipunctures can be made without
of the medial coccygeal muscle. Most of the compressing it. The femoral vessels are covered
intrapelvic portion of the obturator branch ter only by the thin skin and the deep medial fem
minates in the internal obturator muscle. The oral fascia. Upon leaving the femoral triangle at
main part of the obturator branch supplies the the middle of the thigh, the femoral artery and
proximal end of the massive adductor muscle. vein incline laterally along the medial border of
About 5 mm. distal to the origin of the branch insertion of the adductor, where they are cov
which ascends through the obturator foramen, a ered by the caudal belly of the semimembrano
branch about 0.5 mm. in diameter extends lat sus. Upon reaching the popliteal surface of the
erally to enter the trochanteric fossa and supply femur, the femoral vessels are continued be
portions of the muscles which insert there. Its tween the lateral and medial heads of the gas
chief importance concerns the branches which trocnemius as the popliteal vessels. The
364 C h ap ter 4. The H e a r t an d A r te r ie s
L o c a tio n o f r i g h t \
tu b e r is c h ii '
,Ext. pudendal o.
P e r in e a l a. N
Superf. ing u in a l lymph node
/
Caud. su pe rf e p i g a s t r i c
R o o t o f penis _
T e s tis '
Cranial superficial - I
epigastric Q 't v ' C r a n ia l a b d o m in a l
n ip p le
-XIII rib
-Ext. s h e a th o f
r e c t u s abdom inis
Caudal superficial -
epigastric a f v S uperf. abdom inal fa s c ia
r e f le c te d
V a g in a l p r o c e s s t f a t
covering ext. inguinal r in g
-R ound ligament
Femoral v- o f u te ru s
Pectmeus m-
-External
pu d endal a. + v.
Fic. 4-62. Superficial vessels of the abdomen, ventral aspect. (From Miller 1958.)
366 C h apter 4 . The H e a rt and A r te r ie s
branches of the femoral artery in the order in mur. It is accompanied by a satellite vein, which
which they arise are: superficial circumflex iliac, lies distal to it, and the large femoral nerve,
cranial femoral, muscular branches, saphenous, which lies proximal to it. The lateral circumflex
descending genicular, and caudal femoral. femoral of this description is comparable to the
The superficial circumflex iliac artery (a. cir- transverse ramus of man. Hermann (1940) and
cumflexa ilium superficialis) is a small artery Ellenberger and Baum (1943) state that the lat
which runs dorsocranially over the medial sur eral circumflex femoral may arise from the fem
face of the rectus femoris to reach the septum oral separately. There is disagreement about the
between the caudal belly of the sartorius and homology of this vessel in man and the domestic
the tensor fascia lata. It arises from the lateral animals.
surface of the femoral artery close to the cranial The muscular branches (rami musculares) of
femoral artery or it may arise from this vessel by the femoral are variable in origin, number, size,
a short common trunk. Its first branch usually is and distribution. As the femoral artery passes
the principal vessel to the tensor fasciae latae. It distally from the point where the cranial femoral
also sends branches to the underlying rectus fem arises, it lies adjacent to the cranial border of the
oris. The main vessel then bifurcates into dor vastus medialis proximally and inclines laterally
sal and ventral branches, under the thin, caudal around its caudal border distally. Distal to the
belly of the sartorius. The dorsal branch supplies origin of the cranial femoral, a muscular branch
the proximal fourth of the cranial belly of this arises from the lateral surface of the femoral,
muscle and becomes subcutaneous at the cra which bifurcates into cranial and caudal vessels.
nial ventral iliac spine; the ventral branch runs Two separate diverging vessels may arise di
distocranially and supplies the middle portion rectly from the femoral. The caudal vessel may
of the cranial belly of the sartorius. It has a sat end in the femoral sheath or terminate in the
ellite vein. proximal end of the pectineus. The slender cra
The cranial femoral artery (a. femoralis cra nial branch goes to the deep surface of the cau
nialis) leaves the caudolateral surface of the fem dal belly of the sartorius. The next muscular
oral about 5 mm. from the abdominal wall. It im branch is the largest, and most constant. It
mediately disappears between the rectus femoris leaves the caudal side of the femoral, extends
and the vastus medialis. It is about 2 mm. in di distocaudally, and crosses the end of the pectin
ameter at its origin, where it is crossed cranially eus and then the adductor. Upon reaching the
by the saphenous nerve and caudally by the fem cranial border of the gracilis, at the junction of
oral vein. Usually its first branch is small and its proximal and middle thirds, it runs under it to
enters the iliopsoas near its insertion on the tro enter its deep surface. It sends one or more
chanter minor. Another small branch may ex branches to the pectineus and the adductor.
tend lateral to the proximal part of the cranial Usually three branches go to the adjacent mus
border of the vastus lateralis and enter the cau cles before the femoral sinks into the substance
dal, deep part of the tensor fasciae latae. As the of the thigh. Often two of these arise from the
main artery loses contact with the iliopsoas to cranial side of the vessel and after short courses
bend distally in the quadriceps it sends a large disappear into the vastus medialis. One or two
branch caudally, which bifurcates upon reach smaller twigs quite regularly leave the caudal
ing the neck of the femur. The resultant ascend side of the femoral and disappear into the distal
ing branch (ramus ascendens) sends many twigs part of the adductor. Under cover of the semi
to the cranial part of the hip joint capsule, then membranosus, distal to the descending genicular
continues dorsally to supply the insertions of the artery or in common with it, a medium-sized
deep and middle gluteal and the tensor fasciae muscular branch arises from the femoral. It runs
latae muscles. As the descending branch (ramus caudodistally and ramifies in the adductor and
descendens) bends distolaterally around the cau semimembranosus. It anastomoses with the cau
dal surface of the rectus femoris it sends dal femoral artery and is accompanied by a sat
branches to the vasti, rectus femoris, and tensor ellite vein.
fasciae latae. It is the principal source of blood The saphenous artery (a. saphena) (Fig. 4-
to the quadriceps. Circling the lateral surface of 64), less than 1 mm. in diameter, arises from the
the femur in the vastus lateralis distal to the tro medial surface of the femoral just before the
chanter major is the small lateral circumflex fem femoral disappears under the semimembranosus.
oral artery (a. circumflexa femoris lateralis). It It runs distally across the medial surface of the
usually forms a feeble anastomosis with the me semimembranosus, where it and its accompany
dial circumflex femoral artery caudal to the fe ing vein and nerve lie between the converging
A b d o m in a l A o r t a 367
borders of the caudal belly of the sartorius in of the femorotibial joint capsules. The descend
front and the gracilis behind. It supplies small ing genicular is the main blood supply to the
twigs to these muscles. As it passes over the me stifle joint. It is accompanied by a satellite vein.
dial surface of the stifle it sends a single or a The caudal femoral artery (a. femoralis cau-
paired m edial genicular artery (a. genus medi dalis) (Fig. 4-66) is about 2 mm. in diameter as
alis) to the skin and superficial fascia covering it arises from the caudolateral surface of the fem
the medial side of the joint. A proximal branch oral approximately 1 cm. proximal to its en
runs dorsally and anastomoses with the superfi trance into the gastrocnemius to become the
cial part of the deep circumflex iliac artery. Op popliteal artery. Usually it runs caudodistally on
posite or distal to the tibial condyle the saphe the gastrocnemius and sends its first and largest
nous terminates in a small dorsal and a larger branch laterally into the distal end of the biceps
plantar branch. femoris but this branch may come from the fem
The dorsal branch (ramus dorsalis) obliquely oral directly. Most of this vessel ascends in the
crosses the subcutaneous part of the tibia in its muscle but one or more twigs continue through
course distocranially. It then bends around the the muscle to end as cutaneous branches to the
medial border of the cranial tibial muscle and caudolateral part of the thigh. Another twig en
continues distally in the superficial fascia cover ters the insertion of the adductor. The small ar
ing this muscle, to traverse the flexor surface of tery which crosses the lateral surface of the fe
the tarsus. Opposite the tarsus or distal to it, the mur to enter the vastus lateralis comes from
dorsal branch anastomoses with the superficial either the femoral or the caudal femoral. Several
ramus of the cranial tibial. Two or three delicate branches run distally and are distributed to both
rami leave the dorsal branch and supply the fas heads of the gastrocnemius. About 2 cm. from its
cia, periosteum, skin, and cranial tibial muscle. origin the caudal femoral artery crosses the lat
In the proximal part of the metatarsus it termi eral surface of the tibial nerve and sends a large
nates in the three superficial dorsal metatarsal branch proximally into the semimembranosus.
arteries, which are described under the heading, Coming from this muscular ramus, or arising
“Arteries of the Hindpaw.” independently distal to it, is a smaller branch
The plantar branch (ramus plantaris) is the di to the semitendinosus. The terminal part of
rect continuation of the saphenous artery in the the vessel becomes cutaneous in the upper
crus after the dorsal branch arises. It lies medial caudal part of the crural region. The large de
to the tibia opposite the cleft between the me scending branches of the caudal femoral artery
dial head of the gastrocnemius and the bone. It supply the gastrocnemius; the large ascending
soon becomes related to the flexors of the digits, branches are the chief supply to the ham
and with its small accompanying vein and tibial string muscles. Their principal anastomoses are
nerve crosses the medial surface of the tarsus to with the caudal gluteal artery, but there are
enter the metatarsus. It has one prominent also anastomoses with branches of the deep fem
branch in the crus, the peroneal (fibular) artery oral and with the muscular branches of the fem
(a. peronea [fibularis]). This vessel arises near the oral which reach the hamstring muscles. The
middle of the crus from the cranial side of the caudal femoral artery is accompanied by the
plantar branch. It runs distally and crosses the large proximally lying lateral saphenous vein.
tibia obliquely to reach its lateral side. In this This receives the veins which accompany the
course it runs between the tibia and the closely branches of the artery. The main lymphatics
joined tendons of the caudal tibial and long digit from the limb distal to the stifle ascend over the
al flexor muscles. The peroneal artery ends in gastrocnemius in relation to this artery.
the collagenous tissue of the tarsus.
The descending genicular artery (a. genus de- Popliteal Artery
scendens) usually arises from the femoral distal
to the origin of the saphenous, but it may arise in The popliteal artery (a. poplitea) (Fig. 4-66) is
common with it. It runs distally between the vas the continuation of the femoral through the pop
tus medialis and the semimembranosus to the liteal fossa. It begins by entering the gastrocne
medial surface of the stifle joint. In this course it mius and terminates in the interosseous space
sends two or more small, short twigs into the vas between the tibia and fibula distal to their heads.
tus medialis. It lies at a deeper level than does It divides into the small caudal tibial and the
the medial genicular artery. Upon reaching the much larger cranial tibial artery. Passing be
medial epicondyle it divides into articular tween the two heads of the gastrocnemius, it
branches. End-branches supply the medial part crosses the medial surface of the superficial digit-
of the femoropatellar and the medial division (Text continued on page 371.)
368 Chapter 4. The H e a rt and A rte rie s
-M. p e c tin e u s
Deep fe m o r a l a.r v.- -
-M. a d d u c to r longus
Superf. ci r c u m f l e x
i l i a c a.* v.-
C r a n . fe m o r a l a.
L a t c irc u m fle x _
fe m o ra l arv.
Med. c i r c u m f l e x
fe m o r a l a.r V.
F e m o ra l a.w. - - " O b tu ra to r a. v v.
M u s c u l a r br.
D e s c .q e n ic u la r-
a.v v-
F ig . 4-63. Arteries and veins of the thigh, deep dissection, medial aspect.
A b d o m in a i A orta 369
Deep fe m o ra l a.rv.'
M. rectus f e m o r is '
■ M u s c u la r br.
M v a s t u s med.
F e m o r a l o tr v.
---- M. semimembranosus
M u s c u l a r br - M. s e m i t e r d i n o s u s
S a p h e n o u s a.vv.
Med. g e n i c u l a r a r v
M. g a s t r o c n e m i u s
D o rsa l br. s a p h e n o u s a.
Fi b u la r a rv
Fic. 4 -64 Artenes and veins of the thigh, superficial dissection, medial aspect
370 Chapter 4. The H e a rt and A rte rie s
Median s a c r a l -
A o rta
Deep circumflex
iliac
R. int. iliac
Umbi I ical
11iolumbar
(parietal br.
Intiliacl .
(visceral br
Caudal gluteal
R .e x t iliac
Pudendo epiqasfric
tru n k
Caud. abdominal
Superf. circumflex
i I iac
C ran ia l femoral
~Med. circumflex
L at ci rcumflex femoral femoral
F e m o ra l
/o/
F ig . 4-65. Diagram of the arteries of the pelvis and thigh, medial aspect.
A b d o m in a l A o r t a 371
al flexor, and over the flexor surface of the stifle supplies the long digital extensor as it lies in the
joint. It inclines laterally under the popliteus sulcus muscularis of the tibia, and smaller twigs
and, upon leaving it, perforates the origin of the are distributed to the stifle joint capsule. The
flexor hallucis longus to reach the interosseous second muscular branch runs under the long ex
space. It is accompanied by a small satellite vein. tensor and is distributed to the cranial tibial mus
It has small genicular and muscular branches. cle. Small anastomoses exist between the cranial
The caudal genicular arteries (aa. genus cau- tibial and the caudal femoral and the dorsal
dales) are small branches which run to the cau branch of the saphenous. The superficial branch
dal surface of the stifle joint. Opposite the fem (ramus superficialis) is very small in diameter,
oral condyles, a medium-sized genicular branch but long. It reaches the superficial crural fascia
arises from each of the medial and lateral sur between the peroneus longus and the long digit
faces of the popliteal artery and diverge from al extensor at the beginning of the distal third
each other to reach the distal ends of the medial of the crus. Here, in association with the super
and lateral collateral ligaments. These vessels ficial peroneal nerve, it sends a branched twig to
then arborize on the deep surfaces of the medial the lateral malleolus. The artery continues to the
and lateral heads of the gastrocnemius. Two or flexor surface of the tarsus, where it anastomoses
three small articular branches leave the cranial with the dorsal branch of the saphenous artery.
surface of the popliteal and supply the caudal If this does not occur it continues into the hind
part of the femorotibial joint capsule and the paw as the superficial dorsal metatarsal artery II.
cruciate ligaments. The greatest vascular supply
to the stifle joint comes from the caudal side. Arteries of the Hindpaw
The muscular branches (rami musculares) are
represented by two branches to the gastrocne The cranial tibial artery is continued opposite
mius which also go to the collateral ligaments, the talocrural joint as the dorsal p ed al artery (a.
and by a branch which leaves the caudal surface dorsalis pedis) (Fig. 4-70). Several small
of the popliteal artery and runs to the caudal branches arise from it to supply adjacent struc
surface of the popliteal muscle. Before the pop tures. The m edial an d lateral tarsal arteries (a.
liteal artery terminates it gives a branch to the tarsea medialis et a. tarsea lateralis) run deeply
small, proximal tibiofibular joint capsule which to the sides of the tarsus and end in the collateral
continues beyond this to descend in the flexor ligaments. The transverse m etatarsal artery (a.
hallucis longus. metatarsea transversa) (Fig. 4-67) is a small ves
The caudal tibial artery (a. tibialis caudalis), sel which leaves the lateral side of the dorsal
less than 1 mm. in diameter, leaves the caudal pedal artery at the proximal end of the metatar
surface of the popliteal at the interosseous space sus and runs transversely to disappear in the lig
and plunges into the flexor hallucis longus. It amentous tissue of the lateral and plantar sides
runs distally in this muscle, giving off medial and of the proximal end of the metatarsus. It sends
lateral branches. It supplies the nutrient artery two or three branches proximally to be distrib
o f the tibia (a. nutriciae tibiae), which enters the uted to the deep structures of the flexor surface
nutrient foramen on the caudal surface near the of the tarsus. Distally it gives rise to the third
lateral border at the junction of the proximal and and fourth deep dorsal metatarsal arteries.
middle thirds of the tibia. The dorsal pedal artery becomes the perforat
The cranial tibial artery (a. tibialis cranialis) ing metatarsal artery (a. metatarsea perforans)
(Figs. 4-66, 4-67), about 2 mm. in diameter, as it passes from the dorsal to the plantar surface
continues the popliteal artery between the tibia of the hindpaw, by passing between the proxi
and fibula after the caudal tibial arises. It in mal ends of the second and third metatarsal
clines laterally as it runs distally. It crosses under bones.
the peroneus longus to gain the deep surface of The arteries of the metatarsus and digits are
the long digital extensor. Here it is partly sep similar to those of the metacarpus and digits.
arated from the bone by the small, flat extensor Forepaws and hindpaws are similar also in that
hallucis longus which lies lateral to the artery. the main arteries supplying them lie on their
The superficially lying cranial tibial muscle, and flexor sides.
the lateral digital extensor muscle receive almost The superficial dorsal metatarsal arteries II,
their entire blood supply from branches of the III, and IV (aa. metatarseae dorsales superfici-
cranial tibial artery. The largest three or four ales II, III, et IV) (Fig. 4-70) arise subcutane-
muscular branches arise from the first 2 cm. of ously, over the tendon of the long digital exten-
the artery. The first branch runs proximally and (Text continued on page 377.)
372 Chapter 4. The H e a rt and A r te r ie s
Saphenous a.
- -Muscular br.
Desc g e n i c u l a r a
M .g a s t r o c n e m iu s -
-Caud. g en icu la r a.
Caud. tibial a.
Cran. t i b i a l a.------
Tibia
M .g astrocnem ius
Tendon of m. semitendinosus
M t i b i a l is c r a n .
P la n ta rb r F i b u l a r a .- -
saphenous a A c h il ie s te n do n
Tendon o f
m.flex. digit,
lo n g u s
Pla nta r
metata r s a I a.I- Plantar br:
saphenous a.
Plantar
Tendon o f rn.
d ig i t a l aa.I
flex digit s u p e r f
Tendon o f m.
abductor d i g i t i qumti
Superf. p l a n t a r
metatarsal aa II,III IV
Deep p la n t a r
m e t a t a r s a l a .ll
A.to digital
P l a n t a r common footpad
d i g i t a l a a. II, III, IV
L a t p lan tar - -
p r o p e r digital a . l l
M ed p l a n t a r -
p ro p e r digital o IV
P la n ta r br_ _
sa p h e n o u s o
P la n ta r br
sa p h e n o u s a.~
L a t p l a n t a r a .-
M ed p l a n t a r a.
P la n ta r
Tendon of I
m e t a ta r s a l a.1 - -
m.flex. d i g i t
lo n g u s - - 4 Tendon o f m.
fle x d i g it su p p rf.
M e d ia l
P la n ta r d ig ita l a .l- p la n ta r a -------i
Tendon of m.
flex, d ig it, prof.
Lat. p la n ta r a.
P e r f o r a t in g
m e ta ta r s a l a
D eep p l a n t a r «
m eta ta r s a I a a. Zi, III, IV
Tendon o f m.
Tendon of m.
flex. d ig it. s u p e r f
flex, d i g i t prof.
-S u pe rf. p la n t a r
m e ta ta rs a l
aa II, III, IV
P la n ta r com m on « ; £
d ig i to I a a . II, III, IV
Med. p l a n t a r p r o p e r-
d i g i t a l a.IU
D orsal pedal a.
D o rs m e ta ta r s a l a V
S u p e rf, dors.<-
m e ta ta r s a I aa. II, III, IV
Deep dorsal
m e ta ta rs a l aa. 11,111, IV
Dors, common*---
d ig ita l aa 11, 111, IV
Med. d a rs. p roper
d ig ita l a. Ill
P la n ta r com m on
d i g i t a l a 111 Lot. d o rs, p ro p e r
d ig ita l a III
M ed. p l a n t a r
p ro p e r d ig ita l a.IV Lot- p la n ta r p ro p er
d ig ita l a.Ill
Fic. 4-70 '\rtenes of the right hindpaw and first digit, dorsal aspect
A b d o m in a l A o r t a 377
sor, from a small trunk formed by the anastomo larger than the largest tarsal branches. The lat
sis of the dorsal branch of the saphenous and the eral plantar artery usually arises about 1 cm.
superficial branch of the cranial tibial. Typically, proximal to the origin of the medial vessel. It
this bifurcates into medial and lateral branches runs diagonally distolaterally on the plantar lig
at the proximal end of the metatarsus. The me ament under the superficial flexor tendon and,
dial branch becomes the second superficial dor gaining the lateral border of the deep flexor ten
sal metatarsal artery, and the lateral branch di don, runs under it. The medial plantar artery
vides into the third and fourth. Occasionally, an runs at first along the medial border of the deep
arch formed by the anastomoses of the medial flexor tendon, then under it, to anastomose in
and lateral branches gives off the superficial set. the interosseous musculature with the lateral
The superficial branch of the tibial and the dor plantar artery. The plan tar m etatarsal artery I
sal branch of the saphenous may continue inde (a. plantaris metatarsea I) leaves the medial side
pendently to form the lateral and medial arteries of the medial plantar before it turns under the
of the dorsal superficial set. When a first digit or deep flexor tendon. It runs on the plantar sur
dewclaw is present (Fig. 4-70), its dorsal part is face of the first digit as the plan tar digital ar
supplied by the medial and lateral dorsal proper tery I (a. digitalis plantaris I). It anastomoses
digital arteries (aa. digitalis dorsales propriae). with the larger dorsal digital artery I. The short
These vessels arise from the dorsal metatarsal trunk formed by the anastomosis of the two
artery I (a. metatarsea dorsalis I), which comes plantar arteries in turn anastomoses with the
from the dorsal branch of the saphenous and the perforating metatarsal artery to form the plantar
dorsal pedal artery. When the first digit is miss arch (arcus plantaris). Occasionally two anasto
ing, as it usually is, the first dorsal metatarsal ar moses exist between these vessels. The perforat
tery becomes dissipated on the medial part of ing metatarsal limb is much the larger of the two
the metatarsus. The superficial dorsal metatarsal parts which form the arch. The metatarsal part
arteries II, III, and IV anastomose with the deep of the plantar branch of the saphenous artery
dorsal metatarsal arteries in the distal ends of the gives rise to the superficial set of plantar meta
intermetatarsal spaces. tarsal arteries.
The deep dorsal metatarsal arteries II, III, The superficial plantar metatarsal arteries II,
and IV (aa. metatarseae dorsales profundae) III, and IV (aa. metatarseae plantares superfici-
(Fig. 4-67) are similar to the comparable arteries ales) (Fig. 4-68) arise from the terminal part of
of the forepaw. (No first deep dorsal metatarsal the plantar branch of the saphenous as it crosses
artery is present, even when a first digit exists.) the superficial digital flexor tendons. In the clefts
The second deep dorsal metatarsal artery arises between the digits they anastomose with the
from the dorsal pedal artery as it enters the sec larger, deep plantar metatarsal arteries.
ond iptermetatarsal space. The third and fourth The deep plantar metatarsal arteries II, III,
arteries arise from the transverse metatarsal ar and IV (aa. metatarseae plantares profundae II,
tery either separately or from a common stem. III, et IV) (Fig. 4-69) arise from the plantar
The three deep dorsal metatarsal arteries anasto arch. Between their origins, branches arise
mose with their superficial counterparts, to form which supply interosseous muscles. The deep
the dorsal common digital arteries. plantar metatarsal arteries are the chief blood
The dorsal common digital arteries II, III, supply to the hindpaw distal to the tarsus. As
and IV (aa. digitales dorsales communes) are they run distally, they lie between adjacent in
short vessels which give off medial and lateral terosseous muscles. They anastomose with the
dorsal proper digital arteries II, III, and IV (aa. corresponding members of the superficial set
digitales dorsales propriae II, III, et IV). Each near the distal ends of the intermetatarsal
dorsal common digital artery, or one of its spaces. One or two small anastomotic branches
proper branches, anastomoses by a short connec extend dorsally from about the last centimeter
tion deeply in the interdigital cleft with the com of each of these arteries and anastomose with
parable plantar common digital artery. either the deep dorsal metatarsal arteries or the
As the plantar branch of the saphenous artery dorsal common digital arteries. The plantar
approaches the tarsus it gives off several twigs, common digital arteries II, III, and IV (aa. digi
tarsal branches (rami tarsici), to the skin and fas tales plantares communes) are formed by the
cia of the medial surface of the tarsus. Distal and anastomoses of the corresponding members of
medial to the tuber calcanei the m edial an d lat the two plantar series. These and their branches
eral plantar arteries (a. plantaris medialis et a. are similar to the common palmar digital arter
plantaris lateralis) arise. These are only slightly ies. The superficial and deep dorsal and plantar
378 Chapter 4. The H e a r t an d A r te r ie s
arteries of the hindpaw are accompanied by cor the prostate gland of the male or toward the cra
responding nerves. With the exception of the nial part of the vagina of the female. It bifurcates
deep plantar metatarsal arteries, all are accom into cranial and caudal branches. The cranial
panied by satellite veins. branch continues the arch started by the parent
artery and gives origin to the small caudal artery
I n t e r n a l I l ia c A rtery of the ureter, the caudal vesical arteries and the
somewhat larger artery of the ductus deferens of
The internal iliac artery (a. iliaca interna) the male or the uterine artery of the female. The
(Figs. 4-71, 4-72), with its fellow and the caudal branch gives origin to three vessels.
smaller, unpaired median sacral artery, termi These are the middle rectal and urethral in both
nates the aorta at the level of the seventh lumbar sexes, the prostatic in the male, and the vaginal
vertebra. It is about half the diameter of the ex in the female.
ternal iliac, or 2.5 mm. in a medium-sized male The caudal ureteral artery (a. ureterica cau-
or non-gravid female. It varies in length from 0.5 dalis), with its satellite vein, runs forward on the
to 7 cm., and terminates as visceral and parietal dorsal surface of the ureter to anastomose with
branches. The umbilical artery is its only collat the cranial ureteral artery, a branch of the renal.
eral branch. The caudal ureteral artery lies in the lateral
The umbilical artery (a. umbilicalis) arises ligament of the bladder and may arise 1 cm. or
from the internal iliac about 0.5 cm. from its ori more dorsal to the bladder.
gin, but it may arise from the terminal portion of The caudal vesical artery (a. vesicalis caudalis)
the aorta. In the fetus it carried blood to the pla is a branched vessel which contacts the bladder
centa and was the main artery of the pelvis. In at its neck and ramifies over its caudolateral sur
about half of the specimens it sends one or more face. Its medial branches anastomose with their
twigs to the cranial end of the bladder as the cra fellows both dorsally and ventrally on the organ.
nial vesical arteries (aa. vesicales craniales), Cranially, the caudal vesical artery may anasto
which are its only collateral branches. In the mose with the cranial vesical from the umbilical
adult dog the lumen of the artery is obliterated artery. If the cranial vesical is lacking, the paired
completely, or only distal to the origin of these caudal vesical artery supplies the whole organ. It
arteries, forming the lateral u m bilical ligament is accompanied by a satellite vein and lymph
(ligamentum umbilicale laterale). This term re vessels. Sometimes two caudal vesical arteries
fers to the nonpatent remnant of the fetal artery are present on each side.
distal to the origin of the most distal cranial In the male the small artery o f the ductus def
vesical artery when more than one is present. erens (a. ductus deferentis) (Fig. 4-71), which is
Ellenberger and Baum (1943) and Sisson and homologous with the uterine artery of the fe
Grossman (1953) call the urogenital artery, male, is regarded as the terminal branch of the
which supplies much of the pelvic parts of the cranial part of the urogenital, although the larger
urinary and genital systems, the umbilical artery. caudal vesical continues the direction of the par
Since this artery never formed a part of the ent vessel to the bladder. Upon reaching the
vascular path from the fetus to the placenta the ductus deferens at the point where it enters the
term is a misnomer; the artery is not homologous dorsum of the prostate, it runs proximally to
with the comparable vessel of man. ward the testis and anastomoses in the epididy
mis with the testicular artery. It is accompanied
Visceral Branch of the Internal Iliac Artery by a satellite vein, lymphatics, and a nerve com
ponent from the pelvic plexus.
The visceral branch (ramus visceralis) (Figs. The uterine artery (a. uterina) (Figs. 4-72 and
4-71, 4-72) is the smaller, more ventral branch 15-22) is extremely variable in size, depending
of the internal iliac artery. It lies in contact ven- on the stage of genital activity of the uterus. It
trolaterally with its satellite vein as it runs cau arises from the cranial branch of the urogenital,
dally on the terminal tendon of the psoas minor and bends cranially to become related to the lat
muscle. Upon reaching the origin of the levator eral surface of the caudal part of the cervix by
ani, it divides into the urogenital and internal entering the broad ligament. It diverges from
pudendal arteries. the body of the uterus as it runs cranially, so that
The urogenital artery (a. urogenitalis) is about shortly after the uterine horns arise it lies 1 to 4
1.5 mm. in diameter and 2 cm. long. It makes a cm. from the uterine horn in the broad ligament.
sweeping, caudally facing arch as it runs toward It maintains this distance from the uterine horn
A b d o m in a l A o r t a 379
and sends branches to it, which anastomose with The middle rectal artery (a. rectalis media)
uterine branches of the ovarian artery. The uter leaves the dorsal side of the vaginal artery in the
ine artery is also known as the caudal uterine ar female or the prostatic artery in the male. It
tery (EUenberger and Baum 1943) or cervico- arborizes on the wall of the rectum and anasto
uterine artery (Barone and Pavaux 1962). moses with the cranial and caudal rectal arteries.
Smaller arterial branches leave the dorsal surface The internal pudendal artery (a. pudenda
of the uterine artery and supply the broad liga interna) (Figs. 4-71, 4-72) is a direct continua
ment and round ligament of the uterus. Near the tion caudally of the visceral branch of the inter
origin of the uterine artery a branch arises which nal iliac after the urogenital arises. It is about 1
supplies the cranial portion of the vagina. The mm. in diameter and 7 cm. long as it runs along
uterine artery has a satellite vein and accom the dorsal border of the ischiatic spine where it
panying autonomic nerve plexus and lymphatics. lies lateral to the coccygeal muscle and medial
The prostatic artery (a. prostatica) (Fig. 4-71), to the gluteal and piriform muscles. Occasionally
homologous with the vaginal artery of the fe the caudal gluteal and lateral coccygeal arteries
male, is usually less than 1 mm. in diameter and arise from the internal pudendal. The internal
about 1 cm. long as it crosses the rectum. It lies pudendal is free of branches until it reaches the
in the pelvic fascia with the pelvic plexus of ischiorectal fossa, where it gives off a trunk for
nerves and its satellite vein lying caudal to it. the caudal rectal and the perineal arteries
When the prostate is hypertrophied to the extent (Thamm 1941). After this trunk arises, opposite
that it is an abdominal organ the prostatic artery the anal sac or caudal to the caudal border of the
and related structures are carried forward with levator ani, the internal pudendal continues as
the gland. Two or more branches pass over the the artery of the penis or clitoris.
lateral surface of the prostate and others enter The caudal rectal artery (a. rectalis caudalis)
the parenchyma of the organ. Some of these ex may arise from the internal pudendal cranial to
tend through the gland to supply the prostatic the perineal artery, but the two usually arise
portion of the urethra and the colliculus semina- from a common trunk. The caudal rectal runs
lis. The urethral branches (rami urethrales) are medially and divides into dorsal and ventral
the several branches which arise from the caudal branches as it reaches the anal canal just cranio-
extension of the urogenital after the prostatic ventral to the anal sac. In its course medially it
artery arises. The first branch sends twigs to the gives off a lateral branch to this sac. Its dorsal
caudal part of the prostate, as well as to the part lies under the external anal sphincter and
urethra. The remaining branches arborize on the anastomoses with the opposite artery mid-
caudal part of the membranous urethra and its dorsally. It sends branches to and through the
surrounding urethral muscle and, bending dis external anal sphincter to supply the circumanal
tally around the ischial arch, enter the root of the glands, which are usually hypertrophied in old
penis where they anastomose with the artery of male dogs. It anastomoses in a plexiform manner
the bulb. The caudal portion of the urogenital with the cranial rectal artery on the dorsal part
and its branches are accompanied by satellite of the anal canal. The ventral branch anasto
veins, lymphatics, and autonomic nerves. moses with its fellow so that an arterial circle is
The vaginal artery (a. vaginalis) (Fig. 4-72) of formed around the anal opening. This is fre
the female represents the whole caudal portion quently plexiform in nature. In some specimens
of the urogenital of the male. It arises from the the caudal rectal artery arises as an unpaired
urogenital, lateral to the rectum, and runs caudo- vessel from the middle coccygeal artery opposite
ventrally on the rectum to reach the vagina at the sixth coccygeal vertebra.
the junction of its middle and uterine thirds. In The perineal artery (a. perinealis) (Figs. 4-71,
multiparous specimens three to five helicine 4-72, 4-73) usually arises from the internal pu
branches ramify on the distal two-thirds of its dendal in common with the caudal rectal. It
wall and anastomose ventrally and dorsally with courses superficially and supplies the skin and
the opposite branches. Some of these continue fat at the pelvic outlet. Ventrally its deeper
beyond the vagina and ramify on the relatively branches may aid in forming the arterial ring
short urethra as the urethral branches (rami around the anus, after which it leaves the pelvic
urethrales). Cranially the vaginal artery anasto outlet and runs distally to supply the caudal part
moses with the vaginal branch of the uterine of the scrotum as the cau dal scrotal branch
artery, and caudally it runs out on the vestibule (ramus scrotalis caudalis). In the female it sends
as the vestibular branches (rami vestibulares). a long branched vessel distally, which ends in
(Text continued on page 383.)
380 Chapter 4. The H e a rt and A rte rie s
Cran. g l u t e a l
I lio lu m b o r
S a c ra l s p in a l b r\
i P a r i e t a l br. o f int. i l i a c
V e ntra l coccygeal i
j /M e d ia n s a c r a l
C audal g lu te a l ( (
,Rt. int. i l i a c
M e d ia n c o cc yg e a l K
i ,Rt. ext. i l i a c
Vent. lat. c a c c y q e a l \ ' '
v \ ' L u m b a r aa.
D orsal la t.c o c c y g e a l'
.Deep c i r c u m f l e x i l ia c
Super f. l a t c o c c y g e a l '
^ A o rta
Int. p u d e n d a l
__ Caudal m e s e n te ric
C audal r e c t a l ^
------U m b i I i c a l
A .of p e n is ^ ^
" ~ - U re te r
U re th ra l-
" - L e ft c o lic
P e r i n e a l ---- >
v C r a n ia l r e c t a l
A .o f b u l b -
*** v V is c e ra l br. o f int. i l i a c
Deep a. of p e n is -
vU ro g e n i t a l
U re th ra ' iV) ' \
1 i ' ' C ra n ia l v e s ic a l
R t . d o r s a l a .of p e n i s ' t ,
* ' \
/ i I 1 \ ' Ductus d eferens
M id d le r e c t a l1 / » \ \
i 'C a ud al ve s ic a l
U r e t h r a l br.' i
i Caudal u r e t e r a l
p ro s ta te 1
A.of d uctus d e fe re n s
P ro sta tic i
Cron, g lu t e a
1 1 l o lu m b a r
S a c r a l s p in a I far,
i iP a n e ta l b r o f in t ilia c
V e n t c a c c ijc je a l
/ ! Median s a c r a l
C a u d g lu te o ly ( /
/ mR. in t . 11i a c
M e d ia n c o ccyg e a l v ^ / /
' ' /O R. ext. i l i a c
Vent I at. coccygeal s '
L u m b a r aa.
' \ ' i
Dorsal lat coccygeal N \ v ,
Deep a r c u m fle x i l i a c
S u p e r flo t coccygeal
_ A o r ta
Int. p u d e n d a l^
__ - C a u d m e s e n t e r i c
Caudal r e c t a l -
— U m bi h e a l
P e r in e a l - U reter
L e ft col ic
A.of v e s t ib u la r b ulb C ra n ia l r e c t a l
v V isc e ra l b r o f in t i l i a c
A of c h ta n s
m e h orn
tal
U te rin e
V a g in a s C r a n ia l v e s i c a l
U reth ra 1 s Caudal u rete ra l
M id d le r e c t a l1 Caudal v e s ic a l
U r e t h r a l b r a f u r o g e n t to I i 1C e r v i x
V a g i n a l a. ''Vaginal br. of u r o g e n i t a l
Fig. 4 -7 2 . Arteries o f the fem ale pelvis, right lateral aspect.
382 Chapter 4. The H e a rt and A rte rie s
L a t coccygeal a.- -
M caccyqeus^
M. l e v a t o r a n i
Cutaneous b i c a u d g lu te al a*
Caud. re c ta l o.*
Cutaneous br. -
Sacrotuberous t , g n
C au d.glutea l a.
Int. pudendal a
P e r in e a l a.
D o r s a l a. o f p e n is -
M. ischiocavernosus —
Caud. sc r a t a l a -
M. bulbocavernosus
the vulva as the caudal labial branch (ramus rise to the iliolumbar and cranial gluteal arteries.
labialis caudalis). The internal pudendal artery The iliolumbar artery (a. iliolumbalis) (Figs.
is accompanied by the pudendal nerve, and its 4-74, 4-75), less than 1 mm. in diameter, leaves
branches have satellite veins and lymphatics. the pelvic cavity between the iliopsoas muscle
The artery o f the penis (a. penis) (Fig. 4-71) and the base of the sacrum. It arises from the
terminates the internal pudendal artery in the initial part of the parietal branch or from the in
male (Christensen 1954). It begins where the ternal iliac directly. A small twig leaves its cau
perineal artery leaves the internal pudendal or, dal side to run with the lumbosacral plexus of
if this vessel should arise by a common trunk nerves. As it crosses the border of the ilium it
with the caudal rectal, then the origin of the sends the nutrient artery o f the ilium (a. nutricia
trunk marks the beginning of the artery of the ilia) caudoventrally to enter the nutrient fora
penis. It is about 2 cm. long and devoid of col men. The main part of the vessel crosses the
lateral branches. It terminates by bifurcating or, cranioventral border of the wing of the ilium at
more commonly, giving off in succession the the caudal ventral iliac spine and plunges into
artery of the bulb of the penis, deep artery of the the cranial muscles of the rump and thigh.
penis, and the dorsal artery of the penis. Branches are supplied to the iliopsoas and the
The artery o f the bulb o f the penis (a. bulbi quadratus lumborum as they cross the artery.
penis) is a short artery which divides initially The principal branches go to the proximal parts
into two or three branches, which then redivide of the middle gluteal, and other branches supply
within the urethral bulb. The urethral bulb, the abdominal wall. The iliolumbar anastomoses
corpus cavernosum urethrae, penile urethra, and with the cranial gluteal, caudally, and with the
pars longa glandis receive blood through the superficial circumflex iliac, cranially. It is accom
artery of the bulb. A homologous vessel, the panied by a small satellite vein.
artery o f the vestibular bulb, is present in the fe The cranial gluteal artery (a. glutea cranialis)
male. (Fig. 4-74), about 1.5 mm. in diameter, runs
The deep artery o f the penis (a. profunda dorsocaudally across the lateral surface of the
penis) divides into two to five branches and ischiatic nerve and enters the overlying middle
passes through the tunica albuginea to supply gluteal after passing over the cranial part of the
the corpus cavernosum penis and os penis. greater ischiatic incisure near the caudal dorsal
The dorsal artery o f the penis (a. dorsalis iliac spine. Upon reaching the deep surface of
penis) runs on the dorsal surface of the penis dis the middle gluteal it divides. A small branch ex
tally to the bulbus glandis. It anastomoses with tends dorsally between the middle gluteal and
the deep artery and the artery of the bulb. Ac the piriformis, both of which it supplies, and be
cording to Christensen (1954), the artery b i comes superficial at the sacrococcygeal junction.
furcates into deep, superficial, and preputial The main part of the vessel, with the cranial
branches. For the finer distribution of the arter gluteal nerve and a satellite vein, runs cranio-
ies and veins of the penis, and the mechanism of laterally between the deep and middle gluteals.
erection, refer to Chapter 15, Urogenital System. It is the main supply to the middle gluteal and
The artery o f the clitoris (a. clitoridis) (Fig. 4 - anastomoses with the iliolumbar artery, which
72) is homologous with the deep artery of the enters the muscle cranially. It also anastomoses
penis. It is a minute vessel which supplies the fat, with the cranial femoral, which enters the mid
erectile tissue, and integument which compose dle gluteal after crossing the cranial surface of
the clitoris. It is the terminal part of the visceral the hip joint.
branch of the internal pudendal artery. From the ventral surface of the parietal branch
or from the caudal gluteal artery, a long cutane
Parietal Branch of the Internal Iliac Artery ous branch arises which leaves the pelvic outlet
at the ischiorectal fossa and courses toward the
The parietal branch of the internal iliac artery root of the penis in the fat of the pelvic outlet. It
arises as the larger, more dorsal terminal branch, supplies the fat and skin which covers the upper
ventral to the base of the sacrum, caudomedial part of the caudal surface of the thigh. It may
to the iliopsoas muscle. It is about 2 mm. in di anastomose with the perineal artery or may re
ameter and runs caudally toward the ischiatic place it functionally. The parietal branch of the
spine, parallel to the internal pudendal artery. internal iliac terminates at a transverse plane
Before terminating as the large caudal gluteal through the second coccygeal vertebra as the
and small superficial lateral coccygeal, it gives superficial lateral coccygeal and the caudal glu-
384 Chapter 4. The H e a rt and A r te r ie s
M. Sphincter a n i ext.-
M .c o c c y g e u s -----
Cran. g lu te al a^w.
Mm. o b tu ra to r int. f
g e m e lli ^ Ilio lu m b a r a.rv.
Sacrotuberous l i g -
I Br. of superf.
Coud. g l u t e a l a.* v .- ■ circum fl. ilia c a.vv.
I s c h i a t i c n- ~ m ' \
Caud. cutaneous
s u r a l n.
F it. 4-74. Artenes and veins of the gluteal region, lateral aspect
A b d o m in a l Ao r t a 385
Deep circumflex, i l i a c
/ A o r ta
E xt ilia c
- U m b ilic a l
Int. i l i a c
7th lu m b a r
- - Femoral
-Deep fe m oral
- - P a rie ta l b r , int. i l i a c
M e d ia n s a c ra l
Ilio lu m b a r
- 'V is c e ra l br.: int. i l i a c
Sacral spinal branches
" U r o g e n it a l
'•Int. p u d e n d a l
C r a n ia l g l u t e a l
SACRUM
Caudal g l u t e a l
Perin e al
Median c o c c y g e a l " Superf. lat. c occygeal
V e n tr a l c o c c y g e a l ■"
S e g m e n ta l " Median c o c c y g e a l
Vent. lat. c o c c y g e a l Co 9
H E M A L ARCH M u s c u la r br
Segmental
-Vent. lat. coccygeal
Ca 8 - '
Vent coccygeal
V e n tra l aspect
j TRANSVERSE PROC.
Median coccygeal
V e rte b ra l canal I M uscular br.
Dorsal l a t e r a l coccygeal
\
Superf. lat. coccygeal- «r-/ J HEM AL PROCESS
* V\ ®
Co 8
Ventral lat. coccygeal Kri
'V e n t r a l coccygeal
M e d ia n coccygeal
C audal a s p e c t L a te ra l a s p e c t
teal artery. The parietal branch of the internal dinosus, although an extensive anastomosis with
iliac lies medial to the ischiatic nerve and dorsal the caudal femoral and deep femoral arteries
to its satellite vein. exists within these muscles.
The superficial lateral coccygeal artery (a.
coccygea lateralis superficialis) (Fig. 4-75) MEDIAN SACRAL ARTERY
leaves the parietal branch of the internal iliac
about 4 cm. caudal to the origin of the cranial The median sacral artery (a. sacralis mediana)
gluteal. It leaves the pelvis by passing caudo- (Fig. 4-75) is the direct continuation of the aorta
dorsally between the tail and the superficial glu caudally, after the internal iliac arteries arise. It
teal muscle. A cutaneous branch arises at this usually arises opposite the body of the seventh
level and, curving around the sacrotuberous lig lumbar vertebra as an unpaired median vessel
ament and superficial gluteal muscle, enters the which is slightly less than 2 mm. in diameter. It
superficial gluteal fascia. It supplies the skin and crosses under the promontory of the sacrum
fascia of the rump as far forward as the crest of with its dextrally lying satellite vein and enters
the ilium. Branches leave both the dorsal and the the fat-filled furrow between the right and left
ventral surface of the vessel at irregular intervals medial ventral sacrococcygeal muscles. In this
and supply the skin and adjacent fascia of the region it usually gives off two pairs of sacral
tail. At about the sixth coccygeal vertebra it in spinal branches (rami spinales), which enter the
clines dorsally and, lying on the deep coccygeal ventral sacral foramina. One or both of these
fascia dorsal to the transverse processes, runs to may arise from the parietal branch of the internal
the tip of the tail. In the distal two-thirds of the iliac or from the adjacent cranial gluteal artery.
tail it sends many branches to the skin and mus Arising from these spinal branches are twigs to
cles along its dorsolateral aspect. Other branches the adjacent ventral sacrococcygeal muscles.
run across the discs and anastomose with the Variations of the arteries to the tail are numer
median coccygeal artery; shorter branches run ous. Typically, there are seven longitudinal
deeply to anastomose with the deep lateral coc arterial trunks in the proximal third of the tail as
cygeal arteries. Arteriovenous anastomoses exist follows: 1 (impaired) median coccygeal artery,
in the caudal third of the tail. The large satellite 2 (paired) superficial lateral coccygeal arteries, 2
vein lies ventral to the superficial lateral coccyg (paired) dorsal lateral coccygeal arteries, and
eal artery. 2 (paired) ventral lateral coccygeal arteries.
The caudal gluteal artery (a. glutea caudalis) The median coccygeal artery (a. coccygea
(Figs. 4 -7 3 , 4-74, 4-75) is the continuation of mediana) is a direct continuation caudally of the
the parietal branch of the internal iliac distal to median sacral artery at the first coccygeal verte
the origin of the small superficial lateral coccyg bra. It runs mid-ventrally on the coccygeal verte
eal artery. It passes toward the tuber ischii in re brae, where it lies between the right and left
lation to the sacrotuberous ligament caudolater- medial ventral sacrococcygeal muscles. It passes
ally, the ischiatic nerve cranioventrally, and its through the fourth, fifth, and sixth (if present)
satellite vein medially. It gives a muscular branch hemal arches, and then between the successive
to the superficial gluteal, and sends twigs to the hemal processes. Throughout most of its
piriformis, obturator internus, and gluteus me- course, segmental arteries which run caudo-
dius. It anastomoses with the cranial gluteal laterally arise opposite the bodies of the verte
artery in the gluteus medius. The main caudal brae. They pass ventral to the transverse proc
gluteal artery passes over the ischiatic spine with esses of the corresponding vertebra and give fine
the ischiatic nerve and satellite vein, and divides twigs to the adjacent structures. At irregular
into several branches as it enters the biceps fem intervals, usually the length of two or three seg
oris. One branch enters the proximal end of ments, there are ventral branches which supply
the semitendinosus, and another enters the semi the skin. Other branches successively leave the
membranosus. The satellite artery o f the ischia median coccygeal from alternate sides, starting
tic nerve (a. comitans n. ischiadicus), a branch of at the eighth coccygeal vertebra, each anasto
the caudal gluteal, joins the nerve caudal to the mosing with the superficial lateral coccygeal
trochanter major and runs distally with it. The artery of the same side. Only the two superficial
quadratus femoris, obturator internus, and ge- lateral coccygeal arteries and the median coc
melli may also receive branches from the proxi cygeal artery reach the tip of the tail, where they
mal part of the artery. The caudal gluteal is the anastomose. They lose their segmental character
main supply to the biceps femoris and semiten in the last few segments and anastomose with
M e d ia n Sacral Artery 387
each other in a plexiform manner. The right and The only recognizable veins from the tail are the
left ventral coccygeal arteries may be the first right and left superficial lateral coccygeal veins.
branches to leave the median coccygeal.
The paired ventral coccygeal artery (a. coc
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Kennedy, H. N., and A. W. Smith. 1930. An abnormal celiac nal vessels and their relation to the type of posterior vena
artery in the dog. Vet. Rec. (London) 10: 751. cava in the dog (Canis familiaris). Am. J. Anat. 99;1-15.
Krediet, P. 1962. Anomalies of the arterial trunks in the thorax Sisson, S., and J. D. Grossman. 1953. Anatomy of the Do
and their relation to normal development. Thesis, mestic Animals. 4th Ed. Philadelphia, W. B. Saunders
Utrecht, pp. 1-108. Company.
Latimer, H. B. 1961. Weights of the ventricular walls of the Speed, J. G. 1943. The thoraco-acromial artery of the dog. Vet.
heart in the adult dog. Univ. Kansas Science Bull. XLII: J. 99: 163-165.
3-11. Tandler, J. 1899. Zur vergleichenden Anatomie der Kopfarte-
Ljubomudrov, A. P. 1939. The blood supply of the suprarenal rien bei den Mammalia. Denkschr. Akad. Wiss. Wien
glands in the dog. Arkhiv Anat. Grist, i Embryol. 2 0 :220- Math.-naturwiss. Kl. 67: 677-784.
224 (Eng. Summary 381-382). Thamm, H. 1941. Die arterielle Blutversorgung des Magen-
Marthen, G. 1939. Uber die Arterien der Korperwand des darmkanals, seiner Anhangsdriisen (Leber, Pankreas) und
Hundes. Morph. Jahrb. 84: 187-219. der Milz beim Hunde. Morph. Jahrb. 85; 417-446.
Meek, W. J., M. Keenan, and H. J. Theisen. 1929. The auricu Thomas, C. E. 1957. The muscular architecture of the ven
lar blood supply in the dog; I. General auricular supply tricles of hog and dog hearts. Am. J. Anat. 101: 17-58.
with special reference to the sinoauricular node. Am. Trautmann, A., and J. Fiebiger. 1952. Fundamentals of the
Heart J. 4:591-599. Histology of Domestic Animals, translated and revised
Miller, M. E. 1958. Guide to the Dissection of the Dog. 3rd from 8th and 9th German editions, 1949, by R. E. Habel
Ed. Ithaca, N.Y. Pub. by author. and E. L. Biberstein. Ithaca, N. Y., Comstock Publishing
Moore, R. A. 1930. The coronary arteries of the dog. Am. Assoc., Cornell Univ. Press.
Heart J. 5:743-749. Truex, R. C., and L. J. Warshaw. 1942. The incidence and size
Nickel, R., A. Schummer, and E. Seiferle. 1960. Lehrbuch der of the moderator band in man and mammals. Anat. Rec.
Anatomie der Haustiere. Band II. Eingeweide. Berlin, 82:361-372.
Paul Parey. Whisnant, J. P., C. H. Millikan, K. G. Wakim, and G. P. Sayre.
Noer, R. 1943. The blood vessels of the jejunum and ileum; A 1956. Collateral circulation of the brain of the dog follow
comparative study of man and certain laboratory animals. ing bilateral ligation of the carotid and vertebral arteries.
Am. J. Anat. 73: 293-334. Am. J. Physiol. 186: 275-277.
CHAPTER 5
TH E VENOUS SYSTEM
Veins and lymphatic vessels follow the same The precava, or cranial vena cava (v. cava
general course as do the arteries. The accom cranialis) (Figs. 5-1, 5-2), is an unpaired vessel,
panying veins are known as satellite veins, or 1.5 to 2 cm. in diameter, and 8 to 12 cm. long. It
venae comitantes, and often take the same name lies ventral to the trachea, and is in contact with
as the artery they accompany. Although the the esophagus on the left side. It is also in con
smaller satellite veins are frequently double, the tact with the thymus when this gland is fully de
larger veins are single, as are most of the deep veloped. It runs through the precardial medias
veins. All systemic veins have thin walls, and tinum and is the most ventral of the several
most have large lumina in comparison with the structures which course through the thoracic in
arteries. The pressure in the veins is low, and the let. It is formed, at a level cranial to the thoracic
blood flows much more slowly in them than in inlet, by the convergence of the right and left
the arteries. Since there is generally no pulse in brachiocephalic veins. These form an angle,
the venous system, the movement of blood de open cranially, of about 90 degrees, so that each
pends primarily on pressure relations in the tho vein enters the precava at an angle of approxi
rax and on muscular activity. The contraction of mately 45 degrees with the median plane. The
muscles results in compression of the veins, thus precava empties into the cranial part of the right
propelling the contained blood toward the heart. atrium. Stoland and Latimer (1947) describe a
Negative pressure in the thorax during inspira dog with a persistent left cranial vena cava. The
tion and the presence in most veins of semilunar vein was about equal in size to the normal right
valves which prevent back flow augment this ef precava, and was continued caudally in the left
fect of the skeletal and visceral muscles. Veins part of the coronary groove to the coronary si
farthest from the heart contain the most valves. nus. In reviewing the incidence of anomalous
The venous passages in the dura of the central precavae they state that a total of 207 cases have
nervous system are known as sinuses. In the ex been seen, but only three of these have been de
tremities, the veins may be divided into superfi scribed. The precava receives the single azygos
cial and deep sets. The veins of the superficial vein, the paired costocervical-vertebral trunks,
set are large and are clinically important be and the single, or paired, internal thoracic vein.
cause of the frequent necessity of making veni The azygos vein (vena azygos), with its tribu
punctures. In this location they act in cooling taries, is described after the description of the
the blood as they communicate not only with other tributaries of the precava.
the deep veins but also with extensive subcu The costocervical-vertebral venous trunk
taneous, interconnecting venous plexuses. When (v. truncus costocervicalis-vertebralis) (Fig. 5-1)
the animal is cooled, these plexuses and the is formed medial to the proximal end of the first
larger superficial veins contract so that most of rib. The left venous trunk runs lateral to the left
the blood from the extremities must be returned subclavian artery and empties into the dorsolat
to the heart via the deep veins; this prevents eral surface of the cranial part of the precava. It
heat loss. When the animal is warmed, and dur may terminate in the left brachiocephalic vein.
ing work, in short-haired specimens the superfi The right venous trunk, about 1 cm. caudal to
cial veins and their connecting plexuses dilate the left at its termination, crosses the lateral sur
and are plainly visible beneath the skin; this dila face of the trachea and enters the precava ven
tation provides a means of heat dissipation. tral to the brachiocephalic artery and vagus
389
390 Chapter 5. The V e n o u s S y s te m
nerve. The right costocervical-vertebral trunk is also emptied into the supreme intercostal vein
about 3 cm. long; that of the left side is about 4 of the same side.
cm. Both vessels are approximately 5 mm. in di The internal thoracic vein (v. thoracica in
ameter. terna) (Fig. 5-2) is unpaired at its termination in
The vertebral vein (v. vertebralis) begins by the middle of the ventral surface of the precava
the confluence of the vein of the hypoglossal in about half of all specimens. In such instances
canal, if present, and the sigmoid sinus in the it usually ranges from 1 to 4 cm. in length. In the
petrobasilar fissure. The small internal jugular specimens in which it is paired the right vein
vein also arises wholly or partly at this conflu usually enters the precava, whereas the left en
ence. The vertebral vein is about 1.5 mm. in di ters the left brachiocephalic vein. The peripheral
ameter as it runs caudally across the ventrolat part of the internal thoracic vein and its branches
eral part of the atlanto-occipital joint, then are satellites of the comparable parts of the in
under the wing of the atlas to the transverse fo ternal thoracic artery.
ramen. It receives the first intervertebral vein The brachiocephalic vein (v. brachiocephal-
which leaves the vertebral sinus, passes through ica) (Fig. 5-2) merges with its fellow of the op
the intervertebral foramen of the atlas and then posite side anterior to the thoracic inlet to form
through the alar notch, and joins the vertebral the precava. Each is about 2 cm. long and 1 cm.
vein in the atlantal fossa. Worthman (1956) and in diameter, and is formed by the joining of the
Drager (1937) call the initial portion of the ver caudally coursing external jugular and the medi
tebral, the occipital vein, although only a seg ally coursing axillary vein. (Unlike the compa
ment of it parallels the occipital artery. The dog, rable artery, no part of the venous channel com
however, may be regarded as not having a true ing from the thoracic limb normally lies within
occipital vein (v. occipitalis), because the dorsal the thorax. There is, therefore, no basis for nam
nuchal area supplied by the occipital artery is ing a portion of the channel for venous return
drained by the occipital emissary vein, and the from the thoracic limb the subclavian vein.) The
ventral portion supplied by the occipital artery merging brachiocephalic veins lie ventral to the
is drained by the vertebral vein. trachea and esophagus as the most ventral struc
After passing through the transverse foramen tures in this region. In addition to the external
of the atlas from the atlantal fossa, the vertebral jugular and axillary veins, the thyroidea ima and,
vein receives a small muscular tributary, and a occasionally, the caudal thyroid, and the internal
slightly larger communication from the caudal jugular veins enter the brachiocephalic.
portion of the first intervertebral vein and the The v. thyroidea im a is an unpaired vein,
large second intervertebral vein. The portion of about 1 mm. in diameter, which arises primarily
the vertebral vein which passes through the from the deep surfaces of the sternothyrohyoid
transverse foramen of the axis is its smallest part. muscles, but on one or both sides its most cra
There is usually a small anastomotic vein be nial tributary may arise in the thyroid lobe or
tween the second and third intervertebral veins. lobes. It terminates usually in the cranial angle
Most of the blood is conveyed caudally by way formed by the merging brachiocephalic veins.
of the large vertebral venous sinuses. The re The internal jugular vein (v. jugularis in
maining portion of the vertebral vein and its terna), about 1 mm. in diameter, is formed in the
tributaries are satellites of the comparable por petrobasilar fissure by the confluence of the ver
tions of the vertebral artery and its branches. tebral vein, sigmoid sinus, and, occasionally, the
The costocervical vein (v. costocervicalis) and vein of the hypoglossal canal. It regularly re
its three main tributaries, the supreme intercos ceives a relatively large cranial communicating
tal, transverse colli, and deep cervical veins are vessel from the posterior end of the ventral pe
satellites of the comparable arteries. The last, or trosal sinus. Its initial portion may be double.
eighth, cervical intervertebral vein regularly bi The internal jugular lies at first in association
furcates into a cranial communicating vein with the internal carotid artery and then in the
which joins the vertebral vein, and a caudal sheath of the common carotid. In the vicinity of
communicating vein which empties into the the larynx it receives an anastomotic twig from
deep cervical vein. According to Worthman the laryngeal or pharyngeal tributary of the lin
(1956), the supreme intercostal vein receives the gual. Caudal to the larynx it receives the cranial
second and third thoracic intervertebral veins on thyroid vein (v. thyroidea cranialis) from the
each side. In almost half of his specimens the cranial pole of the thyroid lobe. Opposite this
fourth thoracic intervertebral vein on the left tributary there is occasionally a second anasto-
P recava, or C r a n ia l V en a C ava 391
R. subclai/ian a. L, o m o c e r v ic a l a.
R, o m o c e rv ic a l v, - A x illa r y a.
ragj l/e n t e b r a I a,
Dist. c o m m u n ic a t in g ^
b r o f cepha l i e v.
I n t e r n a l j u y u l a r v.--------- In te r n a l th o ra c ic a,
B r a c h io c e p h a lic a.--------- T h o ra c ic d u c t
A x i l l a r y v.
A x i l l a r y v.--------
L, b r a c h i o c e p h a li c v,
V. t h y r o i d e a i m a ~ — S u b c la v ia n a.
In te rn a l th o ra c ic v . - ------ -----L. c o s t o c e r v i c a l - v e r t e b r a l
R. c o s t o c e r v i c o l - v e r t e b r a l t runk
tru n k A o r tic a rch
-----P u l m o n a r y a,
Thoracic o o rta
F ig . 5-1. Diagram of the heart and great vessels, ventral aspect. (Modified from Miller 1958.)
392 Chapter 5. T h e V e n o u s S y s te m
Int. j u g u l a r
Ext ju g u la r- -
Prox. c o m m u n ic a t in g - -
b r o f c e p h a li c
O m o ce rvica l - -
-S u b s c a p u la r
Dist. c o m m u n ic a tin g
b r o f c e p h a lic ~
■Caud. t h y r o i d
T h y r o i d e a im a - -
-L. b ra c h io c e p h a li
A x illa ry
C o s to c e rv ic a t-
-v e rte b ra l tru n k
P recava- - Cephal ic
Int. t h o r a c i c -
-B ra c h ia l
M ed ia n c u b i t a l
M e d ia n - -
- C e p h a l ic
motic connection with the external jugular vein. lacks valves. It receives the small frontal vein (v.
A twig from the medial retropharyngeal lymph frontalis), which is a tributary medial to the
node is also received here. Frequently, in the orbit on the surface of the frontal bone. The
caudal half of the neck, it receives the small cau angularis oculi vein may anastomose with the
dal thyroid vein (v. thyroidea caudalis), which superficial temporal vein via the frontal vein. It
comes from the caudal pole of the thyroid lobe. disappears from the surface of the face by curv
On either or both sides, this vein may terminate ing around the dorsomedial border of the orbital
in the brachiocephalic vein. The internal jugular margin, to become the ophthalmic vein. It usu
vein usually terminates in the caudal portion of ally receives an emissary vein from the superfi
the external jugular vein; rarely it terminates in cial surfaces of the frontal and nasal bones.
the brachiocephalic vein. The ophthalm ic vein (v. ophthalmica) runs
The external jugular vein (v. jugularis ex about 2 cm. posteriorly into the orbit and forms
terna) (Fig. 5-3), unlike that of man, is the main the orbital plexus (plexus orbitalis venosus). Like
channel for return of venous blood from the the vein from which it originates, the plexus lies
head. It begins by the union of the internal and between the periorbita and the medial osseous
external maxillary veins, caudal to the mandibu wall of the orbital fossa. It is 8 mm. at its great
lar gland or at a transverse plane through the cri est width and approximately 4 cm. long. It ex
coid cartilage and the axis. It is about 1 cm. in tends to the orbital fissure, and therefore lies in
diameter and 12 cm. long. In the adult it con the posterior two-thirds of the orbital fossa. The
tains a few nonfunctional valves which are ir plexus becomes consolidated at the orbital fis
regular in their spacing. As the external jugular sure and, after traversing it, joins the cavernous
runs caudally in the superficial fascia it crosses sinus. Dorsally, a small branch runs through the
the lateral surface of the brachiocephalic muscle optic canal to join its fellow and the dorsal petro
obliquely. It lies directly under the skin and is sal sinus of its side; ventrally the plexus is con
commonly used for venipuncture in dogs which tinued posteriorly outside the alar canal and is
are too small for the procedure to be feasible in to be regarded as the beginning of the internal
the smaller veins of the extremities. At the cra maxillary vein. A second connection between
nial border of the shoulder, it receives the proxi the ophthalmic system and the internal maxillary
mal and distal communicating branches of the exists here in the form of a vein which runs
cephalic vein which ascend from the brachium. through the alar canal. This vein also communi
About 2 cm. caudal to the termination of the cates with the cavernous sinus by an emissary
proximal communicating branch, the external vein which traverses the round foramen. The
jugular receives the om ocervical vein (v. omo- ophthalmic vein or its intrinsic orbital plexus
cervicalis), but this is variable. At its termination communicates with the internal maxillary vein,
it usually receives the internal jugular vein on its as well as with the reflex vein at the orbital fora
medial side. men. It also communicates with the superficial
The external maxillary vein (v. maxillaris ex temporal vein, which joins the middle of the or
terna) (Fig. 5-4) is about 5 mm. in diameter and bital plexus dorsally. Two emissary veins join the
2 cm. long. It begins by the confluence of the plexus. One of these is the external ethm oidal
lingual and facial veins ventral to the mandib vein (v. ethmoidalis externa,) a satellite of the
ular gland. It may have one or more tributar like named artery, which passes through the
ies from the capsule of the mandibular gland. ethmoidal foramen. The other emissary vein is
It regularly receives the m an dibu lar vein (v. the fron tal diploic vein (v. diploica frontalis),
mandibularis), which leaves the posterior pole from the diploe of the frontal bone, which passes
of the mandibular gland. It contains a valve at through a small foramen in the supraorbital
its termination. process. The latter vein may join the ophthalmic
The facial vein (v. facialis) (Figs. 5-4, 5-5) vein before the plexus is formed.
begins on the dorsolateral surface of the muz The lateral nasal vein (v. nasalis lateralis) is a
zle, covered by the levator nasolabialis. It is satellite of the lateral nasal artery. It is a tribu
formed by the confluence of the smaller dorsal tary which enters the facial vein.
nasal vein (v. nasalis dorsalis), which drains the The infraorbital vein (v. infraorbitalis) is
dorsolateral surface of the nose, and the larger about 1 mm. in diameter and 1 cm. long. It com
angular vein of the eye. municates with the ventral side of the facial
The angular vein o f the ey e (v. angularis vein, which lies dorsal to the infraorbital fora
oculi) is about 3 mm. in diameter and 2 cm. long. men. It has tributaries from the infraorbital
Blood may flow in either direction in it, as it nerve and adjacent musculature. Posteriorly, it
394 Chapter 5. The V e n o u s S y s te m
unites with the sphenopalatine vein to form a of the ocular muscles to enter the reflex vein. It
short common trunk which anastomoses with also receives tributaries from the maxilla as it
the reflex vein in the anterior part of the ptery curves from the deep surface of the masseter be
gopalatine fossa. fore entering the facial vein.
The m alar vein (v. malaris) is a small tributary The ventral labial vein (v. labialis ventralis), a
which arises mainly in the skin of the lower eye satellite of its artery, runs along the ventral
lid and terminates in the dorsal surface of the fa border of attachment of the buccinator muscle.
cial vein. It receives at its termination a vein which arises
The dorsal labial vein (v. labialis dorsalis) runs in the intermandibular space. This vessel runs
posteriorly along the dorsal margin of attach along the margin of insertion of the digastricus,
ment of the buccinator muscle and enters the then over the lateral surface of the mandible. It
facial vein lateral to the anterior end of the zy may terminate in the facial vein. Throughout
gomatic arch. It drains blood from the upper lip the course of the facial vein small twigs from the
and the dorsal part of the cheek. skin and fascia enter it. Ellenberger and Baum
The angular vein o f the m outh (v. angularis (1943) illustrate a small anastomotic vein located
oris) is a small tributary from the commissure of between the facial and the superficial temporal
the lips which enters the facial vein posterior to vein, as well as small twigs coming from the
the commissure. mandibular lymph nodes.
The reflex vein (v. reflexa) (Fig. 5-4) has no The lingual vein (v. lingualis) (Fig. 5-4) is the
companion artery and is significant for its deep ventral tributary which joins the facial to form
course and many anastomoses. It is called the the external maxillary. It begins in the tip of the
deep fa c ia l vein (v. faciei profunda) by Preuss tongue and as it courses posteriorly it is aug
(1954), who states that it is present in all the mented by numerous tributaries from this organ.
domestic mammals except the ox. It is 2 to 4 It lies in areolar tissue in association with the
mm. in diameter at its terminal end, which lies lingual artery and hyoglossal nerve, lateral to
in the fascia anterior to the masseter muscle, the genioglossal and medial to the hypoglossal
about 1.5 cm. ventral to the zygomatic arch. It muscles. About 1 cm. anterior to the body of the
arises in the ventral part of the orbital and the hyoid bone it crosses the dorsal border of the
adjacent pterygopalatine fossa. The main por hyoglossus and comes to lie dorsal to the mylo
tion of the vein arches dorsomedially around the hyoideus. As it runs out from under the posterior
ventral surface of the cone of eye muscles and border of the mylohyoideus it is joined by the v.
anastomoses with the ophthalmic vein near the sublingualis. This vein begins in the anterior
orbital foramen. A second anastomosis may oc part of the lingual frenulum and runs posteriorly
cur with the ophthalmic about 1 cm. from the on the dorsal surface of the mylohyoideus. It lies
orbital margin. An anastomosis with the super direcdy under the thin mucosa between the
ficial temporal vein frequently occurs as a small lingual frenulum and the fimbriated fold which
vein which obliquely crosses the lateral surface lies lateral to the frenulum. It is occasionally
of the zygomatic arch. A small vein unites the used for venipuncture, but this is ill advised as
reflex with the internal maxillary by running the exceedingly loose tissue which surrounds the
across the lateral surfaces of the pterygoid mus vessel allows considerable hemorrhage to occur.
cles. Anteriorly this vein is accompanied by its satel
Tributaries which enter the reflex vein may lite artery and the lingual branch of the trigemi
form a short venous trunk by the union of the nal nerve. It carries blood from the frenulum
sphenopalatine, infraorbital, and occasionally and the closely adjacent sublingual and mandib
the major palatine. The major palatine is small, ular ducts and the polystomatic part of the
if it is present at all. These veins are satellites of sublingual gland.
the comparable arteries. The main venous drain The hyoid venous arch (arcus venosus hy-
age of the hard palate is by a poorly formed ve oideus) (Fig. 5-3) is a constant large, unpaired
nous plexus which is continuous with the much vein, about 3 mm. in diameter and 3 to 4 cm.
more salient venous plexus of the soft palate. long, which lies ventral to the basihyoid bone. It
Therefore the venous drainage of the hard and usually connects right and left lingual veins
soft palate is not chiefly through satellites of the about 1 cm. caudal to the termination of the
arteries supplying them, but by means of the sublingual veins. Petit (1929) illustrates this ves
veins of the palatine plexuses which drain into sel as extending between the two sublingual
the right and left internal maxillary veins. Con veins. It may be double. It receives on each side
stantly, one or two veins leave the ventral part of the mid line the caudally running small v. sub
t e x t continued on page 398.)
P recava, or C r a n ia l V en a C ava 395
P h a r y n g e a l br- S u b m e n ta l
Hyoid venous a r c h - - L in g u a l
- -L a ryn g ea im p a r
F a c ia l - '
C r a n ia l l a r y n g e a l
L in g u a l
~ —Int. m a x i I l a r y
~~Ext. m a x i I la r y
Thy r a id c a r t i I a g e '
R .c r a n ia l t h y r o id ' P a ra th y r o id g l a n d
T h yro id g l a n d
R. c a u d a l t h y r o i d - -
I------ Ext. j u g u l a r
Tra ch e a -
- -E s o p h a g u s
----- L .in t. j u g u l a r
R. i nt. j u g u l a r —
-----C e p h a l i c , p r o x i m a l
com m unic atin g b r
T hyro id e a im a -
----- O m o c e r v ic a l
L .c a u d a l th y ro id
----- C e p h a l i c , d i s t a l
comm unicating b r
M u s c u l a r br. —
- Ax i I l a r y
B ra c h io o e p h a l ic
Int. t h o r a c i c t r u n k —
j - - -C o s t o c e r v ic a I -v e r t e b r a I tru n k s
P re ca va —
Post. deep t e m p o r a l
' .A la r c a n a l
O rb ita l fissure
tA n t d e e p t e m p o r a l
Med. a u r i c u la r O r b i t a l pflexus
! 1 .E x t e th m o id
I i 1
i i 1 O p h th a lm ic
i i i '
s A n a u la ris o c u li
A n t a u ric u la r
M a jo r p a la tin e
I n te rm e d ia te auric. S p h e n o p a la tin e
Lat. a u r i c u l a r ,JHorsal n a s a l
Transverse fa cTai
S u p e rjy te m p o ra l
R e tro g le n o id
Deep a u r i c u l a r -
0 re a t a u ric u la r.
Vein from
p a la tin e p le x u s . ^ '^
^D o rs a l
__ Int. m n x i l l a r i j la b ia l
\ V ^ ' In fra o rb ita l
I n t. j u g u l a r
Ext, j u g u l a r . \ V \ V e n t r a l la b ia
C ra n ia l t h y r o id / ■
Open fn r .in l
R am us c o m m u n ic a n s / / , / { j
y \ ' S u b lin g u a l
From med. re tro p h aryn g ea l In. '/ i
I \ I \ \ '•P a la tin e p le x u s
Ext. m a x i 11 a r y 1 {
F a c ia l
F rom m a n d i b u l o r g la n d 1 S \ \ ' V
Mandi b u la r a lv e o la r
P h a ryn g e a l br.
14 V
\ \S u jM T i£ n t a l
C ra n ia l la ru n q e a l1 ! \
J | H y o id ven o u s a r c h
L in g u a I
/O rb ita l lig a m e n t
O p h t h a lm i c v.
S u p r a o r b ita l n .(V )
I n f r a t r o c h l e a r n. (V)
sAnqularis o c u l i v.
P a lp e b r a l n ( V I I )
Med. p a lp e b ra l lig.
Superf. t e m p o r a l a. -
- - M a l a r a.
Z y g o m a tic o te m p o r a l
- D o r s a l n asa l v.
M r e t r a c t o r angut i ocul
i - L a t n a s a l v.
Z y g o m a tic o fa c i
I n f r a o r b i t a l v.
D ars■ l a b i a l v.
Deep
Fic.. 5-5. Scheme of the vessels and nerves in the region of the eye.
From maj. p a l a t in e
- B r f r o m deep fa c i a l v.
r sphenopalatine
, - O p h t h a l m ic v.
Deep f a c i a l v. - _
_ - O r b ita l p le xus
O r b ita l plexus - _ / /
\ V - - Optic foram en
Vent, la b ia l v.~ _
— O r b it a l fis s u re
P a latin e p l e x u s — /_ / . __
— Hamulus of pterygoid
Facial v.- - J - -|— M a n d ib u lo r-a lv e o la r v:
- A la r canal
- Superf. te m p ora l v.
Retroglenoid v. — —
- E x t c a r o t i d foramen
G re a t a u r i c u l a r v . - -L in g u a l V.
Ext. m a x i l l a r y v. - -J u g u la r foramen
Int. j u g u l a r vj
Foramen magnum
V e r t e b r a l v.
y -A tla s
Ramus communicans ' ^ ''Transverse foram en
V
^ Cran. th yroid v.
E x t j u g u l a r 1/
A x is - - ^ 50‘*
F ig. 5-6. Veins of the head and neck, ventral aspect.
398 Chapter 5. The V e n o u s S y s te m
mentalis, which usually begins as a single vessel (Fig. 5-4) leaves the skull through the retrogle
in the mid line between the fellow mylohyoid noid foramen and more than doubles the size of
muscles. It receives delicate tributaries from the internal maxillary vein by joining it posterior
both the mylohyoid and the geniohyoid muscle. to the temporomandibular joint. The intra
Entering the caudal surface of the arcus hy- cranial formation of this vein is described with
oideus at the mid line is the delicate unpaired v. the veins of the central nervous system. The
laryngea impar. It anastomoses with the delicate retroglenoid vein is known by many morpholo-
v. laryngea cranialis and usually with end tribu gists as the dorsal cerebral vein.
taries of the v. thyroidea ima. The m andibular alveolar vein (v. alveolaris
The cranial laryngeal vein (v. laryngea cra mandibulae) is the satellite of the comparable
nialis) (Fig. 5-3), a satellite of the like named artery. It leaves the mandibular foramen and at
artery, leaves the larynx ventral to the cranial once receives a branch from the musculature
corner of the thyroid cartilage in company with medial to the mandible, fnainly from the m.
the artery and cranial laryngeal nerve. It joins temporalis as the deep temporal vein (v. tem
the lingual vein about 1 cm. from its termina poralis profunda). Entering the internal maxil
tion. It may send a communicating branch to lary about 5 mm. posterior to the entry of the
the internal jugular vein. The pharyn geal vein mandibular alveolar is the masseteric vein (v.
(v. pharyngea) is a variably formed tributary masseterica). This small tributary comes from
which usually arises in a small venous plexus the upper caudal border of the masseter muscle
located between the vagosympathetic nerve and curves medial to the caudal border of the
trunk and the internal carotid artery on the mandible before it terminates.
lateral wall of the pharynx. It usually sends a The superficial temporal vein (v. temporalis
communicating branch to the internal jugular superficialis) (Fig. 5-4) is about 2.5 mm. in diam
vein. It is a tributary of the cranial laryngeal eter as it terminates in the dorsal surface of the
vein which enters it just lateral to the larynx. internal maxillary. The vein crosses ventral to
The internal maxillary vein (v. maxillaris in the base of the ear, under cover of the parotid
terna) (Fig. 5-4) begins ventral to the alar canal gland. The dorsal tributary arises dorsomedial to
by a posterior continuation and later a consoli the orbit by occasionally anastomosing with the
dation of the extension of the orbital plexus. The small frontal vein, a tributary of the v. angularis
formation and anatomy of the venation in this oculi. The ventral tributary takes a dorsal course
location are complicated and variable. Usually a in the caudal part of the orbital fossa. It rims
small vein lies in the alar canal and receives an under the deep temporal fascia, crosses under
emissary vein from the cavernous sinus through the orbital ligament, and continues medial to the
the round foramen. Anteriorly the vein in the zygomatic arch to anastomose with a branch of
alar canal joins the orbital plexus, whereas pos the reflex vein. This anastomotic channel is over
teriorly it joins the internal maxillary vein. Here 1 mm. in diameter. A small, third anastomotic
also the internal maxillary receives an emissary channel between the v. temporalis superficialis
vein from the oval foramen and the v. meningea and the veins of the orbit is formed as follows: a
media, which is a satellite, usually double, of the branch of the v. temporalis superficialis runs
corresponding artery. About 5 mm. caudal to through the anterior portion of the temporal
the oval foramen two more veins join the internal muscle to enter the orbital fossa and anastomoses
maxillary. One of these is small and comes from with the orbital plexus.
the pterygoid canal; the second is larger, passes The superficial temporal vein, after its forma
through the external carotid foramen, and tion by the three anastomotic branches, runs
connects internally at the confluence of the ven under the deep temporal fascia posteriorly. It
tral petrosal and cavernous sinuses. The internal receives numerous tributaries from the temporal
maxillary vein winds laterally, posterior to the muscle in its course toward the base of the ear.
retroglenoid process, and receives the vein of The v. auricularis anterior is a small vein which
the palatine plexus. begins in the interauricular musculature and
The venous palatine plexus (plexus venosus skin and runs transversely, under the preauricu-
palatinus) (Fig. 5-6) is a rather loose network of lar muscles, anterior to the base of the ear. It re
veins in the soft palate. The largest elements of ceives twigs from the skin and auricular muscles
this plexus are 0.5 mm. in diameter. Anteriorly and the base of the pinna itself. It terminates in
the plexus anastomoses with the sphenopalatine the superficial temporal vein. The v. transversa
and reflex veins. fa c e i is a small tributary which arises in the
The retroglenoid vein (v. retroglenoidalis) fascia ventral to the zygomatic arch. It empties
P recava, or C r a n ia l V en a C ava 399
into the superficial temporal about 1 cm. ventral The first two lumbar, the subcostal, and the
to the termination of the much larger anterior dorsal intercostal veins (vv. lumbales et sub
auricular vein. According to Ellenberger and costales et intercostales dorsales), except the first
Baum (1943), the anterior auricular vein termi three intercostals on the right side and the first
nates in the great auricular vein 50 per cent of three or four dorsal intercostals on the left side,
the time. As the superficial temporal grooves the are received by the azygos or the hemiazygos
anterior border of the parotid gland, it receives vein. The (third) fourth and fifth dorsal inter
one or more rami parotidei from it. costal veins anastomose with each other so that
The great auricular vein (v. auricularis mag- a longitudinal venous trunk is formed which
na) (Fig. 5-4) and its branches, the lateral, inter usually terminates in the proximal end of the
mediate, deep and medial auricular veins, are sixth dorsal intercostal vein. This pattern is usu
satellites of the comparable arteries. The margi ally bilaterally symmetrical in its formation, but
nal veins, the m edial and lateral auricular, anas the left venous trunk is longer than the right be
tomose with each other near the tip of the cause it crosses under the body of the fifth tho
pinna on the posterior, or convex, side. The racic vertebra to reach the azygos vein.
intermediate and deep auricular veins are small. The first two lumbar veins are smaller than
Unlike the great auricular artery, the great auric the others. They are received by the initial part
ular vein anastomoses with the anterior auricular of the azygos vein as it extends forward from an
by means of a venous circle which lies on the anastomosis with the stem which receives the
cervicoauricular and interauricular muscles. This right and left third lumbar veins. The initial part
venous circle receives tributaries from the adja of the azygos vein may be double as it forms this
cent muscles. The great auricular vein termi union. The azygos vein carries most of the blood
nates in the internal maxillary vein. A portion of from the vertebral venous sinuses via the inter
the great auricular vein is bridged superficially costal and lumbar veins to the precava (Bowsher
by parotid gland tissue. 1954). The intervertebral veins of the thorax are
One or two veins enter the internal maxillary single vessels which join the intercostal veins at
near its termination. These come from the dor the highest points of the several intercostal
sally lying skin and underlying brachiocephalic spaces. The lumbar intervertebral veins are
muscle. Occasionally one of these ends in the double as they traverse the intervertebral foram
external jugular vein. A communicating vein ina. They then quickly unite to form the sev
may be located between the internal jugular and eral lumbar veins (Worthman 1956).
the internal maxillary vein. The hemiazygos vein (v. hemiazygos) is about
2 mm. in diameter and is extremely variable. It
lies on the left side of the aorta and connects the
Azygos System of Veins postcava with the azygos vein. It runs from the
left phrenicoabdominal vein near its termination
The azygos vein (v. azygos) (Fig. 5-7) is about in the postcava through the aortic hiatus and
8 mm. wide at its junction with the precava usually anastomoses cranially with the ninth or
where the precava terminates in the right atrium tenth left dorsal intercostal vein close to the
opposite the right third intercostal space. It be vertebral bodies. The hemiazygos vein receives
gins on the median plane ventral to the body of the left subcostal vein and the last two or three
the third lumbar vertebra by anastomosing with left dorsal intercostal veins. Occasionally there
the single trunk formed by the merging of the is an anastomosis between the left phrenicoab
right and left third lumbar intervertebral veins. dominal and the left subcostal vein. In such
Lying in the fat with the lumbar lymphatic specimens the hemiazygos is absent.
trunk, it runs forward, flanked by the tendons of The esophageal and bronchoesophageal veins
the crura of the diaphragm and the psoas major (w. esophageae et bronchoesophageae) are vari
muscles. In the caudal third of the thorax it in able and small. They are satellites of the com
clines slightly to the right, where it lies in the parable arteries. The bronchoesophageal veins,
angle formed by the vertebral bodies and the larger and more constant than the others, termi
aorta. Here, covered only by pleura, it ascends nate in the azygos vein, usually at the level of
to the base of the heart, hooks around the root of the seventh thoracic vertebra. The esophageal
the right lung, and empties into the termination veins, with delicate mediastinal tributaries, ter
of the precava at a right angle. It has the follow minate in the azygos vein caudal to the termina
ing tributaries: lumbar, subcostal, dorsal inter tion of the bronchoesophageal. There are usually
costal, esophageal, and bronchoesophageal veins. two of these, 1 to 3 cm. apart, which cross the
400 Chapter 5. The V e n o u s S y ste m
R. b r a c h i o c e p h a l i c v.
P recava C o s to c e r v ic a l- v e r te b r a l
tru n k
Common t r u n k
o f int. t h o r a c i c vv.
M. lo n g u s c o l l i
-In te rc o s ta l o .v v
E s o p h a g e a l br.~
- - S e v e n th t h o r a c ic
v erte bra
E ig h th r i b -
- C u t edge o f
d ia p h ra g m
M .psoas m in o r -
P o s tc a v a -
L e f t crus of
d ia p h r a g m
R, r e n a l v
right face of the aorta to empty into the ventral cephalica humeri) continues the antebrachial
surface of the azygos vein. Occasionally there is part from the flexor angle of the elbow joint. It
an anastomosis between the azygos or hemi runs proximally at first in the furrow between
azygos vein and the deep circumflex iliac vein. the insertions of the deltoideus and the brachio
cephalicus cranially and the origin of the lateral
VEINS OF THE THORACIC LIMB head of the triceps caudally. At the proximal
border of the lateral head it inclines medially
The veins of the thoracic limb may be divided and runs across the caudal surface of the humer
into superficial and deep sets. us between the long and lateral heads of the tri
ceps. Caudal to the shoulder joint it usually
Superficial Veins of the Thoracic Limb anastomoses with both the axillary and the sub
The cephalic vein (v. cephalica) (Figs. 5-2, 5 - scapular vein. It receives large tributaries from
8) is the only large superficial vein of the tho the triceps in its course through it. It terminates
racic limb. It begins as a transverse vein, the in the axillary vein as its last tributary. It has one
superficial palmar venous arch, which crosses anastomosis with the brachial vein and two with
the palmar side of the distal third of the fourth the external jugular.
The median cubital vein (v. mediana cubiti)
and third metacarpal bones. Here it is well pro
(Figs. 5-2, 5-8) extends between the median
tected by the heavy metacarpal pad. It runs
proximally from the superficial arch on the pal vein at the flexor angle of the elbow joint and
mar side of the interosseous muscles direcdy the cephalic vein of the arm. It is about 2 mm.
caudal to the interosseous space between the in diameter and 2 cm. long. It crosses the distal
second and third metacarpal bones. It passes end of the biceps obliquely as it runs proximo-
superficial to the palmar carpal transverse liga laterally to anastomose with the cephalic vein
ment, parallel to the carpal canal. It usually re near the lateral border of the biceps.
ceives three tributaries here. One of these comes The distal communicating vein (v. communi-
from a band of skin extending to the elbow, cans distalis), deeper than the proximal com
whereas the other two are tributaries from the municating vein, runs proximomedially under
flexor muscles of the antebrachium. Upon enter the brachiocephalicus upon leaving the cephalic
ing the antebrachium it is known as the ante vein at the junction of the middle and distal
brachial part o f the cep halic vein (v. cephalica thirds of the brachium. It enters the external
antebrachii). From the carpus it runs proximo- jugular between the omocervical and axillary
cranially until it reaches the cranial surface of veins. It receives a tributary from the major
the extensor carpi radialis and then it follows tubercle of the humerus, and two or three more
this muscle to the flexor angle of the elbow joint. from the brachiocephalicus and pectoral mus
It is flanked on its medial and lateral sides by the culature.
The proximal communicating vein (v. com-
medial and lateral branches of the proximal col
lateral radial artery and the medial and lateral municans proximalis) leaves the cephalic about
2 cm. proximal to the distal branch and runs
branches of the superficial radial nerve, respec
tively. It is 3 to 5 mm. in diameter and lies di superficially at first on the deltoideus. It then
arches forward and inward, crosses the brachio
rectly under the skin, loosely surrounded by the
cephalicus, and enters the lateral surface of the
superficial fascia. Because of its size, location,
and ease of compressibility, it is the favored site external jugular about 3 cm. cranial to the end of
the distal communicating branch. The proximal
for venipuncture in the dog. The v. cephalica
communicating vein has no muscular tributaries
antebrachii is augmented by receiving the acces
and receives only small vessels from the skin and
sory cephalic vein (v. cephalica accessoria) at
the beginning of the distal fourth of the ante fascia.
brachium. This vein, about 2 mm. in diameter
at its termination, begins on the dorsum of the Deep Veins of the Thoracic Limb
metacarpus and passes proximally over the car
pus and the dorsum of the distal portion of the The deep veins of the thoracic limb are rep
antebrachium before joining the cephalic vein. resented at the proximal end of the carpus by
It is joined by a tributary from the first digit and the radial, ulnar, and the interosseous branch of
skin of the second metacarpal bone at the distal the common interosseous vein. These three
end of the antebrachium. veins lie on the palmar surface of the ante
The brachial part o f the cephalic vein (v. brachium and anastomose distally not only with
402 Chapter 5. The V e n o u s S y s te m
C e p h a lic v - - y f -
Median c u b i t a l v.
C e p h a l i c v.
C ephal ic v.
A c c e s s o r y c e p h a l ic v.
each other but also with the cephalic and acces may enter the axillary or cephalic and the other
sory veins. branch may enter the beginning of the axillary
The radial vein (v. radialis) (Fig. 5-9) is very or termination of the brachial. Usually three
small, being only about one-third as large as its pectoral branches (rami pectorales) enter the
satellite artery. It arises from the deep palmar medial side of the axillary at its termination, and
venous arch (arcus venosus profundus palmaris). another tributary comes from the nerves leaving
It follows the mediocaudal border of the radius, the brachial plexus.
where it is covered by the deep antebrachial
fascia. It joins the small ulnar vein proximal to Veins of the Forepaw
the origin of the terminal radial and ulnar
branches of the brachial artery. The veins of the forepaw (manus) (Fig. 5-9),
The ulnar vein (v. ulnaris ) is about the same like the arteries, nerves, and lymphatics, are di
diameter as its companion artery. It leaves the vided into a dorsal and a palmar set. These are
supracarpal venous arch and runs proximally in not as completely divided into superficial and
the deep digital flexor. It receives tributaries deep series as are the arteries in the metacarpus,
from most of the flexor muscles lying in the ante and only single series exist dorsally and palmarly
brachium. It joins the small radial vein at about in the digits.
the junction of the proximal and middle thirds of The single dorsal digital veins II, III, IV, and
the antebrachium to form the median vein. V (vv. digiti dorsales) begin in the arcus venosus
The median vein (v. mediana) describes a sig digitales formed at the distal ends of the second
moid flexure before becoming the brachial vein phalanges, by the anastomoses of the dorsal and
in the arm. It is continued in the axilla as the palmar sets of digital veins. They occur on the
axillary vein after it traverses most of the bra medial aspects of digits II and III and on the
chium. The median vein receives the palmar an lateral aspects of digits IV and V. These digital
tebrachial vein (v. antebrachialis palmaris) about arches collect small tributaries from the digital
1.5 cm. distal from the place where it collects pads and the corium of the claws. The dorsal
the relatively large common interosseous vein (v. digital veins run proximally on the dorsum of the
interossea communis), which enters its caudal digits and receive communicating branches from
side. The common interosseous vein has one trib the palmar digital veins.
utary, the interosseous branch (ramus inter The superficial dorsal metacarpal veins II,
osseus). This latter branch connects distally III, and IV (vv. metacarpeae dorsales super-
with the supracarpal, the deep palmar, and the ficiales) continue proximally on the extensor ten
proximal palmar venous arches. The median dons from their formation by the confluence of
vein communicates with the cephalic via the the dorsal digital veins and the palmar commun
median cubital vein. It then crosses the biceps icating branches. The lateral or superficial dorsal
in company with the proximal collateral radial metacarpal vein IV runs proximomedially and
artery and becomes the brachial vein (v. brachi joins the superficial dorsal metacarpal vein III.
alis) caudal to the brachial artery. It receives in The trunk formed by this union continues the
succession the bicipital vein (v. bicipitalis), the axis of the fourth vessel and in turn is joined
proximal collateral ulnar vein (v. collaterals by the superficial dorsal metacarpal vein II to
ulnaris proximalis), and the deep brachial vein form the accessory cephalic vein (v. cephalica
(v. profunda brachii), which may be double. accessoria). At the level of the first digit a small
These are satellites of their companion arteries. anastomosis usually occurs between the dorsal
The axillary vein (v. axillaris) (Fig. 5-2) is a metacarpal vein I and the termination of the
continuation of the brachial vein. It receives the superficial dorsal metacarpal vein II on the
cephalic vein, cranial circumflex humeral vein, medial surface of the second metacarpal bone.
lateral thoracic vein, and subscapular vein. The The deep dorsal metacarpal veins II, III, and
subscapular vein (v. subscapularis) receives the IV (vv. metacarpeae dorsales profundae) are
following tributaries, which are satellites of the delicate veins which lie in the dorsal grooves be
corresponding arteries: the cutaneous branches tween the main metacarpal bones. They anasto
(rami cutanei), the circumflex vein o f the scapula mose with the corresponding superficial dorsal
(v. circumflexa scapulae), the thoracodorsal vein metacarpal veins at the junction of the middle
(v. thoracodorsalis), and the caudal circumflex and distal thirds of the metacarpus and with the
vein o f the humerus (v. circumflexa humeri poorly formed dorsal rete of the carpus. The
caudalis). Before terminating, the subscapular blood from the dorsal part of the first digit and
vein frequently bifurcates so that one branch the medial surface of the second metacarpal
404 Chapter 5. The V e n o u s S y s te m
P a lm a r m e ta c a rp a l
S u p e r f i c i a l d o rs a l vv. I I , I I I v I V
D is t a l p a lm a r
m e ta ca rp a l
venous a rc h
vv. II, I I I t IV
Dorsal P a lm a r d i g i t a l vv.
d i g i t a l vv.
DORSAL PALMAR
drains into the accessory cephalic vein some 4 interosseous branch of the common interosseous
cm. proximal to the carpus via the dorsal meta vein. Usually a second venous arch exists here,
carpal vein I. This vein collects a tributary from connecting the cephalic vein with the interosse
the dorsal venous rete of the carpus. ous branch across the superficial surface of the
The dorsal venous rete of the carpus (rete superficial flexor tendon. It lies subcutaneously,
venosum dorsale carpus) is a minute, poorly de just distal to the carpal pad. The palmar skin and
fined plexus of veins on the dorsal surface of the small carpal pad and the skin on the medial sur
distal row of carpal bones. The deep dorsal face of the second metacarpal and digit are
metacarpal veins drain into it, and the accessory frequently drained by a single vein which termi
cephalic, dorsal metacarpal vein I, and the pal nates in the medial surface of the cephalic op
mar set of veins carry blood from it. posite the carpus. In some specimens a com
The palmar set of veins of the forepaw begin municating vein connects the cephalic with the
usually as the single but occasionally double dorsal metacarpal vein I near the level of the
palmar digital veins II, III, IV and V (vv. digi metacarpophalangeal joint.
tales palmares). These commence at the palmar
extremities of the sagittally placed digital venous POSTCAVA, OR CAUDAL VENA CAVA
arches and run proximally on the palmar sur
faces of the proximal interphalangeal joint and The postcava, or caudal vena cava (v. cava
the adjacent phalanges. Usually the middle two caudalis) (Fig. 5-10), begins in contact with the
veins divide on the first phalanges into medial ventral surface of the seventh lumbar vertebra
and lateral branches. The apposed (axial) by convergence of the common iliac veins. It is
branches converge toward an axis through the about 1 cm. in diameter in large dogs and lies in
paw and anastomose with the communicating the furrow formed by the right and left psoas
veins from the dorsal set and with each other or major and minor muscles. At its beginning the
extend singly to the superficial palmar venous aorta lies to the left of it, since the aorta termi
arch and anastomose with it. The abaxial nates ventral to the left common iliac vein. In
branches anastomose with the palmar digital its course cranially it gradually inclines ventrally
veins nearest them. Thus the medial branch until it reaches the medial part of the caudate
anastomoses with the second palmar digital vein lobe of the liver. It then inclines ventrally and
and the lateral branch anastomoses with the to the right at a slightly sharper angle, and
fifth. In this way the palmar common digital deeply grooves or tunnels the caudate lobe of
veins II, III, and IV (vv. digitales communes the liver as it passes in it before reaching the dia
palmares) are formed. These immediately anasto phragm. It passes through the obliquely placed
mose with veins of the dorsal set and then con foramen venae cavae of the diaphragm which is
tinue to anastomose with the distal palmar ve about 3 cm. to the right of the median plane.
nous arch (arcus venosus palmaris distalis). This The intrathoracic portion of the postcava, about
arch is formed by an anastomosis of the cephalic 4 cm. long, lies in a special pleural fold, the plica
vein, medially, and the fourth palmar metacarpal venae cavae, in company with the right phrenic
vein, laterally. It lies deeply under the meta nerve. Here both the nerve and vessel lie in a
carpal pad, on the palmar surfaces of the meta deep groove of the intermediate lobe of the right
carpophalangeal joints. Occasionally the arch is lung before the vein terminates in the caudal
double, and other irregularities exist. part of the right atrium.
The deep palmar metacarpal veins II, III, and Reis and Tepe (1956), after examining 500
IV (vv. metacarpeae palmares) are small satel dogs for variations in the postcava and renal
lites of the deep palmar metacarpal arteries. veins, list only two variants. One of these had
They lie between the fleshy interosseous muscles been described previously by Kadletz (1928) and
and run from the distal palmar venous arch represented a circumaortic venous ring at the
proximally to anastomose with the deep palmar level of the renal veins. The other aberration
venous arch. was a persistence of the left supracardinal vein of
The proximal palmar venous arch (arcus ve the fetus. (The right supracardinal forms the de
nosus palmaris proximalis) lies under the origins finitive postcava.) This was found in 2.6 per cent
of the palmar muscles and follows the distal bor of the 500 dogs examined. Aberrations in the de
der of the thick palmar carpal ligament. Medi velopment of the postcava of the cat are com
ally it connects with the cephalic vein superfici mon, compared with those in the dog. Reis and
ally and with the radial vein deeply; laterally it Tepe (1956) state that there are probably four
anastomoses deeply with both the ulnar and the longitudinal venous pathways in the lumbar re
406 Chapter 5. The V e n o u s S y s te m
gion of the canine fetus. These are the right and verse the inguinal canal. The vein straightens on
left caudal (posterior) cardinal veins and the right approaching the vaginal ring. Throughout its ob
and left supracardinal veins. Any one of these lique intra-abdominal course to the postcava it
venous pathways other than the normal right lies in a plica of peritoneum which may be 4 cm.
supracardinal may persist, or any combination wide near the inguinal canal. The testicular vein
of them, to give any one of 15 different anoma is joined by one or two small tributaries from the
lous postcaval types. The postcava in the dog is adipose and true renal capsules a few centi
essentially devoid of muscle (Franklin 1937). It meters before its termination. It is accompanied,
has the following tributaries, in addition to the except at its termination, by the testicular ar
formative common iliac veins: lumbar, deep cir tery, nerves, and lymphatics.
cumflex iliac, right testicular or right ovarian, re The right ovarian vein (v. ovarica dextra) is
nal, phrenicoabdominal, hepatic, and phrenic shorter and less tortuous than the homologous
veins. right testicular vein. It begins by two or three
The lumbar veins I, II, III, IV, V, VI, and VII tributaries from the right ovary and surround
(vv. lumbales) are satellites of the corresponding ing fat. These are tortuous and plexiform. The
arteries. The first two lumbar veins are tributar caudal or uterine tributary freely anastomoses
ies of the azygos vein on the right side and of the in the broad ligament opposite the cranial end
hemiazygos on the left. The members of the of the uterine horn with the larger uterine vein.
third pair of lumbar veins anastomose with each The renal veins (w. renales) (Fig. 5-10) are
other directly ventral to the body of the third about 8 mm. in diameter. The right renal vein is
lumbar vertebra. From this small venous yoke a about 3 cm. long, whereas the left is 4 cm. long.
small median unpaired vein runs forward to be Each begins at the hilus of the respective kidney
come the azygos vein. In a similar manner a by convergence of the two veins which arise
larger unpaired median vessel runs caudoven- near the poles of the kidney by collecting the in
trally and enters the common trunk formed by terlobar veins. Christensen (1952) describes the
the anastomoses of the right and left fourth lum intrarenal morphology of the renal veins of the
bar veins. This trunk vein is the largest vessel dog. The renal veins may take an oblique course
entering the postcava from the lumbar verte forward to reach the postcava. The renal veins
brae. Because of these venous anastomoses in contain valves at their terminations. The left re
the middle of the lumbar region, blood can flow nal vein receives the left gonadal vein, the left
forward to the heart either by the azygos vein or testicular vein (v. testicularis sinistra) or the left
by the postcava. The members of the fifth and ovarian vein (v. ovarica sinistra). Except for the
sixth pairs of lumbar veins anastomose with each difference in termination, the left gonadal vein
other to form common trunks which enter the closely resembles its fellow on the right, which
ventral surface of the caudal part of the post empties directly into the postcava. The left gon
cava. The right and left seventh lumbar veins adal vein or the left renal vein receives the left
empty into the right and left common iliac veins, cranial ureteric vein (v. ureterica cranialis sinis
respectively. tra) several centimeters before its termination,
The deep circumflex iliac vein (v. circumflexa whereas the right renal vein usually receives the
ilium profunda) is a satellite of the correspond right cranial ureteric vein (v. ureterica cranialis
ing artery and lies caudal to it. The superficial dextra) near the hilus of the kidney. In the 500
and deep tributaries have the same anastomoses dogs examined by Beis and Tepe (1956), double
as do the comparable arteries. renal veins were found five times on the right
The right testicular vein (v. testicularis dex side but never on the left.
tra) enters the ventral surface of the postcava The phrenicoabdominal veins (vv. phrenico-
about 2 cm. caudal to the termination of the abdominales) (Fig. 5-10) are about 4 mm. in di
right renal vein. It collects tributaries from the ameter as each terminates in the lateral surface
testis and epididymis and becomes greatly of the postcava about 1 cm. cranial to the renal
coiled as it continues in the free border of the vein of the same side. Each grooves the ventral
mesorchium. This coiled and flexuous arrange surface of the corresponding suprarenal gland as
ment of the testicular vein is known as the pam the terminal 1 cm. of the vein passes under the
piniform plexus (plexus pampiniformis). The gland. Here it receives the adrenal veins (w.
plexus is intertwined with the testicular lym suprarenales) which, unlike the arteries, drain
phatics, artery, and nerve plexus as they form a entirely into the phrenicoabdominal as it lies in
funiculus which is located cranial to the ductus the groove of the gland (Flint 1900). These veins
deferens and its blood vessels, as they all tra are short and inconspicuous. The formative
P o r t a l V e in 407
phrenic and cranial abdominal parts of the mation of these veins. The cranial mesenteric
phrenicoabdominal vein are satellites of the ar vein is always the largest vessel.
teries of the same names. The cranial mesenteric vein (v. mesenterica
The hepatic veins (vv. hepaticae) (Fig. 5-10) cranialis) is 8 to 10 mm. in diameter at its termi
are embedded, wholly or in part, in the liver nation. It collects approximately 12 jejunal and
parenchyma. They are therefore short and re ileal veins (vv. jejunales et ilei), which are satel
ceive numerous tributaries. They terminate lites of the corresponding arteries and, like
along the lateral and ventral surfaces of the last them, are divided into a proximal and a distal se
4 cm. of the intra-abdominal portion of the post ries. There are only primary formative arcades,
cava. The largest hepatic vein serves the left lat and these are largest in the middle of the series
eral, left medial, quadrate, and part of the right and smaller at each end. Some of the vasa recta
medial lobe. It enters the left ventral part of the are double as they flank the straight arteries.
postcava at the foramen venae cavae and is the The most distal ileal vein anastomoses with the
most cranially located of all the hepatic veins. ileal branch of the ileocecocolic, whereas the
From the right, usually two major hepatic veins most proximal jejunal vein forms an arcade with
enter the postcava, about 2 cm. apart. The cra the caudal pancreaticoduodenal vein (v. pan-
nial tributary comes from the right lateral and creaticoduodenalis caudalis). This latter vessel
partly from the right medial lobe. The caudal is the last tributary to enter the cranial mesen
tributary comes mostly from the caudate lobe. teric and is like the other intestinal vein, except
These vessels are about 3 mm. in diameter. that it enters the cranial side of the cranial mes
There are a score or more of hepatic tributaries, enteric.
ranging under 1 mm. in diameter, which drain The caudal mesenteric vein (v. mesenterica
into the postcava at various places as it courses caudalis) is not a satellite of the like-named ar
through the liver. tery. It begins in the pelvic cavity as the cranial
The phrenic veins (vv. phrenicae) are repre rectal vein (v. rectalis cranialis), which is a satel
sented by a single tributary on each side, begin lite of the cranial rectal artery. In the rectal
ning in the ventral part of the muscular periph plexus of veins at the pelvic outlet it anastomo
ery of the diaphragm. At the lateral junction of ses with the caudal rectal vein, which drains
the muscular and tendinous parts each empties blood into the caval system of veins. The cranial
into the postcava as it passes through the fora rectal vein continues forward from the pelvic in
men venae cavae. There is usually a small trunk let in the left mesocolon to the place where the
vein which empties into the postcava on the tho cranial rectal artery becomes associated with it.
racic side of the foramen venae cavae. This is Here it is continued as the left colic vein (v. col-
formed by two or more tributaries which drain ica sinistra) and collects approximately 25 left
the extensive but flimsy plica venae cavae. colic branches which are satellites of the vasa
recta of the left colic artery. Some of these may
PORTAL VEIN be double. The last tributary to enter the left
colic vein is larger than the others as it collects
The portal vein (v. portae) (Fig. 5-11) with its blood from the left colic flexure as well as the
tributaries forms a portal system. It arises from adjacent middle and left parts of the colon. Its
a capillary bed and ends in a capillary bed. It middle colic tributary anastomoses in an arcade
collects blood from the pancreas, spleen, and all with the middle colic vein. At the left colic flex
of the gastrointestinal tract except the anal ure the caudal mesenteric vein enters the meso
canal. It is about 1.2 cm. in diameter at the porta jejunum, in which it crosses the left face of the
of the liver, where it terminates. It is about 6 cranial mesenteric artery and associated struc
cm. long and lies deeply buried among the ab tures. It joins the cranial mesenteric vein to the
dominal viscera. It runs cranially from its for right of the left limb of the pancreas to form the
mation in the root of the mesojejunum, dorsal portal vein.
to the junction of the right and left limbs of the The common colic vein (v. colica communis)
pancreas. It continues forward from the pan enters the caudal mesenteric vein after receiving
creas in association with the hepatic artery and the ileocecocolic, right colic, and middle colic
plexus of autonomic nerves to form the ventral veins. The ileocecocolic vein (v. ileocecocolica)
boundary of the epiploic foramen. The portal from the ileum, cecum, and colon is the largest
vein is formed by the confluence of the cranial tributary of the common colic. The right colic
and caudal mesenteric and the gastrosplenic vein (v. colica dextra) drains the distal part of the
vein. Great variations exist in the patterns of for right colon, the adjacent right colic flexure, and
408 C h a p te r 5 . T he V e n o u s Sy s t e m
H e p a t ic vv.-
R. c ru s of diaphragm - L
Cran m e se n te ric a -
-L. phrenico-
P o s tc a v a a b d o m in a l v.
-L. r e n a l v.
R. t e s t i c u l a r v. ~L. t e s t i c u l a r v. or
o r o v a r ia n v.- O varian v.
A o rta - -K id n e y
■ -L. t e s t i c u l a r a.
o r o va ria n a.
Caudal mesenteric a.
R. int. i l i a c v.-
Caud. abdom inal a w .
R. ext. 11ia c u-
-Deep femoral v.
Pudendo-epigastric t r u n k -
F ic. 5-10. The postcava and its main tributaries, ventral aspect.
V e in s of the P e l v ic L im b 409
the beginning of the transverse colon. The right VEINS OF THE PELVIC LIMB
colic anastomoses with the colic tributary of the
ileocecocolic by the formation of a weak arcade. Superficial Veins of the Pelvic Limb
The middle colic vein (v. colica media) enters the
common colic less than 1 cm. from its termina
tion. It is formed by the vasa recta from the The large superficial veins of the pelvic limb,
transverse colon and the proximal and distal exclusive of the hindpaw, are the lateral saphe
anastomotic arcades, which are also partly nous and the superficial branch of the deep cir
formed by the right and left colic veins, re cumflex iliac on the lateral side and the medial
spectively. Occasionally the right and middle saphenous, saphenous, and the proximal part of
colic veins form a common terminal trunk. the femoral on the medial side. All of these
The gastrosplenic vein (v. gastrosplenica) veins, except the deep circumflex iliac, are com
(Fig. 5-11) is half to two-thirds as large as the monly used for venipuncture.
cranial mesenteric vein. It is about 5 mm. in The lateral saphenous vein (v. saphena later
diameter and 1.5 cm, long. It is formed by the alis) (parva) (Fig. 5-12) begins by collecting
confluence of the smaller caudally running left its dorsal branch from the flexor surface of the
gastric vein and the larger splenic vein. The left tarsus and its plantar branch from the lateral
gastric vein (v. gastrica sinistra) is formed by surface. The dorsal branch (ramus dorsalis) is 2
several veins which come from the lesser curva to 4 mm. in diameter as it inclines caudally in its
ture of the stomach adjacent to the cardia. Like proximal course. It obliquely crosses the lateral
the corresponding artery it anastomoses with the surface of the distal end of the tibia and is here
right gastric vein. The splenic vein (v. lienalis) frequently used for venipuncture. At the space
arises by two branches which receive tributaries between the deep caudal crural muscles and the
from the long hilus of the spleen. The pancreatic beginning of the calcanean tendon it receives
veins (vv. pancreaticae) are represented by two the smaller plantar branch (ramus plantaris) and
tributaries from the left limb of the pancreas; becomes the lateral saphenous vein. This crosses
they may terminate separately in the last 2 cm. the beginning of the calcanean tendon and re
of the splenic. The left gastroepiploic vein (v. ceives a small tributary from the skin covering
gastroepiploica sinistra) is a satellite of the left the calcanean tuberosity. It continues its subcu
gastroepiploic artery; it comes from the greater taneous course proximally on the caudal surface
curvature of the stomach and collects many epi of the gastrocnemius to the popliteal lymph
ploic tributaries along its course. node. It runs deep to the node and follows the
The gastroduodenal vein (v. gastroduodenalis) intermuscular septum between the m. biceps
(Fig. 5-11) is 3 to 4 mm. in diameter and emp femoris and the m. semitendinosus to terminate
ties into the portal vein about 1.5 cm. from the in the femoral vein in the popliteal fossa.
hilus of the liver. Its chief formative tributary is The superficial branch (ramus superficialis) of
the cranial pancreaticoduodendal vein (v. pan- the deep circumflex iliac vein (v. circumflexa
creaticoduodenalis cranialis). Its pancreatic and ilium profunda) drains the caudal half of the dor
duodenal tributaries begin as small anastomoses sal two-thirds of the skin of the abdominal wall
with the pancreatic and duodenal parts of the and the cranial half of the rump and proximal
caudal pancreaticoduodenal vein (v. pancreati- part of the thigh.
coduodenalis caudalis) near the caudal flexure of The medial saphenous vein (v. saphena me
the duodenum. At its termination it receives a dialis) (magna) (Fig. 5-13) begins by the conflu
small tributary from the left limb of the pan ence of its dorsal and plantar branches medial to
creas. The right gastroepiploic vein (v. gastroepi the stifle joint. The dorsal branch, traced from
ploica dextra) and the right gastric vein (v. gas the flexor surface of the tarsus where it anasto
trica dextra) are satellites of their companion ar moses with the dorsal branch of the lateral
teries. They sometimes unite before blending saphenous, runs proximally across the m. tibialis
with the larger cranial pancreaticoduodenal to cranialis and the tibia. It is about 1 mm. in diam
form the gastroduodenal vein. They anastomose eter. The plantar branch, before joining the dor
with the left gastroepiploic and left gastric veins, sal branch, receives the fibular vein. The medial
respectively. saphenous vein and its main tributaries are sat
The portal vein divides, upon entering the ellites of the saphenous artery and its main
liver, into a small right branch, which is dis branches. A prominent m edial genicular vein
persed in the right lateral and the right medial from the stifle joint joins the saphenous vein
lobe, and a large left branch, which breaks up in above the confluence of the dorsal and plantar
the remainder of the liver. (Text continued on page 413.)
410 Chapter 5. The V e n o u s S y s te m
P o r ta l v.
B e fr a m pancreas
R . g a s t r i c v.
- - L . g a s t r i c v.
Gastroduodenal v.
L.gastroepiploic v.
R.gastroepipioic V.— A- - -Gastrosplenic v.
Cran. p a n c r e a t ic o
duodenal - S p le n ic v.
I -Caud. mesenteric
|- - Common col ic
-M id dle col ic
L. col ic
Cran. mesenteric
J e ju n a l vv.
P o p lite a l ly m p h n o d e --------
Lat. saphenous v.
C alcaneal t r i b u t a r y -
A n a s to m o t ic br.
Superf. d o rs a l m e t a t a r s a l v. I I
Superf. p la n ta r m e ta ta rs a l v.IV
Superf. d o r s a l m e t a ta r s a l v. I l l
F e m o r a l v.
M. s a r t o r i u s Caud. fe m o r o l v.
Med. sophenous v.
----------- M. g r a c i li s
Med. g e n i c u l a r v.
-----------P o p lite a l lym ph node
P la n ta r b r of med. sophenous
D orsal br
o f med. saphenous v.
A n a sto m os is w it h
d o rs a l b r o f lat. s a p h e n o u s v.
Superf. d o r s a l m e t a t a r s a l v .I I- —
■Superf. p l a n t a r m e t a t a r s a l v.
branches. In most specimens a tributary from stifle joint. These enter the popliteal vein proxi
the m. gracilis is received by the medial saphe mal to the joint. A cluster of veins enters this ve
nous about 1 cm. before its termination in the nous channel at the proximal end of the m. gas
femoral. It lies under the thin but strong medial trocnemius, where the popliteal vein becomes
femoral fascia between the caudal belly of the the femoral vein. The largest of these is the lat
m. sartorius and the m. gracilis. It is firmly an eral saphenous, which is three times larger than
chored by fascia between these muscles and is a any other vein distal to the stifle.
common site for venipuncture. It joins the fem The large caudal femoral vein (v. femoralis
oral vein at the apex of the femoral triangle. caudalis), 3 mm. in diameter, drains the biceps
The femoral vein (v. femoralis) (Figs. 5-10, 5 - femoris. It begins as cutaneous twigs from the
13) is accessible for venipuncture as it lies in the caudolateral surface of the thigh. Muscular trib
femoral triangle. This segment of the vessel is utaries ascend from the gastrocnemius. Others
about 8 cm. long and 5 mm. in diameter. It lies enter it from in front and medially from the
caudal to the femoral artery, from which the quadriceps, adductors, and hamstring muscles.
pulse can easily be taken, and cranial to the At the distal end of the femoral triangle, in the
small saphenous nerve. It is sufficiently large and vicinity of entry of the medial saphenous into it,
well attached to permit injections without com the femoral receives a tributary from the adduc
pression. tor and semimembranosus. In its course through
The cutaneous veins of the pelvic limb, exclu the femoral triangle the femoral vein receives a
sive of the hindpaw, are largely tributaries of the tributary which is the satellite of the most proxi
superficial veins. However, the skin of the cau mal muscular branch of the femoral artery. This
dal part of the rump, the region of the pelvic vein arises from the medial surface of the proxi
outlet, and the caudolateral part of the thigh are mal half of the m. gracilis and receives a large
drained by deeply lying veins. A cutaneous area tributary from the proximal part of the adduc
over the caudal part of the middle gluteal mus tor muscles. A small vein frequently arises in the
cle drains into the cranial gluteal vein, whereas region of the subcutaneous inguinal ring and en
a cutaneous tributary which drains into the ter ters the caudal part of the femoral. Entering the
minal part of the superficial lateral coccygeal cranial side of the femoral vein, 1 or 2 cm. from
vein and perineal veins serves the skin of the pel the abdominal wall, is a long vein which begins
vic outlet. A wide caudolateral zone of skin of in the distal part of the cranial belly of the m.
the thigh is drained by three or more tributar sartorius. It runs proximally on the m. vastus me-
ies of a large vein which descends in the biceps dialis and, after receiving a large tributary from
femoris and empties into the terminal part of the the m. rectus femoris, passes under the femoral
lateral saphenous vein. Others are formative artery and unites with the superficial circumflex
tributaries of the caudal and deep femoral veins. iliac vein (v. circumflexa ilium superficialis).
The cranial femoral vein (v. femoralis crani
alis) is the largest and the last tributary of the
Deep Veins of the Pelvic Limb femoral vein, which it enters laterally. As a
satellite of the cranial femoral artery, it begins
The deep veins of the pelvic limb, exclusive in the skin of the proximal, lateral surface of the
of the hindpaw, are largely satellites of the thigh and the adjacent dorsally lying gluteal re
neighboring arteries. The cranial tibial vein (v. gion. Its formative tributaries cross ventral to the
tibialis cranialis) lies medial to its companion ar middle gluteal muscle near its insertion and then
tery, which it approaches in size. It begins on unite and pass the cranial aspect of the neck of
the flexor surface of the tarsus as a continuation the femur and the caudal surface of the origin of
of the superficial dorsal metatarsal vein II with a the m. rectus femoris to enter the lateral surface
contribution from the medial tarsal vein. It re of the femoral at the vascular lacuna. It may re
ceives tributaries from the cranial crural group ceive the superficial circumflex iliac vein at its
of muscles mainly at their proximal ends. It termination, when this vein does not enter the
passes between the tibia and fibula, runs under femoral directly. The femoral vein lies ventro
the m. popliteus, and unites with the delicate medial to the m. iliopsoas as it passes through
caudal tibial vein (v. tibialis caudalis) to form the abdominal wall to become the external iliac
the popliteal vein (v. poplitea). The popliteal vein (v. iliaca externa). This venous trunk is
vein traverses the popliteal notch of the tibia to about 6 mm. in diameter and 3 cm. long. It
enter the popliteal fossa. Here it receives a me arches dorsocranially across the medial surface
dial and a lateral tributary from the sides of the of the m. iliopsoas and ventral to the promon
414 Chapter 5. T h e V e n o u s S y s te m
tory of the sacrum and unites with the internal The vein of the clitoris (v. clitoridis) begins in
iliac vein to form the common iliac vein (v. iliaca the vestibular bulb and is not sharply differenti
communis). It has but one large tributary, the ated caudally from the vestibular plexus. The
deep fem oral vein (v. femoralis profunda), which vestibular plexus (plexus vestibuli) is a closely
enters the caudal side of the external iliac on the knit venous plexus which completely surrounds
abdominal side of the vascular lacuna. Its most the external vulvar opening. It extends about 1
caudal tributaries, like those of its accompany cm. cranially from the dorsal commissure of the
ing artery, are cutaneous twigs from the skin of vulva on its dorsal wall. Ventrally it widens to
the upper caudal part of the thigh. These merge such an extent that it runs proximally on the ure
to form larger vessels as they enter the proximal thra. This network, the urethral venous plexus
part of the hamstring musculature. Within these (plexus venosus urethralis), surrounds the female
heavy muscles further blending occurs so that urethra and splays out on the neck of the blad
the single vein emerges from between the m. der, between the heavy muscular coat and the
pectineus medially and the m. iliopsoas laterally. mucosa. Both the vestibular bulbs and the
At its termination it receives the large pudendo- plexus lie deep to the strong, partly divided con
epigastric trunk (truncus venosus pudendoepi- strictor vestibuli muscle. Between the vestibular
gastricus), which is disposed like the comparable bulbs ventrally, the vestibular venous plexus re
artery and its branches. Two small veins enter flects the division which occurs in the constric
the external iliac or deep femoral vein. One en tor vestibuli muscle in such a way that the
ters caudally from the adipose tissue inside the plexus caudal to the division is thick and annular
pelvic inlet, and the other enters cranially from in nature, whereas that cranial to the division is
the m. rectus abdominis. thin and longitudinal. After leaving the vestibu
The internal iliac vein (v. iliaca interna) (Fig. lar bulb, the vestibular vein receives (opposite
5-10), unlike the internal iliac artery, is not di the ventral part of the external anal sphincter)
vided into visceral and parietal parts. As a single the caudal rectal vein (v. rectalis caudalis), which
vein it lies between these two arteries. Its tribu comes directly from the external anal sphincter,
taries are satellites of the branches of the two anal sac, and the wall of the anal canal. It anasto
parts of the internal iliac artery. It is formed cau moses with the cranial rectal vein via the rectal
dally by the merging of the internal pudendal plexus of veins. Since the cranial rectal vein is
and caudal gluteal veins. indirectly a tributary of the portal and the cau
The caudal gluteal vein (v. glutea caudalis) dal rectal vein is a like tributary of the postcava,
(Fig. 5-14) arises mainly in the m. biceps fem the rectal plexus serves to unite the two systems.
oris with smaller tributaries coming from the The rectal venous plexus (plexus venosus rectalis)
mm. semimembranosus and semitendinosus. It is poorly developed. It lies on and in the muscu
ascends through the lesser ischiatic foramen lat lature of the rectum just cranial to the anal canal.
eral to the internal obturator muscle, medial to It is partly covered by the anal sacs when these
its companion artery and caudodorsal to the are large. The perineal vein (v. perinealis) from
ischiatic nerve. At the pelvic outlet it collects a the cutaneous anal structures ends in the ter
superficial tributary from the proximal caudal mination of the vaginal vein or directly in the
half of the thigh and another one from the inter internal pudendal vein.
nal obturator muscle. Lateral to the m. coccygeus The superficial lateral coccygeal vein (v. coc-
a branch from the fat of the ischiorectal fossa and cygea lateralis superficialis) is the main venous
the first (with, occasionally, the second) sacral in drainage from the tail. It is about 3 mm. in di
tervertebral vein enter the caudal gluteal vein. ameter at the ischiorectal fossa, through which
The internal pudendal vein (v. pudenda in it runs to enter the caudal gluteal vein. It re
terna) is formed at the root of the penis by the ceives segmental branches as it runs forward
convergence of the v. dorsalis penis and truncus from the free end of the tail. At the pelvic outlet
venosus profundus penis et bulbi. These are sat it receives a large tributary from the skin of the
ellites of the comparable arteries. The trunk so tail and the adjacent part of the rump. It con
formed usually receives the perineal vein. (Fora tains valves at intervals of about 1 cm.
complete description of the veins of the penis The urogenital vein (v. urogenitalis) has cra
and male perineum the reader is referred to nial and caudal branches. In the male the cranial
Chapter 15, on Urogenital System.) The internal branch is formed by the convergence of the v.
pudendal vein receives the third and sometimes ductus deferens and the v. ureterica caudalis
the second sacral intervertebral vein. In the fe joining the v. vesicae caudalis. The caudal
male the vein of the clitoris takes the place of the branch is formed by the confluence of the v.
three veins from the penis. prostatica and the v. urethralis. The cranial and
V e in s of the P e l v ic L im b 415
M u s c u l a r ra m u s S u p e rfic ia l g lu te a l m.
In te rn a l pudendal a, * v, 1 i C ra n ia l q lu te a l a. 4 v.
i------r
Superf.
la t coccygeal via. f~ M i d d le
g l u t e a l m.
A. xr v. o f c lito ris C r a n ia l
fe m o r a l a *
Ischiatic n-
S em itendinosus m-
-S a rto riu s m.
S em im em branosus m-
A d d u c to r m.
- L a t cutaneous s u ra l
Caud. c ru ra l abd. m-
- -Q u a d rice ps fe m o ris m.
T ib ia l n-
- F e m o ra / a. + v.
Dist caud. fe m o r a l a - -
Caud. cutan. s u r a l n. - ■ -F ib u la r n.
F ig . 5 - 14. Deep structures of the gluteal and femoral regions, lateral aspect. (From Miller 1958.)
416 Chapter 5. The V e n o u s S y s te m
caudal venous branches may be double as they lite of the iliolumbar artery. It crosses the cranial
unite in the pelvic fascia to form a double uro border of the wing of the ilium and terminates in
genital vein. The formative tributaries of the the lateral side of the internal iliac vein at its
urogenital are satellites of their companion ar termination. It receives the seventh lumbar
teries. intervertebral vein (Worthman 1956).
In the female the development of the uro The common iliac vein (v. iliaca communis) is
genital vein is directly proportional to the geni about 8 mm. in diameter and 5 cm. long. It is
tal activity of the bitch. In late pregnancy and formed by the confluence of the external and
immediately after parturition the uterine tribu internal iliac veins on the tendon of the psoas
tary is larger than all the others together. In the minor about 2 cm. cranial to its insertion. From
nongravid bitch the cranial and caudal tribu their origins the right and left common iliac
taries are similar to those in the male. The cranial veins converge and merge, usually ventral to the
tributary is formed by the caudally running sixth lumbar vertebra, to form the postcava. Oc
uterine vein (v. uterina) meeting at a right angle casionally the union of the common iliac veins is
the caudal vesical vein (v. vesicalis caudalis). unusually far forward, but this anomaly is more
(The caudal ureteral vein [v. ureterica caudalis] common in the cat than in the dog, according to
ends in the proximal part of the caudal vesical.) Darrach (1907) and Huntington and McClure
The cranial and caudal ureteral veins anasto (1920).
mose. The caudal vesical vein anastomoses with The median sacral vein (v. sacralis media)
the cranial vesical vein, if one is present. The (Fig. 5-10) is the unpaired median vein which
uterine vein begins in an anastomosis with the receives the middle coccygeal vein from the tail.
uterine tributary of the ovarian vein. This anasto It runs forward between the right and left ven
mosis takes place opposite the caudal third of tral sacrococcygeal muscles and terminates, ac
the uterine horn. The main vessel follows the cording to Worthman (1956), in both the right
companion artery and therefore lies as much as and the left common iliac vein, or in only one of
3 to 4 cm. out in the broad ligament caudally. them. It is a small vein, about 1 mm. in diameter,
There is, however, a smaller venous channel usually devoid of significant tributaries. In one
which lies in the uterine wall opposite the specimen it was large and collected as single
attached border of the uterine horn and extends short tributaries the first two right and left sacral
throughout its length. intervertebral veins and the last pair of lumbar
This vein is connected, by about five com intervertebral veins, after they had united in a
municating veins, with the main uterine vein common trunk.
which lies in the broad ligament. From both
Veins of the Hindpaw
sides of the uterine wall it receives many tortu
ous tributaries which anastomose with each The veins of the hindpaw (pes) (Fig. 5-15) are
other but do not form a definite uterine plexus. divided into a dorsal and a plantar set. In the
Near the ovarian end of the uterine horn the metatarsus these are further divided into a super
converging uterine veins anastomose with each ficial and a deep series. Anastomoses occur be
other and with the uterine extension of the tween the dorsal and plantar series at both the
ovarian vein. The caudal branch of the urogeni proximal and the distal end of the intermetatarsal
tal vein, the vaginal vein (v. vaginae), arises in spaces. The veins of the hindpaw in the dog have
the submucosa of the wall of the vagina as the been described by Preuss (1942).
fine vaginal venous plexus (plexus venosus The digital venous arches (arci venosi digi
vaginalis). Unlike the comparable artery, it does tales) are anastomoses between the dorsal and
not serve the urethra. Caudally it anastomoses plantar digital veins. One for each digit, they lie
with the vestibular plexus of veins. on the abaxial sides of the distal ends of the sec
The urogenital vein empties into the medial ond phalanges and receive tributaries from the
surface of the internal iliac opposite the cranial claws, digital pads, and terminal phalanges. The
gluteal vein (v. glutea cranialis). single or paired dorsal and plantar digital veins
The cranial gluteal vein is a satellite of the like arise proximally from the arches.
artery and therefore drains most of the proximal The dorsal digital veins II, III, IV, and V (w.
part of the middle gluteal muscle. It usually has a digitales dorsales pedis) arise from the axial and
prominent cutaneous tributary which drains the abaxial sides of the digits. They receive anasto
skin over the proximal dorsal part of the rump moses from the plantar digital veins at the distal
and also receives the first and occasionally the end of the metatarsus, in the respective digital
second intervertebral vein. clefts.
The iliolum bar vein (v. iliolumbalis) is a satel The superficial dorsal metatarsal vein III (v.
V e in s of the P e l v ic L im b 417
metatarsea dorsalis superficialis) is formed by saphenous veins are either confluent or joined by
the axial dorsal digital veins of the third and a short anastomosis at the level of the distal end
fourth digits. The superficial dorsal metatarsal of the tibia. Figure 5-15 represents confluent
veins II and IV are formed by the second and dorsal saphenous veins, whereas Preuss (1942)
fifth dorsal digital veins which anastomose with illustrates a short anastomosis.
the abaxial dorsal branches of the third and The plantar set of veins of the hindpaw begins
fourth dorsal digital veins, respectively. The as single or paired plantar digital veins (vv. digi-
superficial dorsal metatarsal veins anastomose tales plantares). These originate as plantar con
with the deep plantar metatarsal veins near the tinuations of the digital venous arches, and col
distal ends of the metatarsal bones. The third lect tributaries from the digital pads, skin, and
and fourth superficial dorsal metatarsal veins terminal phalanges.
anastomose with each other at the middle of the The superficial plantar metatarsal veins (w.
metatarsus, and the resultant common trunk is metatarseae plantares superficiales) are short
joined near the proximal end of the metatarsus and drain into the distal plantar venous arch
by the second superficial dorsal metatarsal vein. (arcus venosus plantaris distalis). The arch is
The common venous trunk formed by the three continued medially by the dorsal branch of the
superficial dorsal metatarsal veins is the dorsal medial saphenous vein and laterally by the plan
branch o f the lateral saphenous vein (ramus tar branch of the lateral saphenous vein. The
dorsalis v. saphenae lateralis), which is called the plantar metatarsal portions of the medial and
v. metatarsea dorsalis superficialis proximalis III lateral saphenous veins are called the v. meta
by Preuss (1942). This vessel continues proxi tarsea plantaris superficialis proximalis II and the
mally on the long digital extensor tendon to the v. metatarsea plantaris superficialis proximalis
distal end of the crus. Opposite the talocrural IV, respectively, by Preuss (1942).
joint it receives the lateral tarsal vein (v. tarsea The plantar branch of the lateral saphenous
lateralis). The lateral tarsal vein also connects vein (ramus plantaris v. saphenae lateralis) passes
with the plantar branch of the lateral saphenous proximally in the superficial metatarsal fascia
vein and receives tributaries from the joint cap along the lateral surface of the metatarsus. At
sule, skin, and transverse metatarsal vein. the proximal end of the metatarsus it receives
The deep dorsal metatarsal veins II, III, and the proximal plantar venous arch (arcus venosus
IV (vv. metatarseae dorsales profundae) are plantaris proximalis), formed by the deep plan
small veins which lie in the grooves between ad tar metatarsal veins.
jacent metatarsal bones. They enter the trans The plantar branch of the medial saphenous
verse metatarsal vein proximally. vein (ramus plantaris v. saphenae medialis) is the
The dorsal branch of the medial saphenous smallest of the main saphenous branches. It be
vein (ramus dorsalis v. saphenae medialis) paral gins from the larger medial tarsal vein, distal to
lels the tendon of the m. tibialis cranialis as it the medial malleolus. In the crus it is related to
runs proximally over the flexor surface of the the plantar branch of the saphenous artery, and
tarsus. Upon leaving the tarsus it receives the joins the larger dorsal branch of the medial
m edial tarsal vein (v. tarsea medialis) from the saphenous vein opposite the proximal end of the
plantar surface of the tarsus. At the talocrural tibia.
joint the dorsal branch of the medial saphenous The two or three deep plantar metatarsal
vein receives a long anastomotic branch from veins (vv. metatarseae plantares profundae) are
the second superficial dorsal metatarsal vein. small and lie in the intermetatarsal grooves or on
This anastomotic branch is the v. metatarsea the plantar surface of the third and fourth met
dorsalis superficialis proximalis II of Preuss atarsal bones. Near the middle of the metatarsus
(1942). It receives an anastomotic branch from they anastomose with the superficial dorsal met
the proximal plantar venous arch which passes atarsal veins by passing between the respective
between metatarsal bones II and III. metatarsal bones. Proximally the deep plantar
The transverse metatarsal vein (v. metatarsea metatarsal veins enter the proximal plantar ve
transversa) crosses the dorsal surfaces of the nous arch.
proximal ends of the metatarsal bones. It re The medial and lateral saphenous veins are
ceives the threadlike deep dorsal metatarsal the main vessels which return blood from the
veins, and anastomoses with the plantar branch hindpaw. The lateral saphenous is appreciably
of the lateral saphenous vein and with the more larger than the medial saphenous. As its dorsal
proximal, lateral tarsal vein. tributary obliquely crosses the distal lateral sur
The dorsal branches of the lateral and medial face of the tibia, it may be used for venipuncture.
418 Chapter 5. T h e V enous S y s te m
T ra n s v e rs e m e t a ta r s a l - A, , -T ransverse
P r o x i m a l s u p e r f ic i a l
m e ta ta rs a l
Deep d o rs a l -
\r*?i p l a n t a r m e t a t a r s a l I I --------- 5
-Prox. p l a n t a r
m e t a t a rsa I vv.
Deep p l a n t a r - rr' venous a rc h
m e t a ta r s a l vv.
D is t p la n ta r
\--v e n o u s a rc h
- S u pe rf. d o r s a l
m e t a t a r s o l vv. ) \ vy4 -Superf. p l a n t a r
IV, I I I , II m e t a t a r s a l vv. II, III, IV
D o rs a l d i g i t a l vv. - P l a n t a r d i g i t a l vv.
VEINS OF THE CENTRAL nus to bifurcate after the straight sinus has
NERVOUS SYSTEM entered it. The straight sinus is about 1.5 mm. in
diameter and 5 mm. long. It begins at the free
Venous Sinuses of the Cranial Dura M ater posterior margin of the falx cerebri by the merg
ing of the great cerebral vein, ventrally, and the
Within the dura, usually between its periosteal vein of the corpus callosum, dorsally. The straight
and meningeal parts, and in certain places sinus lies in the posterior border of the falx cere
within large osseous canals, there are venous bri as the right and left parts of the tentorium
passages into which the veins of the brain and of cerebelli join it. It lies anterodorsal to the ten
its encasing bone drain. These passages, known torium ossium.
as the sinuses of the dura mater (sinus durae The transverse sinus (sinus transversus) (Fig.
matris), in the dog are not confined exclusively to 5-17) is paired. Each begins mid-dorsally by re
the dura. By means of these passages the blood is ceiving the sagittal and occasionally the straight
conveyed from the brain and skull to the paired sinus, and merges with its fellow to form the con
maxillary, internal jugular, and vertebral veins, flu en ce o f the sinuses (confluens sinuum). This
and to the vertebral venous sinuses. They lack a triple or even quadruple merging of sinuses is
tunica media and tunica adventitia in their walls, located within the dorsal part of the occipital
and they do not have valves in their lumina. bone and may be asymmetrical. From the con
They are divided into dorsal and ventral sets, fluens sinuum the transverse sinus runs laterally
which freely intercommunicate. The dorsal set in the transverse canal for about the proximal
consists of the unpaired dorsal sagittal and two-thirds of its length and then continues in the
straight sinuses, and the paired transverse sinus. transverse groove. It terminates at the distal end
The ventral set consists of the double, unpaired of the transverse groove by dividing into the
intercavernous, and the paired cavernous, sig temporal and sigmoid sinuses. The temporal
moid, occipital, and dorsal and ventral petrosal sinus, larger than the sigmoid, continues in the
sinuses. The cranial venous sinuses have been direction of the transverse sinus through the
studied in a variety of vertebrates by Hofmann temporal meatus, whereas the sigmoid sinus
(1901), and in the dog by Zimmermann (1936), bends downward and backward on its way to
and Reinhard, Miller, and Evans (1962). A com the petrobasilar fissure.
parative developmental study of the cranial ve Both the transverse and the sigmoid sinus
nous system in man and dog was made by Padget have a connection with the occipital emissary
(1957). vein (v. emissaria occipitalis). This vein lies on
The dorsal sagittal sinus (sinus sagittalis dor the posterior surface of the skull and drains
salis) (Fig. 5-16) begins by the confluence of the blood from the deep muscles on the cranial part
right and left rhinal veins (vv. rhinales), which of the neck. Right and left veins lie ventral to
come from the osseous nasal septum and its the ventral nuchal line as they form a prominent
covering mucosa, and the olfactory bulbs and anastomosis with each other dorsally. The oc
their dural coverings. From near the middle of cipital emissary vein drains the area supplied by
the cribriform plate, where the sagittal sinus is the dorsal part of the occipital artery. The
formed, it runs posteriorly in the attached edge smaller linkage with the transverse sinus occurs
of the falx cerebri. It therefore lies directly ven lateral to the confluence of the sinuses. The
tral to the sagittal suture and the interparietal larger connection between this vein and the
process of the occipital bone. The sagittal sinus sinus system passes through the supramastoid
collects the dorsal cerebral and most of the foramen as it joins the first bend of the sigmoid
diploic veins and is usually joined by the straight sinus. The anterior side of the proximal third of
sinus near its termination. It measures about 3 the transverse sinus receives an occipital diploic
mm. in diameter at the foramen impar, which it vein, and a second such vessel may enter the
traverses to form a junction with the right and sinus as it leaves the transverse canal. The short,
left transverse sinuses. posteriorly running dorsal petrosal sinus enters
The straight sinus (sinus rectus) (Fig. 5-16) the transverse sinus near the ventral end of the
usually drains into the posterior part of the sagit transverse groove.
tal sinus before it enters the foramen impar, but The tem poral sinus (sinus temporalis) (Figs.
it may pursue an independent course through 5-16, 5-17) of Zimmermann (1936) is the an-
the accessory foramen impar and join the conflu teroventral continuation of the transverse sinus.
ence of the sinuses within the occipital bone. It lies in the temporal meatus and therefore is
Another frequent variation is for the sagittal si placed between the petrous and squamous parts
420 Chapter 5. T h e V e n o u s S y s te m
T h a la m o s tr io te v.
Dors, s a g i t t a l sirius,
fD o rs a l c e r e b r a l v.
O c c ip ita l c e re b r a l ^ , 1 '
I n t e r n a l c e r e b r a l v.
G r e a t cerebral v.t
V. o f corpus c a llo s u m
S t r a i g h t s in u s
P o s t e r i o r c e r e b r a l v. \ Dorsal p e tro sa l sinus
\
T ra n s v e rs e sinus Cavernous s in u s
Dors, p e tr o s a l sinus s. \
Midd le m e n in g e a l v.
O c c ip ita l em issary v.^
Temporal sinus Optic foramen
' , From deep temporal v.
Supramastoid fo r a m e n ^ /
- O phthalm ic v.
S ig m o id s in u s ^ ^
- - F r o m deep fa c ia l v.
Condyloid v. in c a n a l _ __
— O r b it a l plexus
Vent, o c c i p i t a l sinus- - -
" ~ ~ ~ - O r b it a l fis s u r e
J u g u la r fo r a m e n - • Foramen rotundum
Hypoglossal foramen''
canal
V e r te b r a l v.
Foramen ovale
In t ju g u la rv .' / ext. c a r o t id fo ra m in a
\ \
J u g u l a r fo ra m e n 1 J ' \ ^Ventral p e tr o s a l sinus
\ \
I n t. m a x i l l a r y v 1 C a ro tid c a n a l
' \
R e tro gle n oid v. * fo r a m e n 1 0f p a i a t i n e p le x u s
F ig . 5-16. Diagram of the cranial venous sinuses, lateral aspect. (From Reinhard, Miller, and Evans 1962.)
P a r i e t a l e m i s s a r y v.
D o rs a l s a g i t t a l s i n u s ( i I n t e r n a l cerebral v.
G r e a t ce re bra l v. ! Vein of corpus c allo sum
S t r a i g h t sinus Falx c e r e b r i
Confluens sinuum
T e n to riu m c e r e b e l I i
O c c ip ita l d i p l a i c v.
O c c ip ita l e m is s a ry v - O ptic foramen
T e m p o ra l s i n u s
O r b it a l f i s s u r e
V e r t e b r a l sin u s~ ~
V V ~ A n te rio r a l a r f o r a m e n
V e rte b ra l v - -
I n t j u g u l a r v.
sS c
Dorsal p e tr o s a l s in u s
R e tro g le n o id Post, c e re b r a l v.
F ig . 5-17. Cranial venous sinuses, lateral aspect. (Right cerebral hemisphere removed.) (From Reinhard, Miller and Evans
1962.)
V e in s of the C en tra l N erv o u s Sy st e m 421
of the temporal bone. It receives no tributaries. acute angle ventrally, lateral to the temporal
At the retroglenoid foramen it becomes the meatus.
retroglenoid vein (v. retroglenoidalis), which, The ventral petrosal sinus (sinus petrosus
after a course of about 1 cm., empties into the ventralis) (Fig. 5-18) extends between the pos
internal maxillary vein as one of its largest tribu terior end of the cavernous sinus and the ventral
taries. The retroglenoid vein forms a shallow end of the sigmoid sinus. It lies in the petro
vertical groove on the posterior side of the retro basilar canal and is an intraosseous posterolateral
glenoid process. It arises from a small plexus of extension of the cavernous sinus. A smaller ve
veins which lies between the cartilaginous ex nous channel lies in the laterally adjacent and
ternal acoustic process and the ventral part of parallel carotid canal. It connects the same par
the squamous temporal bone. The plexus re ent sinuses as does the ventral petrosal; this
ceives one or two tributaries from the muscula venous channel contains the internal carotid
ture of the cranial part of the neck. artery.
The sigmoid sinus (sinus sigmoideus) (Fig. 5 - The paired cavernous sinus (sinus cavemosus)
18) is the roughly S-shaped posteroventral con (Fig. 5-18) plays a key role in the ventral vena
tinuation of the transverse sinus. It begins by tion of the brain. The right and left sinuses lie on
forming an arc around the proximal end of the the respective sides of the floor of the middle
petrous temporal bone. The first arc is continued cranial fossa and extend from the orbital foram
by the second arc, which lies medial to the petro- ina to the petrobasilar canals. Anteriorly, each
occipital synchondrosis. The sinus terminates communicates through the orbital foramen with
after traversing the jugular foramen by continu the orbital plexus of veins. Laterally, each gives
ing as the internal jugular vein. At this junction off emissary veins which run through the round,
the ventral petrosal sinus enters the venous oval, and the external carotid foramina to enter
channel from in front and the vertebral vein the internal maxillary vein. Posteriorly, each is
leaves from behind. The sigmoid sinus receives continued by the ventral petrosal sinus and indi
one or two delicate meningeal veins from the rectly is connected with the vertebral venous
medulla oblongata. Its largest connection is with sinus. Laterally, the middle meningeal vein, a
the condyloid vein (v. condyloidea). From the satellite of the middle meningeal artery, enters
junction of the two arcs forming the sigmoid the cavernous sinus opposite the round foramen.
sinus the condyloid vein passes through the The two cavernous sinuses are connected medi
condyloid canal, and continues as the ventral ally, by means of the large but short anterior and
occipital sinus, which becomes the vertebral posterior intercavernous sinuses (sinus inter-
venous sinus. Pilcher (1930), by means of dye in cavernosi), in front of and behind the stalk of the
jections into the sagittal sinus, demonstrated dorsum sellae. The expanded dorsal part of the
that the sagittal, transverse, sigmoid, and verte dorsum sellae covers the middle portion of
bral sinus system is the main venous drainage the usually larger anterior intercavernous sinus,
from the brain. Occasionally there is an osseous which lies directly posterior to the hypophysis.
canal between the condyloid canal and the hy The usually smaller posterior intercavernous
poglossal foramen. This canal conducts the vein sinus runs across the caudal surface of the base
o f the hypoglossal canal (v. canalis hypoglossi), of the dorsum sellae and is only about 2 mm.
which extends from the condyloid vein to the long. It unites the right and left cavernous si
dorsal surface of the initial part of the vertebral nuses where they lie closest together. This sinus
vein in the petrobasilar fissure. may be absent. A third delicate intercavernous
The dorsal petrosal sinus (sinus petrosus connection may exist anterior to the hypophysis.
dorsalis) (Fig. 5-16) is a posterior extension, be The cavernous sinus contains, free in its lumen,
ginning at the free border of the tentorium cere- the anterior portion of the middle meningeal
belli opposite the distal end of the petrosal crest, artery and the anastomotic ramus of the external
of the basal vein of the cerebrum and the vein ophthalmic artery. The cavernous sinuses con
which runs with the trigeminal nerve (Zimmer tain no trabeculae, but a few stabilizing threads
mann 1936). The sinus almost at once comes to attach to the arteries as they perforate the wall
lie on the lateral surface of the pyramid, which it of the sinus (Zimmermann 1936).
grooves. About 6 mm. from its junction with the The ventral occipital sinus (sinus occipitalis
transverse sinus it receives the posterior cerebral ventralis) (Fig. 5-16) is the venous link between
vein. This vein is 1 mm. in diameter and is as the condyloid vein and the vertebral sinus. It is
large as the sinus it enters. The confluence of the about as wide as it is long, measuring approxi
dorsal petrosal and transverse sinuses forms an mately 5 mm. in each direction. It is transversely
422 C h apter 5 . The V e n o u s S y s te m
compressed and converges toward the opposite of the frontal lobe. Usually the largest and long
sinus as it lies on the medial surface of the lateral est dorsal cerebral vein arises in and on the gyri
part of the occipital bone. The occipital sinus dorsal to the rhinal fissure by anastomoses with
becomes the vertebral sinus as it leaves the fora the posterior and ventral cerebral veins. It runs
men magnum to run across the medial surface of over the coronal and posterior sigmoid gyri to
the lateral mass of the atlas. It is always con reach the cruciate sulcus. It collects many tribu
nected to its fellow sinus ventrally by the trans taries from neighboring gyri and sulci along its
versely running, flat ventral interoccipital sinus, course. The last tributary to enter it is the parie
and there may be a dorsal connection also. The tal diploic vein just before it enters the sagittal
posterior part of the medulla lies between the sinus from the cruciate sulcus. One or two small
right and the left occipital sinus. dorsal cerebral veins enter the posterior half of
The ventral interoccipital sinus (sinus interoc- the dorsal sagittal sinus from each hemisphere.
cipitalis ventralis) (Fig. 5-18) is a stout, flat The posterior cerebral vein (v. cerebri poste
transverse venous passage which connects the rior) (Figs. 5-17, 5-18) drains most of the cortex
right and left ventral occipital sinuses just inside of the temporal lobe. One branch arises from the
the foramen magnum. It lies on the floor and ad dorsolateral portion of the lobe, and the other
jacent sides of the occipital bone. Its anterior branch comes from the piriform area. This
border is irregular, as it apparently sends small branch forms a groove on the lateral surface of
finger-like processes between the two layers of the pyramid, as it runs dorsoposteriorly to unite
the dura. with the more dorsal branch before entering the
The dorsal interoccipital sinus (sinus interoc- dorsal petrosal sinus. Sometimes this union fails,
cipitalis dorsalis) may be absent. It is a dorsal and the branches terminate independently in
transverse channel or plexus which unites the the dorsal petrosal sinus.
right and left ventral occipital sinuses. It may be The great cerebral vein (v. cerebri magna)
coextensive with the first dorsal internal verte (Fig. 5-16) is the sole channel for return of ve
bral venous plexus which lies under the cranial nous blood from all the deep or central gangli
lip of the arch of the atlas. These plexuses, which onic veins of the cerebrum. It begins by the con
are lateral to the junction of the brain stem and fluence of the dorsally located vein of the corpus
spinal cord, might be entered in cisternal punc callosum and the ventrally located internal cere
tures at the atlanto-occipital joint. bral veins and the thalamostriate vein. It runs in
the triangle formed by the cerebral hemispheres
Veins of the Brain and the vermis of the cerebellum caudally. At its
termination it receives a tributary from the oc
The veins of the brain (venae cerebri), like the cipital lobe. Bedford (1934) experimentally pro
dural sinuses into which they drain, do not con duced occlusions of the great cerebral vein in
tain valves, and their walls contain no muscular the dog and noted the rapid establishment of a
coat. They empty into the sinuses in a direction collateral circulation.
usually opposite to the flow of blood in the si The vein of the corpus callosum (v. corporis
nuses. Those from the cerebral hemispheres may callosi) (Fig. 5-17), unpaired, begins by collect
be divided into cortical and central veins. The ing a small tributary from the medial surface of
cortical veins may be divided further into dorsal, each hemisphere, anterior to the corpus callo
and posterior cerebral veins. The central veins, sum. It runs posteriorly, dorsal to the corpus cal
which drain into the great cerebral vein, are the losum, where it is connected to the free margin
corpus callosal, basal, internal cerebral, and of the falx cerebri by arachnoid. During this
thalamostriate veins. The cerebellar veins are course it receives minute tributaries from the
divided into dorsal and ventral veins. The veins medial surfaces of the hemispheres. At the sple-
of the brain stem are the medullary and pontine nium it receives the single choroidal vein (v.
veins. choroidea), which is a posterior continuation of
The dorsal cerebral veins (vv. cerebri dor the choroid plexus of the third ventricle.
sales) (Fig. 5-16) are paired but are not bilater The thalamostriate vein (v. thalamostriata) is
ally symmetrical. All enter the sagittal sinus. another tributary which enters the posterior end
They drain the cortex of nearly the whole cere of the vein of the corpus callosum. It arises in
brum. Although for the most part they lie in the the thalamus and corpus striatum.
sulci, they often run across the gyri. From one to The internal cerebral veins (vv. cerebri inter-
four dorsal cerebral veins enter the anterior half nae) (Fig. 5-17) are present in the dog as a clus
of the sagittal sinus. These come from the cortex ter of tributaries from the dorsal midbrain. Dor-
V e in s of th e C en tral N ervo u s S y stem 423
„ F o r a m e n ro tu n d u m
Ant. I n t e r c a v e r n o u s s i n u s
Cavernous s in u s
- Ventral p e t r o s a l sin us
Post, c e r e b r a l v - -
in p e t r o b a s i l a r c a n a l
D o r s a l p e t r o s a l s in u s ^ — 'Temporal sin u s
" - - J u g u l a r fo ram en
J u g u l a r foram en " S ig m oid sin u s
O ccipital e m i s s a r y v.
S i g m o i d sin u :
^ C o n d y l o id v. in c a n a l
T r a n s v e r s e sii
npu x Ventral o c c i p i t a l s in u s
D orsal s a g i t t a l s i n u s ( cut ) / \ '
sal to the mesencephalon, right and left vessels of the cranium and the caudal part of the frontal
anastomose with each other, as well as with the sinus. It runs anterolaterally, and leaves the skull
basal vein on the respective side. by the small frontal foramen to enter the angular
The cerebellar veins, like the cerebral veins, vein of the eye ventral to the zygomatic (supra
lie in the pia and tend to follow the sulci. They orbital) process. Posteriorly it anastomoses with
are divided into dorsal and ventral sets. either the anterior part of the sagittal sinus or a
The dorsal cerebellar veins (vv. cerebelli dor- large dorsal cerebral vein by a single or a double
sales) are right and left vessels which drain into vessel.
the right and left transverse sinuses by one or The parietal diploic vein (v. diploica parie-
two stems adjacent to the confluence of the si talis) arises in the diploe of the medial part of the
nuses. They arise and lie one on either side in parietal bone 1 to 2 cm. from the mid line by
the fissure between the two lateral hemispheres anastomosing with the occipital diploic vein or
and the central vermis. The numerous fine trib veins. It terminates in the mid-dorsal part of the
utaries which enter them lie mostly in the fine sagittal sinus or in the adjacent large dorsal
sulci between the thin, closely packed gyri. Two cerebral vein. When single, it lies in the diploe
or more venous threads run across the gyri of the of the cerebral juga opposite the ectolateral
vermis from origins in the caudal colliculi. sulcus. When it is double, the two parts are con
The ventral cerebellar veins (w. cerebelli nected by a dense rete.
ventrales) are one or two minute vessels, on The occipital diploic vein (v. diploica occipi
each side, which lie between the lateral hemi talis) arises from an anastomosis with the parietal
spheres and the medulla. They collect veins diploic vein and enters the transverse sinus
from the brain stem primarily, but also from the about 2 cm. from the mid line. It is frequently
cerebral hemispheres, and drain into the sigmoid double or triple; when it is triple the second
or occipital sinuses. vessel enters the sinus close to the mid line, and
The medullary and pontine veins lie on the the third enters the proximal part of the middle
ventral and lateral surfaces of the medulla ob third. Zimmermann (1936) illustrates a small
longata and pons. Those from the pons are trans diploic vein in the interparietal process which
verse, whereas the medullary veins lie lateral to enters the confluence of the sinuses.
the pyramids. They are about 0.2 mm. in di
ameter and lie about 4 mm. from the basilar
Meningeal Veins
artery. They are moderately sinuous and collect
many fine twigs from each side of the ventral
The meningeal veins (vv. meningeae) lie in
median fissure. The lateral tributaries may also
and drain the dura mater. The anterior menin
come from a longitudinal vessel which crosses
geal vein is a delicate vessel which begins in the
the olive. It receives fine tributaries from the
dura covering the frontal lobe. It frequently
cerebellar hemispheres. The medullary veins
leaves a faint shallow groove on the cerebral sur
empty laterally into the occipital sinuses. The
face of the frontal bone. It perforates the inner
pontine veins, one on each side, arise mid-ven-
table of the frontal bone and joins the frontal
trally and run laterally over the pons or between
diploic vein in the region of the ethmoidal fossa.
the pons and the trapezoid body to reach the
transverse fissure. They drain into the sigmoid The middle meningeal vein (Fig. 5-19) is more
sinus. extensive in its ramifications within the dura.
After the major tributaries converge over the
lateral surface of the brain, the vein courses ven-
Veins of the Diploe troanteriorly along the petrous temporal bone.
Within the cranium, at the foramen ovale, it
The diploic veins (w. diploicae) (Fig. 5-19) joins a venous sinus which connects the cavern
are present only in those places where there is ous sinus with the internal maxillary vein.
cancellous or spongy bone (diploe) uniting the
two tables of the skull. There are thus no diploic
veins in the lateral wall of the brain case, where Veins of the Spinal Cord and Vertebrae
the two tables are fused, or anteriorly, where the
two tables of the frontal bone are widely sepa The vertebral vein system constitutes an alter
rated to form the frontal air sinuses. The dog has nate route for the return of blood from the body
frontal, parietal, and occipital diploic veins. to the heart, via anastomoses with anterior sys
T he fron tal diploic vein (v. diploica frontalis) temic veins and the azygos vein, which in effect
drains into the angular vein of the eye from the bypasses the caval system. The vertebral sinuses
diploe located in the triangle between the roof are in direct communication with the cranial
V e in s of th e C en tral N ervo u s Sy st em 425
F r o n t a l d i p l o i c v.
D o r s a l c e r e b r a l v.
- - D o r s a l s a g i t t a l s in u s
- - F r o n t a l d i p l o i c v.
F ron tal d i p l o i c v.
D o r s a l c e r e b r a l v.~ ■Parietal d i p l o i c vv.
M id d le m e n in g e a l v- -
P a r i e t a l e m i s s a r y v.
- - T r a n s v e r s e s in u s
O c c ip it a l d i p l o i c vv. -
I n t r a o s s e o u s p a r t of
D orsal c e r e b e lla r t r a n s v e r s e s in u s
venous sinuses and, since no valves exist in connect with the lumbar veins by means of the
either, blood may flow cranially or caudally, de ventral venous plexuses. The sacral and coccyg
pending on pressure relations. Batson (1940, eal vertebrae usually have no basivertebral veins.
1957), in discussing the concept of the vertebral The intervertebral veins (w. intervertebrales)
vein system and its active physiological role, are present at every intervertebral foramen, pro
makes reference to anatomical and clinical ob viding communication between the vertebral
servations in both man and animals. Drager sinuses and extravertebral veins. The first few
(1937), Worthman (1956), and Reinhard et al. are single on each side, but most are double
(1962) have described and illustrated the verte with one part lying in the caudal and the other
bral sinuses in the dog. in the cranial notch of the contiguous vertebrae.
The vertebral venous sinuses (sinus venosi When they are double, the emerging roots of
vertebrales) (Figs. 5-20, 5-21, 5-22) are paired, the spinal nerve he between them, or may be
thin-walled, flattened, valveless vessels which ringed at the intervertebral foramen by dorsal
extend from the skull to the coccygeal vertebrae. and ventral anastomoses between the double
They lie on the floor of the vertebral canal in the veins. In this way, a venous cushion surrounds
epidural fat. As paired trunks coursing through the nerve roots at their union. This may in
the vertebral canal they diverge from each other clude the dorsal root ganglion. Even when the
at the intervertebral foramina and approach intervertebral veins are single, they may arise as
each other over the vertebral bodies. They are paired veins. This arrangement is regularly found
largest in the cervical region. Within the arch of in the caudal few segments of both the cervical
the atlas, where they originate as continuations and the thoracic region. The intervertebral veins
of the ventral occipital sinuses, they may appear take the names and numbers of the interverte
ampullated (Worthman 1956). Their diameter is bral spaces through which they pass, except for
reduced at the junction of the last cervical and the first two sacral intervertebral veins, which
first thoracic vertebrae, and remains constant pass through the two ventral sacral foramina on
from there to the level of the fourth or fifth lum each side. They have major extravertebral
bar vertebra. Caudal to this level the vertebral si anastomoses as follows;
nuses decrease in size, and they may fuse within Cervical I though VIII, with the vertebral vein.
the fourth to sixth coccygeal vertebrae, or termi Thoracic I, II, and III, with the costocervical
nate as fine venules in the tail musculature. and supreme intercostal vein.
Along the course of the vertebral sinuses there Thoracic IV (left side), about 50 per cent with
are frequent anastomoses between the right and the supreme intercostal vein.
left channels. Some of these anastomoses are Thoracic IV or V (left side) through thoracic IX
superficial, whereas others are beneath the dor or X, with the azygos vein.
sal longitudinal ligament or within the vertebral Thoracic IX or X (left side) through thoracic
body. XIII, with the hemiazygos vein.
Within the vertebral canal the vertebral Thoracic IV (right side) through lumbar III, with
sinuses receive the spinal veins (vv. spinales), the azygos vein.
which follow the nerve roots to the interverte Lumbar IV (V) and V (VI), with the postcava.
bral region on each side. Lumbar VI and VII, with the internal iliac or
The basivertebral veins (vv. basivertebrales) directly into common iliac or the postcava.
are usually paired tributaries which arise within Lumbar VII with the internal iliac.
the vertebral bodies or from the soft tissues ven Sacral I (II), with cranial gluteal vein.
tral to the vertebrae or from anastomoses with Sacral II (III), with internal pudendal vein.
paravertebral veins. They ascend through osse Coccygeal I through IV, with the middle sacral
ous canals in the vertebral bodies and join the and the internal iliac vein.
longitudinal vertebral sinuses. In the cervical In the thoracic and lumbar regions the inter
region they begin in the ventral vertebral venous vertebral veins empty into the intercostal vein
plexuses by an anastomosis with muscular tribu before joining the larger channels.
taries of the vertebral veins within the longus The arcuate veins (vv. arcuata) of Ellenberger
colli muscle. In some cranial segments of the and Baum (1943) and Reinhard, Miller, and
thoracic region no basivertebral veins are evi Evans (1962) are called interarcuate veins by
dent; more caudally, single basivertebral veins, Drager (1937) and Worthman (1956). As the
at their beginnings, anastomose with intercostal main components of the internal vertebral
veins. In the lumbar region the basivertebral plexus they are most prominent in the cervical
veins are largest. They are usually paired and and thoracic regions. They arise in the epaxial
V e in s of th f C en tral N erv o u s System 427
Vv of ext. vertebral
Ii in tervertebral v III
'Exit o f cervical n V II 'R v e rte b ra l sinus
V ertebral v. V e r t e b r a l v.
L. v e n t r a l o c c i p i t a l sinus
Fic, 5-20 Cervical vertebral veins, nght lateral aspect. (From Reinhard. Miller, and Evans 1962.)
Thoracic v erte b ra I
A rcu a te v I
Thoracic v erteb ra XIII ivhich receives V. o f dorsal
Intcrspmous v. Vv. of aorsal e x t vert, plexus
L. vertebral sinus int. vent plexus
fiasivertcbral V
In te rc o s ta l v XII' ’AzLjCjos v C o s to c e rv ic o l v
Laterol tr a n s v e r s e branch1 Intervertebral v., V e r t e b r a l v.
th o racic VI
R. v e r t e b r a l s in u i ,
S a c ru m I I lu m b a r v e r te b rc ^
Coccygeal vein'
I ■ ' P o s tc a v a J Intenvert. v.t lumbar /
i 1 i
ft in te rn a l iha c vein1 1
iL
1 common iliac vein
1Anastomotic branch,
Middle s a c r a l vein R ex te rn a ! iliac vein postcava to azygos v.
Fic 5-22 Lumbar, sacral and coccygeal vertebral veins right lateral aspect (From Reinhard, Miller, and I.vans 1962,1
430
L y m p h o id T is s u e 431
after ligating and resecting 3 to 5 mm. of the dilated portion of the lymph channel receives the
large lymphatic trunks in the leg of the dog. intestinal trunk, which runs dorsally from the
Since lymphatics contain numerous valves, abdominal viscera. It is formed by the lumbar
they are difficult to inject in a retrograde direc lymph trunks, which are the forward continua
tion. The valves usually possess two cusps, but tions of the lymph vessels from the pelvic limbs,
they may consist of a single flap. They may best via the efferents from the iliac lymph nodes.
be observed by producing congestion after liga The exact origin of the thoracic duct is some
tion, by injecting dye particles peripherally, or what arbitrarily assigned, as the morphology of
in edematous material. Prier, Schaffer, and Skel- the cisterna chyli is so erratic. Typically, the cis
ley (1962) have performed direct lymphangiog terna chyli is an elongated sac, dilated in the
raphy in the dog, using a radiopaque medium middle and constricted at both ends, which lies
injected into metatarsal lymphatics. Clinical ap dorsodextral to the aorta from the fourth to the
plications of lymphography include identifica first lumbar vertebra. The thoracic duct is con
tion of lymph node lesions, evaluation of lym sidered to begin between the crura of the dia
phatic blockage, and visualization of the prog phragm where the cisterna attains its minimum
ress of therapy on lymphatic lesions. (Fischer width. It runs cranially from a point opposite
1959; Fischer and Zimmerman 1959). the first lumbar vertebra on the right dorsal bor
Research on the anatomy and physiology of der of the thoracic aorta and the ventral border
the lymphatic system has resulted in a consid of the azygos vein to the sixth thoracic vertebra.
erable fund of knowledge, although many ques Here it inclines to the left, running between the
tions remain unanswered. Drinker (1942) com azygos vein and the aorta, and then obliquely
mented on the observations of Aselli (in 1627) crosses the ventral surface of the fifth thoracic
and of Pecquet (in 1651) on the lymph vessels vertebra to enter the precardial mediastinum. It
and nodes in the dog. Aselli described and illus runs cranioventrally in the precardial mediasti
trated the mesenteric lacteals and mesenteric nal septum, and in about one-half of the speci
lymph node (“pancreas of Aselli”); Pecquet de mens it terminates singly at the junction of the
scribed the receptaculum chyli (“cistern of left external jugular vein with the precava
Pecquet”) and the thoracic duct, including its (Baum 1918). In many specimens the cranial
termination. Rusznyak, Foldi, and Szabo (1960) portion of the thoracic duct bifurcates or bifur
credit Rudbeck (in 1652) and Bartholinus (in cates. These branches are usually connected by
1653) for having recognized the lymphatic sys cross branches so that a coarse, widespread
tem as an entity. The English edition of the plexus is formed. The termination of the end
monograph on lymphatics and lymph circula parts of the thoracic duct varies as to both the
tion by Rusznyak, Foldi, and Szabo (1960), re veins they empty into, and the places on the ves
vised and enlarged from the Hungarian, Ger sels at which they empty. When the thoracic
man, and Russian editions, contains an extensive duct is single, it usually presents a terminal dila
bibliography (over 1700 citations). Frequent ref tion immediately before it becomes constricted
erence is also made therein to the morphology to perforate the venous wall. Similar ampulla
and experimental physiology of the lymphatic like dilations may be present on each of the
system in the dog. A classical topographical de branches when the thoracic duct terminates in
scription of the lymph nodes and vessels in the multiple vessels. Freeman (1942), who examined
dog was written by Hermann Baum in 1918. the termination of the thoracic duct in 25 dogs,
The illustrations used in this chapter have been found that it had no branches in 3 and was di
reproduced from that work, “Das Lymphgefass- vided in 5, and that it sent branches to the right
system des Hundes,” through the kind permis side in 8 , to the azygos vein in 5, and to both the
sion of Springer Verlag, Berlin. right side and the azygos vein in 4.
The thoracic duct (ductus thoracicus) (Fig. 6 - Lymphoid tissue is present in all classes of
5 is the chief channel for return of the lymph of vertebrates, but reaches its highest development
the body. Lymph from the right thoracic limb in mammals, in which circumscribed organs, the
and shoulder and the right side of the neck and lymph nodes, occur along the lymph vessels.
head is returned via the right lymphatic duct. The spleen, thymus, and bone marrow also con
The thoracic duct begins in the sublumbar re tain lymphoid tissue, although bone marrow
gion, or between the crura of the diaphragm as a functions primarily in forming erythrocytes, and
cranial continuation of the cisterna chyli. This leukocytes rather than lymphocytes. Yoffey and
432 Chapter 6. T he L y m p h a t ic System
L y m p h o id T is s u e 433
Lymph Nodes
%
31 S
ICl The lymph node (nodus lymphaticus) is the
o
% -o structural and functional unit of the lymphatic
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xia xia system. It serves two important functions. It acts
8 So b
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£■oJG ®
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c S £
ay SC a °_
■=Rr~" cuCu 1 I f ! ja white blood cells. Lymph nodes are located in
Ir -3
'cS fe
£ d aj d . „ ® those places where they are afforded maximum
’S %2 ^ a £ £ Z £ i £ ’SZ £ n ”1 protection yet produce minimal interference
with the functioning of the skeletal, muscular,
f i l l si &■! j-gj Jb ^C£u.s~v a% «
ij'E fc-tsi si f
-n S v o la j - 5- cSs - -S a _^j ^ *- o -1_- — 1—£■5 ► and blood vascular systems. They are thus found
!=■ S ' l. l* -l0 s< e' 0l <se ^ . ooX Oi l< S • 'S O •& in the small fat storehouses at the flexor angles of
cb u i c e 1 "+H
Cu
I ^"- " s t a j a j * i- l* lB n 2i il l. l
rt
Oh CQ joints, in the mediastinum and mesentery, and
i O] CO ' ' t l O CD in the angles formed by the origin of many of the
larger blood vessels. Each node consists of a cap
sule, containing elastic and smooth muscle fibers
(Trautmann and Fiebiger 1952), and an internal
434 Chapter 6. T he L y m p h a t ic System
framework consisting of septa and trabeculae. of the head. The tonsillar ring of lymph tissue is
There is a convex surface, and a small flat or described with the pharynx.
concave area, the hilus, which is usually not The parotid lymph node (nodus lymphaticus
prominent. Internally the node contains a poorly parotideus) (Figs. 6-2, 6-3) is a bean-shaped
defined cortex and m edulla. The structural unit node located under the anterodorsal border of
of the lymph node is the lymph nodule. Each the parotid gland on the posterior parts of the
nodule contains light-colored central areas in zygomatic arch and adjacent masseter muscle.
which the lymphocytes are formed. Most nod In a medium-sized dog it is about 1 cm. long,
ules are located in the cortex, where they are 0.5 cm. wide, and 0.3 cm. thick. Occasionally
partly surrounded by the lymph sinus. The a second or even a third node may be present
lymph sinus is a lymph-circulating space located (Baum 1918).
under the capsule and partly along the septa and The afferent lymphatics to the parotid lymph
larger trabeculae. The lymphoid tissue of the node come from the cutaneous area of the pos
medulla is in the form of anastomosing cords of terior half of the dorsum of the muzzle and the
lymphocytes with few nodules. side of the cranium, including the eyelids and
associated glands, the external ear, the tempo
Lymph Follicles romandibular joint, and the parotid gland. Its
two or three efferent ducts run between the
Solitary and aggregated follicles are found digastric muscle and the parotid gland to the
mainly in the wall of the digestive tube. They large retropharyngeal lymph node.
differ from nodes, in that lymph vessels arise in, The mandibular lymph nodes (nodilymphatici
rather than pass through them; thus they con mandibulares) (Figs. 6-2, 6-3) form a group of
tain only efferent lymphatics. The various ton two or three nodes, or rarely as many as five,
sils and the aggregated follicles are described which lie ventral to the angle of the jaw. A flat
with the digestive system. The solitary and small tened three-sided node, with borders about 1
aggregated follicles are particularly abundant in cm. long, lying above the external maxillary vein
the cecum, rectum, anal canal, prepuce, and and a long, ovoid node lying below the external
third eyelid. The lymphoid tissue of the third maxillary vein constitute what is probably the
eyelid is located on its bulbar side. It occasion most common arrangement. The node lying ven
ally becomes infected and hypertrophied, caus tral to the external maxillary vein usually is over
ing follicular conjunctivitis. It may protrude 2 cm. long and about 1 cm. wide. It is flattened
around its free border in the form of a reddened transversely. In more than one third of the speci
tumor (tumefied orbital gland). The lymph folli mens examined by Baum (1918), two or more
cles of the prepuce are commonly infected in old nodes replaced this single node on one or both
male dogs, and an almost continuous purulent sides. In only 16 out of 36 specimens was the
exudate is discharged from the sheath. The sub- grouping of the mandibular nodes on the two
mucous lymph follicles of the large intestine can sides similar. The dorsal node may be double,
be seen clearly in the cadaver under proper light but this condition is rare.
ing conditions if the intestine is inflated. The afferent lymphatics to the mandibular
lymph nodes come from all parts of the head not
REGIONAL ANATOMY OF THE drained by the afferent lymphatics of the parotid
LYMPHATIC SYSTEM node. There is overlapping in the areas of drain
age, so that the eyelids and their glands, and the
The larger lymph vessels and the lymph nodes skin of the dorsum of the cranium and the tem
of the body will be described regionally, under poromandibular joint drain into both nodal sta
the following headings: head and neck, thoracic tions. The efferent lymphatics of the mandibular
limb, thorax, abdominal and pelvic walls, genital lymph nodes go primarily to the ipsilateral
organs, abdominal viscera, and pelvic limb. The medial retropharyngeal lymph node. The nodes
spleen and thymus are separately discussed later. composing the group are connected with each
other, as well as with the contralateral medial
Lymph Nodes and Vessels retropharyngeal lymph node. The 8 to 10 effer
of the Head and Neck ent lymphatics anastomose with each other and
form a plexus as they pass over the pharynx. Be
The parotid lymph node and the mandibular fore reaching the medial retropharyngeal node
groups of lymph nodes are the only lymph nodes they unite to form three to five small trunks
R e g io n a l A n a t o m y of th e L y m p h a t ic Sy st e m 435
which enter the ventrolateral surface of the pole of the ipsilateral medial retropharyngeal
node. lymph node or its efferent ducts.
The medial retropharyngeal lymph node Yoffey and Drinker (1938) state that the
(nodus lymphaticus retropharyngeus) (Figs. 6-2, tracheal duct (their cervical duct) is usually
6-3) is the largest node found in the head and single as it leaves the caudal pole of the node,
neck. It largely and sometimes completely but it may be double or in the form of a plexus.
takes the place of the cranial members of both As it runs down the neck it lies in or adjacent to
the superficial and the deep series of the cervical the lateral wall of the carotid sheath. These in
and the pharyngeal nodes as found in man. It is vestigators found no additional nodes along the
an elongated, transversely compressed node, course of the tracheal duct. The left tracheal
with a more pointed caudal end, and is about 5 duct usually terminates in the thoracic duct. The
cm. long and nearly 2 cm. wide. It lies under the right tracheal duct usually terminates in the
wing of the atlas in the triangle bounded by the angle formed by the merging of the right exter
m. digastricus cranially, the m. longus colli dor nal jugular and the right axillary vein to form the
sally, and the pharynx and larynx ventrally. The brachiocephalic vein, but the termination may be
mastoid parts of both the m. brachiocephalicus to one side of the angle. Baum (1918) states that
and m. sternocephalicus largely cover the node the efferent lymph duct from the large superfi
laterally, although its cranioventral part is related cial cervical nodes doubles the size of the right
to the mandibular gland. Coursing along its tracheal duct as it enters this channel 2 or 3 cm.
medial surface is the terminal portion of the from the first rib in large dogs. The resultant
common carotid artery, as well as the hypo vessel is known as the right lymphatic duct
glossal, vagus, and sympathetic nerves, and the (ductus lymphaticus dexter). It is about 5 mm.
internal jugular vein. wide and 1 cm. long; it empties into the right
In 10 out of 47 specimens Baum found two axillary vein or the angle formed by the merging
medial retropharyngeal lymph nodes present on of this vein with the right external jugular. Com
one or both sides; in one-third of his specimens monly the right lymphatic duct bifurcates and
a lateral retropharyngeal lymph node was pres then reunites, forming a circle before it termi
ent. This node is less than 1 cm. in diameter and nates. Sometimes the forked condition persists,
lies at the dorsal border of the mandibular gland so that the right lymphatic duct enters the ve
in the fat posterior to the cartilaginous external nous system, in front of the first rib, by two chan
acoustic meatus. It is completely or partially nels.
covered by the posterior part of the parotid The superficial cervical lymph nodes (nodi
gland. The afferent lymphatics of these two lymphatici cervicales superficiales) (Fig. 6-4)
small accessory nodes come from the structures usually consist of two nodes, one lying dorsal to
lying adjacent to them. Their efferent vessels the other in the adipose tissue on the serratus
drain into the large medial retropharyngeal ventralis and the scalenus in front of the supra-
lymph node. spinatus. They are covered superficially by the
The afferent lymphatics of the medial retro thin cleidocervicalis, the omotransversarius and,
pharyngeal lymph node come from all the deep at their dorsal end, by the trapezius. The omo
structures of the head which have lymphatics. cervical artery and vein lie medial to the caudal
Thus, the tongue, the walls of the oral, nasal, and parts of the nodes as they course in front of the
pharyngeal passages, the salivary glands, and the shoulder in the groove between the shoulder and
deep parts of the external ear drain into this neck. The more ventral superficial cervical
node. It also receives afferent lymphatics from lymph node may encroach on the trachea on the
the larynx, esophagus, and the noncutaneous, right side, and the esophagus and trachea on the
nonmucous structures of the neck which have left side. Occasionally a single node is present on
lymphatics. The efferent ducts from the parotid each side, but more commonly three or more
and mandibular lymph nodes drain into the me nodes replace the usual two. Most of the nodes
dial retropharyngeal lymph node also. Yoffey and are oval and somewhat flattened. They are col
Drinker (1938) found, lying between the hamulus lectively about 3 cm. long and less than 1 cm.
and the pharyngeal opening of the auditory tube, thick.
four or five lymph vessels which came from the The afferent lymphatics come mainly from
floor and side walls of the nose. The right and the skin of the posterior part of the head, includ
left tracheal ducts (ductus trachealis dexter et ing the pharyngeal region, and a part of the
sinister), 2 to 4 mm. wide, arise from the caudal pinna, the lateral surface of the neck, and the
(Text continued on page 439.)
436
Chapter 6.
The
L y m p h a tic
S y s te m
F ig . 6-2. Lymph vessels of the tongue, the tongue muscles, the soft palate, and the larynx of the dog. (After Baum.)
437
h. M. rectus lateralis (each partially removed) 1. Lymph vessel of the sublingual gland which
i. Lacrimal gland r, f . Sublingual glands goes to a mandibular lymph node (cut)
438
Chapter
6.
The
L y m p h a tic :
S y s te m
Fic.. 6-4. Medial retropharyngeal lymph nodes and cervical lymph nodes of the dog. (After Baum.)
a, a'. Medial retropharyngeal lymph nodes 1. Thyroid gland
b. Cranial cervical lymph node 2. Axillary vein (cut)
c, c'. Caudal cervical lymph nodes 3. External jugular vein (cut)
d, d\ d ". Superficial cervical lymph nodes 4. Internal jugular vein
e. Axillary lymph nodes 5. First rib
e'. Accessory axillary lymph node 6. Trachea
f. Left tracheal duct 7. Esophagus
g. Efferent vessel of the superficial cervical lymph 8. M . serratus ventralis
nodes 9. M. scalenus
i. Thoracic duct with terminal branches 10. M. stemothyroideus
k, k', k ", k '". Lymph vessels of the larynx 11. M. stemohyoideus
1. Lymph vessel which runs to a cranial mediastinal 12. Muscles of the pharynx
lymph node 13. M, longus capibs
m to m Mandibular lymph nodes 14. M. digastricus
n. Efferent vessels of the mandibular lyinph nodes
which go to the medial retropharyngeal lymph
node of the opposite side
R e g io n a l A n a t o m y of the L y m p h a t ic S ystem 439
whole thoracic limb, except a variable region on pharyngeal node. Those from the caudal deep
the medial side of the brachium and antebrachi cervical lymph nodes empty into the right
um, the shoulder and the cranial part of the tho lymphatic duct, or into the thoracic duct on the
racic wall. Mahomer et al. (1927), by making in left. On either side they may empty into the
jections into the thyroid gland of dogs, revealed tracheal duct or into a cranial mediastinal node.
efferent connections with the cervical lymph
nodes, the cervical lymphatic trunks, and with
the veins at the base of the neck. The efferent Lymph Nodes and Vessels of the
ducts connect members of the group when more Thoracic Limb
than one node is present, and as one to three
trunks they descend over the serratus ventralis The axillary lymph node (nodus lymphaticus
and the scalenus to merge with the tracheal duct axillaris) (Fig. 6-4) usually is the only lymph
on the right side to form the right lymphatic node of the thoracic limb. In Baum’s (1918)
duct. On the left side they empty into the tho series of 43 specimens, double nodes were found
racic duct. On either or each side they may in 10 , and in 6 of these the double nodes oc
empty into the external jugular vein directly. curred on both sides. The main axillary lymph
The deep cervical lymph nodes (nodi lymphat- node is usually in the form of a disc about 2 cm.
ici cervicales profundi) (Fig. 6-4) are located in diameter, although the diameter may range
along the cervical portion of the trachea on each from 0.3 to 5 cm. It lies 2 to 5 cm. caudal to the
side. They are exceedingly small, and vary in shoulder joint in the angle formed by the diverg
number. They are customarily divided into ing brachial and subscapular blood vessels. It is
cranial, middle, and caudal nodes. In the dog bounded laterally by the teres major, medially
one or more of these nodes are frequently ab by the transversus thoracis, and ventrally by the
sent. They range considerably in size; they may dorsal border of the deep pectoral muscle. The
be barely visible, or several millimeters long. accessory axillary lymph node (nodus lymphat
The smaller nodes are spherical to ovoid; the icus axillaris accessorius), when present (prob
larger ones are usually elongated and parallel to ably more often than Baum’s figures indicate),
the long axis of the trachea. lies caudal to the principal node in the fascia be
If a cranial deep cervical lymph node is pres tween the adjacent borders of the deep pectoral
ent, it is located between the caudal end of the and latissimus dorsi muscles, caudal to the mus
medial retropharyngeal lymph node and the thy cles of the brachium. It varies in size from less
roid gland. It lies either dorsomedial to the than 1 mm. to 1.5 cm. When it is large the main
thyroid gland along the carotid sheath, or on the node is correspondingly reduced in size. In 4 of
pharynx cranial to the thyroid. It was absent in 29 specimens Baum found a third node in close
19 of Baum’s 64 dissections. The middle deep relation to the main axillary node. In only one of
cervical lymph node was present in only 4 of 50 his specimens was it present on both sides.
specimens Baum examined, and in only 1 of The afferent vessels of the axillary lymph node
these 4 was it present on both sides. It usually or nodes come mainly from the thoracic wall and
lies along the carotid sheath, but it may lie ven the deep structures of the thoracic limb. Those
tral to the trachea in the middle third of the of the thorax extend beyond it so that vessels
neck. The caudal deep cervical lymph node was arise from the deep structures of the neck and
present in 17 of 56 specimens examined by from the abdomen. Both the thoracic and the
Baum, but in only two specimens was it present cranial abdominal mammary glands of each side
bilaterally. In 11 of 17 specimens a single node have lymphatics which drain into the axillary
was located on the ventral surface of the trachea, nodes. An anastomosis between the afferent
and in others there were two or more nodes lying lymphatics of the axillary nodes and those of the
on the ventral surface of the caudal third of the superficial inguinal nodes sometimes occurs be
cervical part of the trachea. tween the cranial and caudal abdominal mam
The afferent lymph vessels to the deep cervi mary glands. The axillary and accessory axillary
cal lymph nodes come from the larynx, thyroid lymph nodes are connected with each other by
gland, trachea, esophagus, and the last five or six lymph vessels. The efferent lymph vessels from
cervical vertebrae. The efferent lymphatics of the axillary nodes of each side course cranially
each cranially located node become a part of the and unite with each other to form one or more
afferent lymphatics of the node located next anastomosing larger trunks which lie on the
caudally. In this way the cranial deep cervical transversus thoracis. The efferent trunks pass
node receives lymphatics from the medial retro medial to the axillary vein, curve around the first
440 Chapter 6. T he L y m p h a t ic System
rib, and empty by one or several branches on the lymphatics from the sternal node on each side
left side into the thoracic duct, left tracheal duct, run in front of the internal thoracic blood ves
left external jugular vein, or into all of these. On sels, where they form a plexus. The right vessels
the right side the axillary efferent lymphatics terminate in the right lymphatic duct, the left in
empty into the right tracheal duct, the right the thoracic duct. Many variations exist.
lymphatic duct, the right external jugular vein, The intercostal lymph node (nodus lym
or into all three of these. phaticus intercostalis) (Figs. 6 -5 to 6-7) was
found in only 14 of 54 specimens by Baum
Lymph Nodes and Vessels of the Thorax (1918), and in only 2 of these was it present on
both sides. This small spherical node lies in the
The lymph nodes of the thorax may be di vertebral end of either the fifth or the sixth inter
vided into parietal and visceral groups. The costal space under the sympathetic trunk, cau
parietal group includes the sternal and inter dal to the intercostal artery. According to Baum
costal nodes; the visceral group includes the it receives a portion of those lymph vessels which
mediastinal and tracheobronchial nodes. The pass into the thoracic cavity through the last six
parietal nodes are smaller and less constant in to eight intercostal spaces. It probably also re
number and location than are the visceral nodes. ceives lymphatics from the ribs, vertebrae,
The sternal lymph node (nodus lymphaticus pleura, and aorta. These vessels lie on the tho
sternalis) (Figs. 6-5, 6 - 6 ) is usually represented racic vertebrae and longus colli muscle. The ef
by a single node on each side. In others a single ferent vessels go to the mediastinal nodes.
median node, which may be located either right The mediastinal lymph nodes (nodi lym-
or left of the median plane, serves both sides. phatici mediastinales) (Figs. 6 -5 to 6-9) vary in
The node may be lacking completely, or, in rare number and shape. Most of them are associated
instances, a double node may be present on one with the large vessels of the heart which run
side. through the dorsal part of the precardial medias
When one node is present on each side, it lies tinum. Unlike most other animals, the medias
immediately cranial to the transversus thoracis tinal lymph nodes in the dog are confined to the
muscle and medial to the second costal cartilage precardial mediastinum or surface of the heart.
or second interchondral space cranioventral to Although they lie in the precardial mediastinal
the internal thoracic blood vessels. Typically, septum, most nodes are not visible through the
the node is ellipsoidal in shape and 2 mm. to 2 pleura from both sides, even in emaciated speci
cm. in length. If a single node is present it may mens. For this reason right and left nodes are
be dumbbell-shaped. When two nodes are pres described. In young animals some of these nodes
ent on one side they may lie close together and are partly embedded in the thymus.
be mistaken for a single node. On the left side the mediastinal lymph nodes
The afferent lymphatics of the sternal node on vary in number from one to six and in length
each side lie under the transversus thoracis in from less than 1 mm. to 3 cm. Most of these
the fat lying between this muscle and the dorsal nodes are oblong, and they lie along the precava
surfaces of the sternal ends of the costal carti and the brachiocephalic, left subclavian, and
lages. Occasionally a single vessel is present. It costocervical arteries. When several nodes are
arises in the abdominal wall, perforates the dia present on the left side, one of these is located
phragm near the middle of the costal arch, and, opposite the first intercostal space either cranial
running under the pleura in the phrenicocostal or caudal to the left costocervical vein. If only a
sinus, extends forward and downward to dip single node exists, it is always located cranial to
under the transversus thoracis. According to the costocervical vein. Small kernel-like nodes
Baum (1918), it receives tributaries from the lying between the dorsally lying left subclavian
ribs, sternum, serous membranes, thymus, adja and the ventrally lying brachiocephalic artery,
cent muscles, and mammary glands. Stalker and or in the left groove between the trachea and
Schlotthauer (1936) state that lymphatics were esophagus, are not apparent unless they are ex
not found to penetrate the thoracic or abdominal posed by removal of the overlying pleura and fat.
walls. Clinical observations indicate that the On the right side the mediastinal lymph nodes
sternal lymph nodes receive no afferent vessels usually number two or three, with a maximum of
from the mammary glands. In the absence of the six. The disc-shaped node lying between the
sternal lymph nodes the afferent vessels which right costocervical vein and the precava is most
would otherwise drain into them drain into the constant. It may be double. In large dogs it
mediastinal nodes. The one to three efferent measures over 1 cm. in diameter and may partly
R eg io n a l
A natomy
of
the
L y m p h a t ic
S ystem
F ig. 6-5. Lymph vessels of the mediastinum, pericardium, diaphragm, aorta, and esophagus of the dog; the left lung is completely removed, the left
wall of the thorax is almost completely removed, as is the M. transversus thoracis. (After Baum.)
1. First rib 12. Twelfth rib d. d'. Lymph vessels which run to the cranial
2. M. longus colli 13. Thirteenth rib lumbar aortic lymph nodes
3 , 3 \ 3 Z. Aorta 14. Costocervical vein e. Lymph vessels which enter the abdominal cav
4. Precardial mediastinum 15,1 5 '. Thoracic duct ity through the diaphragm and em pty into the
5. Pericardium and cardial mediastinum 16. Union of the axillary and jugular veins with the splenic, gastric, left portal, or cranial lumbar
6 ,6 ', 6\ Postcardial mediastinum subclavian vein aortic* lymph nodes
7. 7 ', 7 s. Diaphragm a. a 1, a2. Cranial mediastinal lymph nodes f. Lymph vessel which enters the abdominal cav
8. Esophagus b. Left tracheobronchial lymph node ity with the esophagus
9. Precava b'. Middle tracheobronchial lymph node g. Intercostal lymph node
441
10. Brachiocephalic artery c. Sternal lymph node h. An efferent vessel which goes to the right side
11. Left subclavian artery and appears as numl>er 10 on Fig. 6 -6 .
442
Chapter 6.
The
L y m p h a tic .
S y s te m
Fic 6-6. Right side ot thf thoracic cavity of the dog. (After Baum.)
a. Left ventricle q. Aorta 4 Lymph vessels of the esophagus which enter the
h. Right ventricle r. Trachea abdominal cavity
c. Right auricle s. Right main bronchus 5. Lymph vessels of the esophagus which, turn to the
cl, d'. Coronary sulcus s'. Right eparterial bronchus left and empty into the left tracheobronchial
e. Right longitudinal sulcus t, Esophagus lymph node
f. Postcava u. M. longus colli 6. Sternal lymph node
g. Precava v. Left M. transversus thoracis 7. Lymph vessels which empty into the gastric,
h. Azygos vein w, w 1, w\ Pars costalis, pars lumbal is. and tendinous splenic, portal, or cranial lumbar lymph node
i. External jugular vein parts of the diaphragm 8. Lymph vessels which go to the cranial lumbar
i'. Internal jugular vein x. Sternum lymph node
k. Internal thoracic artery and vein y, y'. Dorsal and ventral piece of the first rib 9. Intercostal lymph node
1. Right sulxdavian artery z. Right M. transversus thoracis (cut) 10. Efferent vessel of a left cranial mediastinal lymph
m. Right axillary artery and vein node
n. Right costocervical vein 11. Thoracic duct
o. Right vertebral artery 1 ,2. Middle and right tracheobronchial lymph node 12. Right tracheal duct
p. Right common carotid artery 3 to 35. Cranial mediastinal lymph nodes
R e g io n a l A n a t o m y of th e L y m p h a t ic System 443
cover both veins between which it lies. Addi to the esophagus dorsally. Its apex fits snugly
tional nodes are frequently found along the dor into the angle formed by the bifurcation of the
solateral surface of the trachea, between the trachea. The vagi lie in contact with the limbs of
costocervical and azygos veins. A node may lie the node as they become intimately related to
between the precava and the brachiocephalic the ventrolateral surfaces of the esophagus.
artery ventral to the trachea. Sometimes the tracheobronchial group includes
The afferent lymphatics, according to Baum a fourth node just cranial to the right node in the
(1918), come from the muscles of the neck, tho angle formed by the azygos vein at its entrance
rax, and abdomen, the scapula, the last six cervi into the precava. In other specimens the middle
cal vertebrae, the thoracic vertebrae, ribs, tra node and either the right or left node form one
chea, esophagus, thyroid, thymus, mediastinum, confluent mass.
costal pleura, heart, and aorta. Lymphatics do The bronchopulmonary lymph nodes (nodi
not invade the central nervous system or the lymphatici bronchopulmonales) are often absent.
contractile elements of skeletal muscles. The They were present on one side in only 14 of 41
mediastinal nodes receive efferent vessels from specimens examined by Baum (1918), and were
the intercostal, sternal, middle and caudal deep never present on both sides. They are small
cervical, tracheobronchial, and bronchopulmo nodes which lie on the dorsal surfaces of the pri
nary nodes. The efferent lymphatics of all nodes mary bronchi between the peripheral ends of
caudal to the relatively constant node located in the right and left tracheobronchial nodes and
front of the costocervical vein on each side drain the parenchyma of the lungs. They receive drain
into it. From this node on the left side arise effer age from the lungs and their efferent vessels go
ent lymphatics which empty into either the tho to the tracheobronchial nodes.
racic duct or the left tracheal duct, or into both. The afferent vessels to the tracheobronchial
On the right side similar efferent vessels go to lymph nodes come from the lungs and bronchi
the right lymphatic duct or the right tracheal primarily, but also from the thoracic parts of the
duct, or to both. aorta, esophagus, and trachea, and from the
The tracheobronchial (bronchial) lymph heart, mediastinum, and diaphragm. The two to
nodes (nodi lymphatici tracheobronchiales) four efferent lymphatics from each tracheo
(Figs. 6-5, 6 - 6 , 6 - 8 , 6-9) include all nodes bronchial node go pardy to another node of this
which lie on the initial parts of the bronchi at group and partly to the mediastinal nodes. Dogs
the bifurcation of the trachea. The nodes, which which have lived in dusty or smoky environ
are constantly present, are known as the right, ments have deeply pigmented tracheobronchial
left, and middle tracheobronchial lymph nodes. nodes. Inhaled foreign materials, regardless of
The right and left tracheobronchial lymph their nature, are filtered out of the lymph as it
nodes are similar in size and location. Each lies passes through these nodes; this accounts for the
on the lateral side of its respective bronchus, but markedly dark color of both the tracheobronchial
also on the trachea, to a small extent. Dorsally, and bronchopulmonary nodes in some speci
the right node is located ventral to the azygos mens.
vein; the left node has a similar relation to the Kubik, Vizkelety, and Balint (1956) have de
beginning of the thoracic aorta. These nodes are scribed the lymphatic drainage into each of the
0.5 to 3 cm. long and are ellipsoidal in shape, several lymph nodes located near the hilus of the
with truncated caudal extremities. The left node lung. Kubik and Tombol (1958) have investi
is more angular, as it is wedged in the space gated the tracheobronchial nodes through which
bounded medially by the left primary bronchus the lymph from the lung passes in its course to
and trachea, dorsally by the aorta, and ventrally the venous system. Miller (1937) has also de
by the pulmonary vein from the left apical and scribed the lymphatics of the dog’s lung.
cardiac lobes of the lungs.
The m iddle tracheobronchial lym ph node Lymph Nodes and Vessels of the
(nodus lymphaticus tracheobronchialis media) is Abdominal and Pelvic Walls
always the largest node of this group. It is in the
form of a V as it lies in the angle formed by the These nodes, like those of the thorax, can be
origin of the primary bronchi from the trachea. divided into parietal and visceral groups. The
The left limb of the V lies on the dorsal surface parietal group includes the lumbar, iliac, sacral,
of the right pulmonary vein, from the diaphrag and deep inguinal. The visceral group is divided
matic lobe; the right limb lies along the right largely into subgroups which serve specific
cranial surface of this vein. The node is related organs.
444
Chapter
6.
The
L y m p h a tic
S y s te m
F ig . 6 - 7 . Lymph vessels of the pleura of the dog. (After Baum.)
a, a'. Dorsally running lymph vessels; they fonn vari 1 , 1'. Cranial mediastinal lymph nodes 7. M. transversus thoracis icut)
ous small branches which run in part (b) crani- 2. Sternal lymph node 8. 8'. Left and right first ribs
ally to the cranial mediastinal lyinph nodes (1, 3. Esophagus (cut) 9. Ninth rib
1 ) and in part (c) caudally to the cranial lumbar 4. Trachea (cut) 10. Thirteenth rib
lymph node 5. Diaphragm (cut and reflected) 11. Intercostal lymph node
d, d Ventrally running lymph vessels which empty 6. M. longus colli 12. Internal thoracic artery and vein
into the sternal lymph node (2). Som e of them
(d'l first run on the diaphragm
R e g io n a l
A n a to m y
of
the
L y m p h a tic
System
Fi(,. 6-8. Lymph vessels of the lungs and bronchial lymph nodes of the dog. (After Baum.)
a, a a-. Apical, cardiac, and diaphragmatic g. Pulmonary veins 6, &. Subserosal lymph vessels which course
lobes of the left lung along the attachm ent of the pulmonary lig
b, b ‘,b *. Similar lobes of the right lung 1, 2 ,3 . Left, right, and middle tracheobron ament
c. Intermediate lobe chial lymph nodes 7. Subserosal lym ph vessels which pass deeply
d. Trachea 4, 4'. Pulmonary lymph nodes 8 ,8 '. A left and a right mediastinal lymph
e, e'. Left and right main bronchus 5. Subserosal lymph vessels hich pass around node
f. Pulmonary artery and its branches the acute margin to the diaphragmatic sur
face and there pass deeply
445
44 6 Chapter 6. T he L y m p h a t ic Sy stem
The lumbar lymph nodes (nodi lymphatici the dorsal half of the abdomen, the pelvis, and
lumbales) (Figs. 6-10, 6-12 to 6-14) are small the pelvic limb. It also receives afferent vessels
nodes which lie along the aorta and postcava from the genital system and the caudal part of
from the diaphragm to the deep circumflex iliac the digestive and the urinary system, as well as
arteries. Except for a paired node near the dia efferent vessels from the deep and superficial
phragm, they are erratic in their development inguinal, left colic, sacral, and internal iliac
and are often absent. Baum (1918) was able to nodes. The efferent vessels from the external
demonstrate as many as 17 individual nodes in iliac lymph node or nodes drain forward to form
some specimens. Because of their small size and the lumbar lymph trunk or drain into the caudal
their similarity in color to the fat in which they members of the lumbar lymph nodes if these
are embedded, they are easily overlooked. The nodes are present.
most constant in size and position is the paired The internal iliac lymph node (nodus lym
lumbar node, which has somewhat different re phaticus iliacus internus) (hypogastric of Baum)
lations on either side. The left node is occasion (Fig. 6-13) is usually a small paired node which
ally double. It is 1 to 2 cm. long and lies between lies in the angle between the internal iliac and
the left crus of the diaphragm and the left sub the median sacral artery, ventral to the body of
lumbar muscles, dorsal or caudal to the left the seventh lumbar vertebra on the ventral sac
renal artery. Its cranial pole usually touches or rococcygeal muscle. The node is unusually vari
extends dorsal to the left phrenicoabdominal able. There may be three nodes, one behind
artery. The right member of this pair, usually another, on one side, or the nodes may be
smaller than the left, may have its cranial pole double on each side. In six of Baum’s specimens
located dorsal to the right phrenicoabdominal only a single node served both sides.
artery and vein just after these vessels have The afferent lymph vessels come from the
crossed the respective dorsal and ventral sur thigh, pelvis, pelvic viscera, tail, and a portion of
faces of the right adrenal gland. the lumbar region; the efferent lymph vessels
The afferent lymphatics to the lumbar nodes drain into the sacral nodes.
come from the lumbar vertebrae, the adrenal The sacral lymph nodes (nodi lymphatici
glands, and the abdominal portions of the uro sacrales) (Figs. 6-10, 6-13, 6-15, 6-17) are lack
genital system, including the testis of the male. ing about half of the time. They are not sharply
They receive efferent vessels from more caudally differentiated from the internal iliac nodes when
located nodes. Efferent vessels from these nodes more than one iliac node is present. They lie ven
also empty directly into the lumbar lymphatic tral to the body of the sacrum or ventral to the
trunks; those from the more constant, cranially ventral sacrococcygeal muscle. Small nodes,
located nodes drain directly into the cisterna when present, lie on each side of the median sa
chyli. cral artery. Occasionally there is a single small
The external iliac lymph node (nodus lym node, located in the fat ventral to the artery; in
phaticus iliacus externus) (medial iliac of Baum) other dogs one or two nodes are present on one
(Fig. 6-13) consists of a large constant node side. Baum (1918) occasionally found a small
located between the deep circumflex iliac and sacral node located between the piriformis and
the external iliac artery. This is usually single, the sacrococcygeus ventralis muscles, closely as
but it may be double on one or both sides. It is sociated with the internal iliac artery and vein.
about 4 cm. long, 1 cm. wide, and 0.5 cm. thick. The afferent lymph vessels come from the ad
It is irregular in outline, and more often its cra jacent musculature and viscera; the efferent
nial end extends under or over the deep circum vessels go as a plexus to the iliac nodes.
flex iliac vessels rather than behind the external The deep inguinal lymph node (nodus ingui-
iliac artery. It is bounded deeply on the right nalis profundus) (Fig. 6-13) was found by Baum
side by the postcava, which lies dorsal and to (1918) in 18 of 50 dogs examined. A node was
the right of the aorta. Each node lies in the fur present on each side in 5, on the left side in 1,
row between the psoas major and the aorta and and on the right side in 12. It lies on the ventral
postcava, ventral to the bodies of the fifth and surface of the tendon of the psoas minor at its
sixth lumbar vertebrae. Caudally, it more often insertion and is flanked by the internal and ex
bends laterally and follows along the cranial ternal iliac veins as these join to form the com
border of the external iliac artery rather than mon iliac vein.
running under it. Afferent lymph vessels drain into it from the
The external iliac lymph node (or nodes) re pelvic limb and it lies in the course of the effer
ceives afferent lymph vessels from all parts of ent lymphatics from the popliteal and superficial
R e g io n a l A n a t o m y of the L y m p h a t ic S ystem 447
inguinal nodes. These lymph vessels, according hepatici) (portal of Baum) (Figs. 6-10 to 6-12,
to Baum, may bypass the node. 6-14) usually consist of right and left nodes, lying
one on each side of the portal vein, 1 or 2 cm.
from the hilus of the liver. They may, however,
Lymph Nodes and Vessels of the vary greatly in number, form, and size. The node
Genital Organs on the left is longer and larger than that on the
right. It lies in the lesser omentum, dorsal to the
The lymphatics of the female genital organs common bile'duct. It is about 3 cm. long and ir
(Fig. 6-15) empty into the lumbar, external iliac, regular in form; the caudal part which reaches to,
internal iliac, superficial inguinal, and sacral extends along, or even extends beyond the gas-
lymph nodes. A fine lymphatic network in the trosplenic vein may be separated from the main
mesosalpinx and fat surrounding the ovary drains lymphoid mass and form one or two additional
into lumbar lymph nodes in the region of the nodes. When single, the left node is about 3 cm.
renal artery and vein. The lymphatics of the cra long. On the right there may be one to five nodes,
nial half of the uterus empty into the lumbar and of various sizes and forms. They lie on the right
external iliac lymph nodes, while vessels of the side of the portal vein opposite the left node and
caudal half of the uterus drain into internal iliac dextral to the gastrosplenic vein at its termina
and sacral lymph nodes. Lymphatics of the va tion. They are closely related to the intermedi
gina enter the internal iliac lymph nodes, while ate part of the pancreas and may be flattened as
those from the vestibulum vaginae in addition to they lie between the layers of peritoneum. Oc
entering the internal iliac and sacral nodes also casionally the two nodal masses are joined cra
pass to the superficial inguinal node. Occasion nially.
ally lymphatics from the vagina or vestibulum The afferent lymph vessels of these nodes
vaginae bypass the internal iliac and sacral lymph come from the stomach, duodenum, pancreas,
nodes and empty directly into the external iliac and liver. The several nodes are connected by
lymph node. The lymphatics of the vulva and cli lymph vessels. The efferent vessels from the
toris pass for the most part into the superficial right node unite into four to eight vessels which
inguinal lymph node. run over both surfaces of the portal vein to the
The lymphatics of the male genital organs cranial mesenteric artery, where they help form
(Fig. 6-16) empty into the same lymph nodes as the intestinal trunk or the network of lymphatics
do those of the female. The scrotum is drained which represents this trunk.
by a coarse network which enters the superficial The splenic lymph nodes (nodi lymphatici
inguinal node. The lymphatics of the testis and
lienales) (Figs. 6-10 to 6-12) are a group of three
epididymis extend in the spermatic cord into the
to five nodes that lie along the course of the
abdominal cavity to enter the external iliac and splenic artery and vein and their terminal
lumbar lymph nodes. For the most part they ac
branches in the dorsal wall of the greater omen
company the blood vessels of the spermatic cord.
tum. Most are small nodes which can be more
Lymphatics of the prostate gland form a coarse
easily palpated than seen in obese specimens.
network on the surface of the gland from which
Usually the largest node lies on the cranial side
several vessels on each side drain into the exter
of the splenic vessels or in the angle of their di
nal and internal iliac lymph nodes. Lymphatics
vision, about 2 cm. from the termination of the
of the prepuce and penis enter the superficial
gastrosplenic vein. The length of this node may
inguinal lymph node.
be as much as 4 cm., but it is more commonly
1.5 cm.
Lymph Nodes and Vessels of the The afferent vessels to the splenic lymph
Abdominal Viscera nodes come from the esophagus, stomach, pan
creas, spleen, liver, omentum, and diaphragm.
The lymph nodes of the abdominal viscera of Their efferent vessels help to form the intestinal
the dog are not numerous, in comparison with trunk or the lymphatic plexus which frequently
those of other species. The following node groups replaces it.
serve the abdominal viscera: ( 1) hepatic, (2 ) The mesenteric lymph nodes (nodi lymphatici
splenic, (3) mesenteric, and (4) colic. To these mesenterici) (Figs. 6-10, 6-11) are the largest
may be added the inconstant gastric, duodenal, lymph nodes of the abdomen. Usually two large
and omental nodes. nodes are present, right and left. Baum (1918)
The hepatic lymph nodes (nodi lymphatici describes these nodes, under the name of jejunal
448 Chapter 6. T he L y m p h a t ic System
449
450 Chapter 6. T he L y m p h a t ic System
Fic:. 6-11. Lyinph vessels of the small intestine and omentum of a dog lying on its back. (After Baiun.)
a. Duodenum h. Spleen (covered in part by omentum) 5. Kight colic lyinph node
b, b\ Jejuimm i. Intestinal mesentery 6 ,6 - „6\ Mesenteric lymph nodes
t . Ileuin I. Cut edge of abdominal wall 7. Middle colic lymph node
d. Cecum 8. Intestinal trunk
e. e'. Colon 1. Omental lyinph node 9. Lymph vessel of the duodenum which
f. Dorsal wall of omentum through which 2. Duodenal lyiuph node goes to the right jejunal lymph node (6)
the stomach can l>e seen 3. Rij;ht hepatic lymph node 10. Cranial mesenteric artery
g. Pancreas 4. Splenic lymph node 11. Jejunal lymph trunk
Fic. 6-12. Lymph vessels and lyinph nodes of the stomach, spleen, pancreas, duodenum, and large intestine of the dog. (After Bauin.)
a. Duodenal lyinph node the mesenteric lyinph nodes (removed) 3 ,3 ' Pancreas
b. Right hepatic lymph nude m. Lyinph vessels of the anus and rectum 4. Spleen (with splenic veins displaced)
c. Left hepatic lymph node n. Lymph vessels which go directly to the 5. Ileum (cut)
d. d'. Splenic lymph nodes lutnbar cistern 6. Cecum
e. Right colic lymph node o. Gastric lyinph node 7 ,8 ,9 . Colon
£ Middle colic lymph nodes P Lyinph vessels of the rectum which course 10. Rectum
g. Left colic lymph nodes over the dorsal surface of the rectum to the 11. Left colic vein
h. Lumbar lymph nodes internal and external iliac lymph nodes 12. Middle colic vein
i. External iliac lyinph nodes 13. Ileocecocolic vein
k Internal iliac lymph node 14. 14'. Portal vein
U '. Lyinph vessel o f the duodenum and 1. Stomach 15. Ventral wall o f the omental sac (reflected)
lymph vessel of the pancreas which go to 2. Duodenum (cut) 16. Mesentery of the colon
451
Fic. 6-13. Lymph vessels of the kidney of the dog; the lymph nodes lying ventr.il to the aorta and
its terminal branches. (The right kidney [f] is displaced toward the pelvis.) (After Baum.)
a. a , a . Diaphragm I P . Left and right cranial liunhar lymph 8. Deep inguinal lymph nodes
b. Psoas muscles nodes (the right node is covered by the 9. Cisterna chyli
c. Lateral abdominal wall postcava) 10. Lumbar trunk
d. e. Muscles of the tail 2. Lum bar lymph nodes which are located 11. Efferent vessels of the superficial inguinal
f, f\ kidneys near the renal artery and vein and medial femoral lyinph nodes (from
g. Postcava 3 ,3 '. Lum bar lym ph nodes (those lal>eled 3 them a part [ 1 PJ enters the deep inguinal
h* \lHlominal aorta are covered by the postcava) lymph nodes (8|)
i. Hight deep circumflex iliac artery and vein 4 , 4\ 4 External iliac lymph nodes 12. Lymph vessels which enter the abdominal
k. Hight external iliac artery and vein 5. Internal iliac lymph nodes cavity from the thorax with the major
1,1'. Right internal iliac artery and vein 6. Middle sacral lymph nodes splanchnic nerve and sympathetic trunk
7. Lateral sacral lymph node 13. Lymph vessels of the diaphragm
452
R e g io n a l A n a t o m y of the L y m p h a t ic System 453
The left colic lym ph nodes (nodi lymphatici Lymph Nodes and Vessels of the Pelvic
colici sinistri) (Fig. 6-12) number two to five, Limb
and lie in the caudal part of the left mesocolon
near the pelvic inlet. The largest member of the The popliteal lymph node (nodus lymphaticus
group is usually located in the angle formed by popliteus) (see Fig. 6-1) is the largest node of
the terminal branches of the caudal mesenteric the pelvic limb. It is rarely double, and is con
artery. The others are mostly located between stant in location. It is an oval node about 2 cm.
the cranial rectal artery and its satellite vein long, which lies in the fat depot between the
which lies on the intestine. They form an inter medial border of the biceps femoris and the lat
connected chain of oval nodes which do not eral border of the semitendinosus, as these mus
extend further caudally than the pelvic inlet. cles diverge from each other. The node and its
Each node is not more than a few millimeters surrounding fat are subcutaneous caudally as
long. they lie in the popliteal space caudal to the stifle
The afferent lymph vessels to the colic lymph joint.
nodes come from the ileum, cecum, and colon. The afferent lymph vessels to the popliteal
The middle colic node receives efferent vessels node come from all parts of the pelvic limb dis
from the left colic nodes. The efferent lymph tal to the location of the node. The 8 to 10 effer
vessels from the right and middle colic nodes ent vessels accompany the medial saphenous
empty into the intestinal trunks. Those from the vein in the popliteal space and unite to form 2 to
left colic nodes go as groups of one to three ef 4 trunks proximal to the origin of the m. gastroc
ferent vessels from each node, and either enter nemius. They continue proximally, close to the
the external iliac, the lumbar, or the middle colic femur, where they lie between the m. semimem
node, or empty into the intestinal trunk directly. branosus and m. adductor. They reach the apex
At various places in the omentum and mesen of the femoral triangle, traverse it and the vascu
tery are inconstant or small nodes, as follows: lar lacuna within the femoral canal, and, after
The gastric lyinph node (nodus lymphaticus crossing the medial surface of the pelvis, empty
gastricus) (Fig. 6-12) lies in the lesser omentum into the external iliac lymph node.
close to the lesser curvature of the stomach near The medial femoral lymph node (nodus lym
the pylorus. It occasionally is absent, or double. phaticus femoralis medialis) is exceedingly in
The duodenal lyinph node (nodus lymphaticus constant and small. Baum (1918) found it only
duodenalis) (Figs. 6-11, 6-12) is quite constant five times in 40 dissections, and then only once
and lies between the initial part of the duode on both sides. It is never over a few millimeters
num and the right limb of the pancreas. Some in diameter, and lies in the fat under the deep
times it lies ventral or ventromedial to the py medial femoral fascia at the distal part of the
lorus, but separated from the closely adjacent femoral triangle. The femoral vessels lie in front
pancreatic lobules. It may be double. of it, and it is intercalated in the main lymph ves
The omental lymph node (nodus lymphaticus sels from the popliteal node. It may receive affer
omentalis) (Fig. 6-11) is the smallest and the ent lymphatics from the medial side of the pelvic
most inconstant of the small variable nodes of limb (Baum 1918).
the abdominal viscera. It lies in the dorsal wall The superficial inguinal lymph nodes (nodi
of the greater omentum 2 to 5 cm. from the duo lymphatici inguinales superficiales) (Figs. 6-15,
denum. A small node may be found in the ven 6-16), usually two in number, begin a few milli
tral wall of the omental sac, a few centimeters meters cranial to the vaginal process and lie in
from the pylorus. the fat which fills the furrow between the ab
The afferent vessels to the omental lymph dominal wall and the medial surface of the thigh.
node come from the omentum; those to the duo In the male, right and left nodes lie along the
denal node come from the duodenum, pancreas, dorsolateral borders of the penis. When a single
and omentum; and those to the gastric node node is present the external pudendal vessels lie
come from the esophagus, stomach, liver, dia lateral to it, but when two nodes are present
phragm, mediastinum, and peritoneum. The ef these vessels usually run between the cranial
ferent vessels from the omental node go into the and caudal poles of the nodes, or the cranial
right hepatic or right colic node; those from the node lies medial to the vessels in a deeper loca
duodenal node go to the right hepatic node; and tion than the caudal node. Their shape and size
those from the gastric node go to the left hepatic vary between wide extremes, but the nodes are
or splenic node, or both (Baum 1918). usually oval and about 2 cm. long.
(Text continued on page 458.)
53
n
c
>
r
>
z
>
H
O
C
■n
H
1
n
r
*
2
X
>
H
n
Fic. 6-15. Lymph vessels of the female genital organs of the dog. (After Baum.) C/3
C/i
1. External iliac lymph node a. Left kidney (reflected) h Bladder H
2,3. Lumbar lymph nodes b. Left ovary (reflected) i. Urethra n
4. Internal iliac lyinph node c. Left uterine hom (reflected) k. Mesosalpinx
5. Superficial inguinal lymph nodes c'. Right uterine hom 1. Broad ligament of the uterus
7. Lateral sacral lymph node d. Body of the uterus m. Lateral ligament of the bladder
8. Lymph vessel which courses to the other e. Vagina n. Ventral abdominal wall
surface of the uterine horn f. Vestibule o, o'. Pelvis, transected
g. Vulva
Cn
Cn
4^
Or
o>
Fi< . 6-17 Lvrnph vessels of the urinary and male genital organs of the dog. {After Baum.)
1. External iliac lymph node b. Ventral abdominal wall (cut) m. Lateral ligament of the bladder
2. Lumbar lymph nodes c. Pelvis, transected n. Ureter
3. Internal iliac lymph node d. Lumbar muscles o. M. coccygeus (cut)
4. Superficial inguinal lyniph nodes e. Aorta p. Cut surface of M . adductor
5. Lateral sacral lyinph node f. Postcava q. M. bulbocavernosus
6 ,6 '. Lymph vessels of the preputial folds g. Left external iliac artery r. M . ischiocave rnosus
7. Lymph vessels of the urethra h. Internal iliac artery s. Penis
8. Lyinph vessel of the bladder i. Bladder t. Clans penis
k. Prostate u. Bulbus glandis
a. Ilium I. Urethra v. Prepuce (opened and reflected)
458 Chapter 6. T he L y m p h a t ic Sy st em
The afferent vessels to the superficial inguinal side of the floor of the abdomen. Depending on
lymph nodes come from the ventral half of the the fullness of the stomach, this may reach any
abdominal wall, including the caudal abdominal level from the infrasternal fossa to a transverse
and inguinal mammary glands. In the male, the plane caudal to the umbilicus. The ventral ex
afferent vessels come from the penis and the skin tremity may be uniformly rounded and approxi
of the prepuce and scrotum. Other afferent ves mately twice as wide as the dorsal half of the or
sels come from the ventral part of the pelvis, the gan. Usually the most cranial portion of this ex
tail, and the medial side of the thigh, stifle joint, tremity is pointed in a direction which ranges
and crus. The superficial inguinal lymph nodes from cranioventral to craniodorsal.
receive the efferent vessels from the popliteal The parietal surface (facies parietalis) faces
node and thus serve as one of the nodal stations the diaphragm and the lateral abdominal wall on
for the whole pelvic limb. The efferent vessels the left side. It extends from the vertebral ends
unite to form one or two trunks which pass of the last two ribs ventrolaterally, mainly oppo
through the inguinal canal with the pudendal site the eleventh left intercostal space. As the
vessels and finally break up in the external iliac distal portions of the last few costal cartilages
lymph nodes. bend forward and downward to form the costal
arch, the spleen continues tangentially across
SPLEEN the medial surface of the costal arch and ob
liquely downward, along the medial surface of
The spleen (lien) (Figs. 6 -1 0 , 6-11, 6-18) is the cranial part of the abdominal wall. It is at
situated in the left hypogastric region, approxi first related to the diaphragm and then to the
mately parallel to the greater curvature of the transversus abdominis muscle. The parietal sur
stomach. It is gray-brown in color, and often has face is slightly convex transversely. The proxi
a purple cast. Although it is irregular in shape mal part is most convex longitudinally as it
and outline, it is considerably longer than it is bends medially toward the median plane.
wide, and slightly constricted in the middle. The The visceral surface (facies visceralis) of the
distal fourth to half is widest and most variable spleen is divided into two nearly equal longitudi
in position. The location of the organ is depend nal parts by the long hilus (hilus lienis) of the or
ent on the size and position of the other abdomi gan. The area cranial to the hilus is related to the
nal organs, particularly the stomach, to which it greater curvature of the stomach; that caudal to
is loosely attached. It is rather firm in consist the hilus is related to the left kidney proximally.
ency, especially when contracted, and has a It is related to the colon at its middle and to the
thick trabecular framework. When enlarged, the mass of the small intestine distally. Even in
spleen may be slightly sigmoid in shape, with hardened specimens, the organs related to the
the ventral end, or free extremity, lying on the spleen do not make clear-cut impressions on it,
floor of the abdomen, frequently extending as the fatty omentum prevents direct contact of
across the mid-ventral line to the right side. It is the visceral surface with the adjacent viscera.
roughly tongue-shaped and presents two ex The cranial and caudal borders (margo crani
tremities, two surfaces, and two borders. In alis et caudalis) of the spleen are thin and irregu
cross section the spleen is triangular. Its longest lar in contour. They may contain shallow or
surface is the outer or parietal surface. deep fissures. There is always a concavity of the
The dorsal extremity (extremitas dorsalis), cranial border proximal to the expanded distal
rounded and wedge-shaped, lies ventral to the end. This may involve the whole border or be re
left crus of the diaphragm, between the fundus placed by an angular depression. In such in
of the stomach and the cranial pole of the left stances the proximal two-thirds of the cranial
kidney. Because of the relatively fixed position border is usually sigmoid in shape. This is
of the left kidney, the position of this part of the masked by a sweeping cranial concavity involv
spleen is the least variable. ing the middle half of the organ. The cranial bor
The ventral extremity (extremitas ventralis) is der may be infolded.
most variable, in both position and shape. When The spleen is attached to the greater curva
the spleen is maximally contracted it is com ture of the stomach by the gastrosplenic liga
pletely hidden under the middle of the caudal m ent (lig. gastrolienale), a part of the greater
border of the rib cage. When it is maximally dis omentum which runs between the stomach and
tended, not only does most of the organ extend the spleen. Proximally, the ligament attaches the
beyond the rib cage, but the ventral extremity greater omentum to the diaphragm.
moves beyond the mid-ventral line to the right The internal structure of the spleen consists
S pleen 459
h o r t g a s t r i c aa. r vv.
S p l e n i c a. f v. ' /
' /
V
/
/
To p a n c r e a s '
To s p t e n o c o h c hg. r -
gr. o m e n t u m
/
To g r o m e n tu m - --
Fic. 6-18. The spleen, showing its blood supply (The cranial border is reflected laterally)
460 Chapter 6. T he L y m p h a t ic Sy stem
of the parenchyma, the red and white splenic long, narrow reticuloendothelial cells. The si
pulp (pulpa lienalis), and a framework which nuses coalesce into veins of the red pulp, and
consists of a. fibrous tunic (tunica fibrosa), or cap these finally merge to become the trabecular
sule, which is rich in elastic and smooth muscle veins. The venous pathway through the splenic
fibers, and trabeculae (trabeculae lienis), which capsule parallels the arterial inflow.
are large and fibromuscular. The trabeculae The exact method whereby blood passes from
form a complicated network within the organ. the arterial to the venous side of the capillary
Some join the veins, strengthening their walls, bed is still in doubt. Three theories have been
and others are independent of them (Trautmann postulated, as follows:
and Fiebiger 1952). The larger intrasplenic ar 1. The “open” circulation theory was held
teries lie mainly in the trabeculae. The collage largely by Mall (1903), who did his research on
nous fibers of the trabeculae continue directly the dog. According to this theory there is no en
into the reticular fibers of the splenic pulp. dothelial continuity between the arterial and
The w hite pulp in the dog consists of diffuse venous sides of the capillary bed, but rather the
and nodular or follicular lymphoid tissue. The arterial capillaries open directly into the red
nodules (folliculi lymphatici lienales) are usually pulp and the blood gradually filters into the
less than 1 cm. in diameter and are not grossly sinuses through the clefts between the reticulo
visible. The germinal centers of these nodules endothelial cells which form their walls. Robin
are lighter in color than the surrounding pulp. son (1930), working with cats, also believed in
Lymphatic tissue is also elaborated along the ar the “open” circulation theory. He cited Mall’s
teries (diffuse lymphatic tissue). The red pulp statement that only about 5 ml. of blood per
consists of the venous sinuses and the cellular minute passes through the spleen of the dog as
tissue filling the spaces between them. It in partial evidence in support of this theory.
cludes many lymphocytes, free macrophages, 2. According to the “closed” circulation
and all the elements of the circulating blood. theory, the arterial capillaries communicate di
The nongranular leukocytes are the most numer rectly with the lumen of the venous sinuses.
ous of these free cells (Bloom and Fawcett 3. The compromise theory was set forth by
1962). Kniseley (1936), who made his observations on
The blood vessels of the spleen are the splenic living, unstimulated spleens. According to this
artery from the celiac artery, and the splenic theory, the walls of the venous sinuses quantita
vein, which drains into the gastrosplenic vein. tively separate the cells of the blood from the fluid
Blood enters the organ by way of about 25 of the blood by continuous filtration processes.
splenic branches (rami lienales), which pass The salient observations of Kniseley’s (1936)
through the long hilus. Once within the capsule experiments were these: The sinuses become
they course in the trabeculae, branching re filled with whole blood by the closing of a physi
peatedly and becoming smaller. When they ological sphincter at the venous or efferent end
reach a diameter of 0.2 mm., they leave the tra of the sinus. As more blood is allowed to pass
beculae and become surrounded by lymphoid into the sinus, the fluid of the blood diffuses
tissue (white pulp), in which they continue to di through the sinus wall, bringing about a great
vide. According to Bloom and Fawcett (1962), concentration of the cellular residue. Appar-
when they reach a caliber of 40 to 50 microns endy the fluid of the blood re-enters adjacent
they leave the lymphatic tissue and enter the capillaries, as these have been observed to con
red pulp. Here they branch into small straight tain relatively cell-free blood. One stage merges
vessels, called penicilli. After these have divided into the next and, at the same time, other capil-
and become smaller, their walls become greatly lary-sinus units are not undergoing cyclic
thickened. Further divisions reduce the arterial changes at all. In one cat the time interval for
capillary to a caliber of not over 10 microns each cycle varied from a few minutes to as long
(Bloom and Fawcett 1962). as 10 hours.
The venous side of the vascular pathway MacKenzie et al. (1941) also observed the
through the spleen begins in the venous sinuses. spleen in cats and other small mammals by trans-
These sinuses (sinus lienis) play an active role as illumination. They observed that in unstimulated
a part of the reticuloendothelial system. They cat spleens contractions take place every 40 sec
occupy more space than does the solid part of onds and last about 25 seconds. MacKenzie
the red pulp, among which they profusely an et al. (1941) state that the pulp spaces consti
astomose. They range from 12 to 40 microns in tute the blood depots, whereas Kniseley (1936)
width, and have walls which are composed of emphasizes the packed nature of the erythro
T h ym us 461
cytes in the sinuses during the storage stage. material. It is laterally compressed, and lies in
MacKenzie et al. state that there are no pre the cranial ventral part of the thoracic cavity.
formed intact connections between the arterial Because it is predominantly lymphoid in struc
and venous systems; Kniseley describes the man ture and hormone production is questionable,
ner in which the cells penetrate the wall. Mac it is described under the vascular rather than the
Kenzie et al. state that the well-developed peri endocrine system.
arterial sheaths of the cat filter and distribute The organ, relatively large at birth, grows rap
much of the blood which enters the red pulp; idly during the first few postnatal months so that
Kniseley attributes only a strong sphincter-like it reaches its maximum development before sex
action to them. Barcroft and Florey (1928) state ual maturity, or between the fourth and fifth
that a perfectly well defined lymphatic system postnatal months, just before the shedding of the
drains the spleen. These investigators ligated the deciduous incisor teeth. The thymus begins to
splenic veins and injected trypan blue through involute with the changing of the teeth. Al
the splenic artery in the cat. A lymphatic vessel though the process is rapid at first, the organ
was seen running from the hilus to one of the usually does not atrophy completely even in old
mesenteric lymphatic nodes. age. As it decreases in size and loses its lymphoid
True accessory spleens are rare in the dog. structure, it is replaced by fat.
Armstrong (1940) described three animals hay The thymus is located almost entirely in the
ing scores of accessory spleens as a result of precardial mediastinal septum. Its cranial end,
postnatal trauma. Many small splenic grafts pars cervicalis, is 0.5 to 1 cm. long and lies cra
were dispersed in the omentum. Some spleens nial to a transverse plane through the thoracic
were divided and others showed extensive cica inlet. The main portion of the organ, pars tho-
trix formation. Dogs usually do not possess racalis, extends from the thoracic inlet to a
hemal lymph nodes. curved line opposite the left side of the heart.
The nerve supply to the spleen is from the ce This place is usually located opposite the fifth
liac plexus, and consists chiefly of non-myeli- costal cartilage.
nated postganglionic sympathetic fibers. The Each polygonal lobule, which may measure
few myelinated fibers present are probably affer over 1 cm. in length, is separated from adjacent
ent axons. The vagi also send fibers to the lobules by a delicate but distinct connective tis
spleen. The nerves to the spleen form the splenic sue capsule, so that the lobules move freely on
plexus of nerves which entwine the splenic ar each other. The whole organ is further divided
tery. into right and left lobes (lobus dexter et sinister).
Several diverse functions have been attributed The division between the two lobes is distinct
to the spleen of the dog: caudally, but cranially it is not always possible to
1. It stores and concentrates the erythrocytes determine to which lobe the adjacent lobules
and releases them during times of need. belong, because the lobes are united here by es
2. It filters the blood, as the lymph nodes fil sentially the same amount and kind of connec
ter the lymph, and removes the worn-out eryth tive tissue that unites the lobules.
rocytes from the circulation. From these cells it The left lobe extends further caudally than
produces bilirubin, which is collected by the the right. It occupies a special outpocketing of
liver. From the hemoglobin it extracts the iron, the precardial mediastinal portion of the pleura
which is released here, and is again used by the of the left pleural sac. Its caudal border is gently
red bone marrow in the production of new curved and thin as it lies between the left thora
erythrocytes. cic wall and the left ventricle. In a small dog this
3. It produces many of the lymphocytes and portion of the left lobe may have a dorsoventral
probably most of the monocytes, and has an im dimension of 5 cm. and may be 5 mm. thick. The
portant function in the production of antibodies. left lobe becomes slightly narrower cranially as
The spleen is not essential to life or even to it lies in the mediastinal septum, loosely fused to
health, as most of its normal functions are taken the right lobe.
over by other tissues in its absence. The right lobe of the thymus usually butts
against the cranial surface of the pericardial sac
and is expanded laterally. The thickest part of
THYMUS the thymus is located here. When maximally de
veloped in the beagle, the thymus has the follow
The thymus (Fig. 6-19) is a light gray, dis ing measurements: 12 cm. long by 6 cm. wide
tinctly lobulated organ with a pink tinge in fresh by 3 cm. thick. When fully developed in this
462 Chapter 6. T he L y m p h a t ic Sy stem
lEsophagus
L . s u b c la v ia n o.x i
breed, it weighs about 50 gm. The right lobe (Baum 1918). Both parasympathetic (vagal) and
accounts for 60 per cent of the weight, and the sympathetic nerve fibers supply the organ, and
left lobe for 40 per cent. Variations in size and are probably vasomotor. The basic cell unit of
form are common. Latimer (1954) has studied the thymus is a small lymphocyte, sometimes
the fetal development of the thymus in the dog. referred to as a thymocyte. These cells do not
Dorsally the thymus is related to the phrenic differ from small lymphocytes found in other or
nerves, precava, and trachea. The apical lobes of gans of the body. The thymic corpuscles (Has-
the lungs produce large smooth impressions on sall’s bodies) are spherical or oval structures in
its lateral sides when it is maximally developed. the medulla of the gland, composed of concen
The internal thoracic vessels form deep clefts, or trically arranged cells. The centers of many thy
even run through the cranioventral portion of mic corpuscles consist of degenerated cells
the gland. The thymus receives its chief blood which may be hyalinized, cystic, or calcified.
supply from one or two thymic branches which The function of the thymus is not clear. A sur
go to each lobe from the ipsilateral internal tho vey of the thymus and its relation to lymphocytes
racic artery. Occasionally, an additional thymic and immune reactions by Arnason, Jankovic, and
branch leaves the brachiocephalic artery on the Waksman (1962) indicates that the thymus ap
right side and the subclavian on the left side. pears to be a principal source of the small lym
The veins from the thymus are satellites of the phocytes of the blood, spleen and lymph nodes.
thymic arteries. The lymphatics from the thy The thymus plays a major role, particularly in
mus form four to six vessels which empty into early life, in maintaining the immunologic in
the cranial mediastinal and sternal lymph nodes tegrity of the organism.
T h ym us 463
IN T R O D U C T IO N TO THE
NERVOUS SYSTEM
B y R A L P H L . K IT C H E L L
The nervous system has been by custom cate shorter than axons. In some neurons the axon
gorically divided into central and peripheral arises from the cell body at the axon hillock.
systems. This division is not made on the basis of The cytoplasm of many nerve cells contains
definitive structural or functional considerations, an abundance of cytoplasmic chromidial sub
but merely for convenience in discussion. The stance, called Nissl substance (Figs. 7-1, 7-2).
central nervous system is defined as consisting This material stains blue with basic dyes such as
of the brain and the spinal cord. The peripheral thionin, cresyl violet, toluidin blue or methylene
nervous system includes the cranial and spinal blue. Electron microscopy shows chromidial
nerves, their ganglia, and peripheral portions of substance to be a highly developed and organ
the autonomic nervous system. ized endoplasmic reticulum consisting of densely
The central nervous system is composed of packed, narrow tubes with fine granules associ
large numbers of cells called neurons bound to ated with the tubes. Histochemical and other
gether by a framework of supporting cells called studies have demonstrated that the granules con
neuroglia. The neuron is the genetic, structural, sist primarily of ribonucleic acid. The cytoplasm
functional and trophic unit of the nervous sys also contains mitochondria and Golgi substance.
tem. It is the fundamental unit of the nervous Mitochondria are numerous in the cell body and
system from which more complex structural and in the dendrites, but are fewer in number in the
functional concepts must originate. axon. Neurofibrillae run in all directions within
the cell body. They are also present in the den-j
drites and axon. Specialized receptors and nerve
STRUCTURE OF NEURONS endings may be found as terminals of certain
nerve cells having specific functions.
A neuron consists of a cell body (neurocyton
or perikaryon) and its processes (Fig. 7-1). There FUNCTION OF NEURONS
are wide variations in the shapes and sizes of
neurons. Neurons are often classified on the Nerve cells have the ability to respond rapidly
basis of structural characteristics such as the to a stimulus, and if the response initiated by the
shape of the cell body and the number, shape stimulus is of sufficient magnitude, the impulse
and ramifications of the processes. Dependent is conducted by the axon independent of the
upon the number of processes arising from the stimulus. The physicochemical reactions in
cell body, a neuron may be termed unipolar, volved in the conducted response are termed the
bipolar or multipolar. nerve impulse (Katz, 1961; Tasaki, 1959). The
Neuronal processes may be classified on a nerve impulse involves the axon membrane,
structural basis as dendrites or axons. Dendrites which has special electrical properties and selec
have an internal structure similar to that of the tive permeability to various ions. The axon mem
cell body, whereas axons are more specialized. brane separates two aqueous solutions, one in
Dendrites are more numerous and are usually side the axon membrane and one outside, which
464
F u n c t io n of N eu ro n s 465
=>
g
z
UJ
O '''''COLLATERAL BRANCH
AXON NONMYELINATED
---- COLLATERAL BRANCH
- A X O N MYELINATED
AXON
MYELINATED AND
COVERED WITH
NEURILEMMA
(Na+) (Na+)
+ + +\>- ■^r^- + + + + + + + + + + + + + + + + + + + + + + + + + + +
rr~
A X O N I ____
- \+ +
+ + +/X*r Titf- + + + + + + + + + + + + + ++ + + + ++ + + + + + + +
(Na+) (Ha*)
+ + + + + + + + + + + + + + t----c A + + + + + + + + + + + + + + + +
AX0N*r (**)+ +. I I I I I I I I I I I I I I I t
+ + + + * + + + + + + + + + 1---- S~sA- + + + + + + + + + ++ + + + + +
+ + + + + + + t - -'55 k + + + + + + + + + + + + + + +++++
w///////////^////m^tm^m
v^2n+ + \~
AXON ■&+ +T-
A +l 1
+ + + + + + + I — JL+ + + + + + + + + + + + + + + I — J l^+ + + + + +
F ig . 7-3. Schematic representation of nerve impulse. A, Depolarizing electrotonic effects result in sodium ions entering the
axon. The resting potential is gradually altered until threshold is reached. At this time sodium ions enter the axon in great num
bers, resulting in an action potential. B and C, propagation of the nerve impulse is illustrated. Sodium ions enter the axon in ad
vance of the nerve impulse, and potassium ions flow out after the impulse has passed. The outflow of potassium ions results in a
brief refractory period. (Modified from Katz, 1961.)
468 Chapter 7. In tro d u c tio n to th e N e rv o u s S y s te m
DORSAL ROOT
F ig . 7-4. Diagram illustrating relations of afferent neurons to dorsal roots, interneurons within central nervous system, and ef
ferent neurons to the ventral roots of the spinal cord. The arrows indicate the direction nerve impulses are usually transmitted.
Dendritic
zone
Axon -
Origin
Axon
Telodendria
F ig . 7-5. Diagram of various receptor neurons, interneurons, and motor neurons, illustrating that the impulse origin, rather
than the location of the cell body, is a useful method in relating neuron structure to function. (Modified from Bodian 1962.)
G en era l St r u c tu r a l and F u n c t io n a l O r g a n iz a t io n of P e r ip h e r a l N e r v e s 469
dendria are the usually branched and variously (3) General visceral afferent fibers supply re
differentiated terminals of axons which show ceptors located in visceral structures. (4) Special
membrane and cytoplasmic differentiation re visceral afferent fibers carry impulses away from
lated to specialized transmission or neurosecre receptors in tissues of pharyngeal arch origin
tory activity (Figs. 7-5, 7-14, 7-15, 7-16). and the olfactory mucosa. (5) Somatic efferent
fibers are motor fibers to skeletal muscles. (6 )
SHEATHS ASSOCIATED WITH AXONS General visceral efferent fibers are motor fibers
going to cardiac muscle, smooth muscle, and
The cell membrane of an axon is surrounded glands. These fibers are also referred to as auto
by a lipid membrane, the myelin sheath, which nomic nerve fibers. Autonomic nerve fibers leave
is formed by neurilemmal or glial cells (Figs. 7- the central nervous system in nerves from three
1, 7-7). The myelin sheath cannot be seen with separate regions of the body. These are the
the light microscope in axons less than 1 micron cranial and upper sacral regions (parasympa
in diameter. The largest axons, with their myelin thetic fibers) and thoracolumbar region (sym
sheaths, may be 20 microns in diameter. The pathetic fibers). Autonomic fibers in peripheral
neurilemma is a thin layer of flat cells so arranged nerves are of two groups according to their
that they enclose the myelin sheath and the axon. tandem arrangement (Fig. 7-9). Preganglionic
Many axons less than 1 micron in diameter may fibers have their cell bodies located within the
be surrounded by a common neurilemma (Fig. central nervous system. These nerve fibers pro
7-6). Within the central nervous system, the ject to parts of the peripheral nervous system to
glial cells surround the myelin sheath and the terminate near the cell body of a postganglionic
axon. The neurilemma (or glial network), myelin fiber. Preganglionic neurons can be regarded as
sheath and the axon are collectively referred to efferent interneurons. Postganglionic neurons
as a “nerve fiber.” Nerve fibers less than 1 mi have their cell bodies located in autonomic
cron in diameter are frequently referred to as ganglia outside the central nervous system (Fig.
nonmyelinated (unmyelinated or nonmedul- 7-9). Their axons terminate in the structures
lated) fibers, and fibers having a diameter greater they innervate (see Autonomic Nervous System).
than 1 micron are referred to as myelinated (7) Special visceral efferent fibers are motor
fibers. Osmic acid, when used as a stain, colors fibers to muscles of pharyngeal arch origin.
the myelin black, leaving the axon unstained Historically, and traditionally, afferent fibers
(Fig. 7-10). Silver precipitation methods, as well have been referred to as sensory fibers and
as other staining techniques, affect only the axon, efferent fibers as motor fibers. However, physio
leaving the myelin unstained (Fig. 7-8). The logical activity in afferent fibers does not mean
latter techniques are used to demonstrate the that the stimulus producing this activity will be
presence of axons less than 1 micron in diameter. perceived. This is true for activity in both so
The myelin sheaths are interrupted at regular matic afferent and visceral afferent fibers; how
intervals. These interruptions are termed nodes ever, in man, in only a very few instances will
of Ranvier (Fig. 7-7). The distance between activity in general visceral afferent fibers result
nodes (internodal segment) is related to the di in perception of the stimulus.
ameter of the fiber. Fibers having large diameters The terms “motor” and “efferent” are often
have longer internodal segments than fibers used interchangeably. Some authors will refer
having small diameters. In peripheral nerves, to a nerve supplying a muscle as a “motor”
one neurilemma cell forms the sheath for one nerve. This is incorrect and misleading, since, in
internodal segment. most instances, significant numbers of fibers in
these nerves are general somatic afferent fibers.
FUNCTIONAL CLASSIFICATION OF
NERVE FIBERS GENERAL STRUCTURAL AND FUNCTIONAL
ORGANIZATION OF PERIPHERAL NERVES
Nerve fibers in peripheral nerves can be func
tionally classified according to the structures A cranial or spinal nerve consists of its root
which they supply (Fig. 7-9). (1) General so (radix) or roots (radices), associated cranial or
matic afferent fibers carry impulses away from spinal ganglia, the nerve itself and its peripheral
receptors located in the skin and skeletal struc branches. Cranial nerves vary in the number of
tures. (2) Special somatic afferent fibers carry roots, some having only one root and others hav
impulses away from receptors located in special ing two or more roots (see Chapter 10).
sensory areas such as the eye and the internal ear. All spinal nerves have two roots, a dorsal root
Chapter 7. I n t r o d u c t io n to the N ervou s S y stem
470
SCHWANN CEU
cyto plasm
F ig . 7-9. Schematic diagram illustrating four primary branches o f a spinal nerve and the functional classes o f nerve fillers found
in these branches. The dorsal and ventral branches of a spinal nerve contain general somatic afferent, general visceral afferent, so
matic efferent, and general visceral efferent fibers (all contain postganglionic sympathetic fillers. The ventral branches of the first
three sacral nerves also contain preganglionic parasympathetic fibers). The meningeal rami are very small nerves going to the me
ninges. They contain general somatic afferent, general visceral afferent, and general visceral efferent fibers (not shown). Note that
preganglionic sympathetic fibers are present in the ramus eommunieans of L 5 (T 1 to L 5 or 6), but are absent in the ramus com-
municans of other spinal nerves.
Stru ctu res M e d ia t in g R e c e p t io n , C o n d u c t io n , and R espo n se 473
LOCATION F SECTION
STAINING
C —5 T—9
TECHNIQUE
DORSAL ROOT VENTRAL ROOT DORSAL ROOT VENTRAL ROOT
rfe f vQ k
WOLTER
i$£
v * • -« * • L -
Tv*. '*’• * ' * * § : *1 •<
• .V » * V
1M •
HOLMES • > W v
- 4*** • *#
- - 1 M f A-li .v>-_
0 18 36 54 72
MICRONS
A B c n
Fig. 7-10. Cross section of portions of dorsal and ventral spina] nerve roots of a dog. A. Dorsal root, third cervical spinal nerve.
B, Ventral root, third cervical spinal nerve. C. Dorsal root, ninth thoracic spinal nerve. D, Ventral root, ninth thoracic spinal nerve.
Upper row. Wolter's modification of YVeigert-Pal technique. Lower row, Holmes's technique. Histograms represent the myelinated
fiber diameter spectrum of each root. Note the approximately unimodal distribution of sizes of myelinated fibers in the dorsal roots
as compared to the bimodal distribution in the ventral roots. The relatively larger number of fibers in the 2 -6 micron diameter
groups in the ventral roots of T 9, as compared to similar groups in C 3, is the result of the presence of preganglionic sympathetic
fibers in T 9 and the absence of such fil>ers in C 3.
EPINEURIUM
PERINEURIUM
PERIPHERAL -
SPINAL
SENSORY ENDING
ENDING
AXON
ONE MYELIN SHEATH
NERVE •
FI8ER NEUROLEMMA
of schwann)
Fig. 7-11. Diagram showing the various parts of a sizable peripheral nerve. (After Ham and Leeson 1961.)
474 Chapter 7. I n t r o d u c t io n to the N ervous S ystem
DIAMETER ( j j ) DIAMETER ( j i )
Fic. 7 12. Histograms showing the frequency distribution of various diameters of myelinated fibers in the communicating
branches of the ninth thoracic and the third sacral nerves. The larger numbers of fibers in the 2- and 4-inicron diameter groups of
the ninth thoracic are due to the presence of preganglionic sympathetic fil>ers in this ramus.
Epi dermis
Sweat
Gi an * Dermis
Subcutaneous
Fat
tween a specific physical or chemical stimulus neurons or efferent neurons. Likewise, a single
and a set of receptors having a similar structure. interneuron or a single efferent neuron may have
Numerous studies have shown that a number of many boutons (up to a thousand or more) of
receptors, regardless of their structure, are dis telodendria from many afferent or other inter
tinctly responsive to specific physical or chemi neurons forming synaptic junctions with its
cal stimuli and much less responsive to other dendritic zone (receptor portion of the cell
physical or chemical stimuli (Gray, 1959). In membrane; Figs. 7-15, 7-16).
other words, a good correlation exists between The effect of a nerve impulse (or impulses) in
a specific receptor and a specific stimulus (phys presynaptic terminals upon the generation of
iological specificity), but a good correlation does impulses in an axon of an interneuron or an
not always exist between the specific stimulus efferent neuron is a graded effect similar to that
and the specific structure of a group of receptors occurring in the receptor zone of an afferent
(anatomical specificity) (Melzak and Wall 1962). neuron. It is doubtful whether one nerve im
A receptor responds to a specific stimulus in pulse in one presynaptic terminal (or bouton)
a graded manner dependent upon the physio will result in the generation of a nerve impulse
logical status of the receptor as well as upon the or impulses in the axon of an interneuron or of
intensity and duration of the stimulus being a motor neuron. A presynaptic fiber may be
applied. All receptors, therefore, have the regarded as excitatory or inhibitory to a second
properties of selectivity, sensitivity, and adap neuron. Some investigators interpret their
tation. If the activity generated in the receptor findings such as to deny that afferent neu
in response to the stimulus reaches sufficient rons have synaptic endings which are inhibitory.
magnitude, this will result in the generation They propose that the inhibitory effects of af
of an impulse in the distal segment of the ferent neurons are through excitatory effects on
axon. Each impulse is conducted toward the intemeurons which in turn produce only inhibi
telodendria at a given rate for each nerve fiber. tory effects on other neurons. Other investigators
The rate of conduction for any segment of the conclude that an afferent neuron, or an inter
nerve fiber can be related to the fiber diameter neuron, may be both excitatory and inhibitory.
and in turn to the internodal length. The activity Regardless, the evidence available indicates that
in an axon seen after the application of a specific the presynaptic excitatory and inhibitory effects
stimulus to its receptor is governed almost en produced at the receptor membrane (subsyn-
tirely by the response of the receptor. An axon aptic membrane) interact in such a manner that,
reflects activity in the receptor by an increase (or if the excitatory effects are of sufficient magni
decrease) in the number of impulses transmitted tude, depolarization of the initial segment of an
per unit of time (frequency of nerve impulses). axon will occur and a nerve impulse will be gen
The pattern of the frequency of the response erated in the axon of the interneuron or efferent
may also be of some significance. neuron (Fig. 7-14). The nerve impulse initiated
The telodendria of an axon of an afferent in the axon of an interneuron or motor neuron
neuron or an interneuron contacts the dendritic is an “all or nothing” phenomenon similar to
zone (or zones) or receptor portions of an inter that discussed earlier for an afferent axon.
neuron or an efferent neuron. This functional Intemeurons may conduct the impulse di
connection between two nerve cells is called a rectly to efferent neurons or to other inter
“synapse” (Fig. 7-14). A synapse is formed by neurons and to the brain. The former pathway
the presynaptic fiber (telodendrion) and the may result in a reflex response (Fig. 7-4), where
postsynaptic membrane of the second neuron. as the latter pathways may result in other reflex
A gap, no more than 200 angstroms in width, responses or subtle changes in the excitatory
separates the two structures. Many of the telo state (either raising or lowering it) of other inter
dendria have terminal enlargements called neurons (Fig. 7-14). Very complex interrela
boutons (Figs. 7-14, 7-15, 7-16; boutons termi- tions among neurons within the central nervous
naux if the nerve fibers ends there or boutons en system are believed to be involved in the per
passage if the nerve fiber goes to another ception of the stimulus. Earlier referen ce was
neuron). The telodendria. may terminate by m ade to the difficulties encountered in correlat
means of many small fibers which intermingle, ing specific stimuli to the specialized mor
but establish no protoplasmic connections, with phology of receptors. Also it was stated that a
the arborizations of the dendrites. The teloden given receptor, regardless of its morphology,
dria of an axon from a single afferent neuron or when stimulated by appropriate stimuli, gives a
an interneuron contacts a large number of inter specific physiological response to a specific stim-
(Text continued on page 479.)
476 Chapter 7. I n t r o d u c t io n to the N erv o u s Sy stem
F ig . 7-14. Schematic diagram illustrating synaptic relations between a bouton and a neurocyton of an interneuron (or a motor
neuron). Presynaptic activity in a bouton may be either excitatory or inhibitory. Excitatory and inhibitory synaptic activity inter
act algebraically to effect generator activity in the somatic-dendritic segment; thus a gradation of activity can occur. Lines of cur-
rent-flow are shown which occur when a synaptically induced depolarization of the somadendritic membrane electronically
spreads to the initial segment of the axon. If the activity induced in the initial segment of the axon is of such magnitude as to cross
the threshold of that segment, a nerve impulse will be generated in the axon.
Str u c t u r e s M e d ia t in g R e c e p t io n , C o n d u c t io n , and R espo n se 477
F ig . 7-15. Diagram illustrating boutons on the surface of interneurons or motor neurons. A, Boutons terminaux. B, Boutons en
passage.
478 Chapter 7. I n t r o d u c t io n to the N ervou s S y stem
F ig . 7-16. Photomicrographs illustrating boutons on a lower motor neuron in the dog. Each arrow points to one of the many
boutons on the surface of the cell membrane. Silver technique.
St r u c t u r es M e d ia t in g R e c e p t io n , C o n d u c t io n , and R espo n se 479
THE BRAIN
B y HERM ANN M EYER
The central nervous system consists of the brain major parts: the telencephalon, diencephalon,
and the spinal cord. The brain develops from the mesencephalon, metencephalon, and the myel-
rostral intracranial portion of the early neural encephalon in rostrocaudal sequence. The pat
tube, while the spinal cord is derived from the tern of this subdivision of the brain is based on
remainder of the neural tube. the early development of the rostral part of the
Structurally, the central nervous system con central nervous system (Fig. 8-1).
sists of gray matter and white matter. The gray At the rostral intracranial end of the neural
matter is formed by the aggregation of the cell tube three vesicPes develop. In rostrocaudal
bodies of the neurons within the central nervous sequence they are called the prosencephalon or
system. The white matter consists mainly of forebrain, the mesencephalon or midbrain, and
myelinated nerve cell processes or portions of the rhombencephalon or hindbrain. In the
processes inside the central nervous system. further development the prosencephalic vesicle
In the spinal cord the cell bodies form cen divides into the telencephalon and the dien
trally located continuous gray columns within cephalon. The mesencephalon does not divide.
the white matter. In the brain the spatial rela The rhombencephalon again gives rise to two
tion of gray and white matter is different. The parts: the metencephalon and the myelencepha-
columns of gray matter, as they are found in the lon.
spinal cord, are intersected by white matter and From a purely macroscopic point of view the
form discrete entities of gray matter. Such ac brain may be subdivided into three components:
cumulations of cell bodies within the central the cerebrum, the brain stem, and the cerebel- -
nervous system are called nuclei. In addition to lum (Fig. 8-2).
these nuclear masses, gray matter is located The cereFrum is the largest, most rostral part
peripherally in the cerebral hemispheres and the c of the brain. It is derived from the telencephalon.
cerebellum as cerebral cortex and cerebellar The hrain stem includes the entire diencephalon
cortex respectively. The white matter between and mesencephalon, the ventral portion of the
the nuclear masses of the brain and within the metencephalon and the entire myelencephalon.
spinal cord consists in part of fiber groups with It connects the cerebrum with the spinal cord
common connections and functions. These and the cerebellum. The cerebellum is located
fiber groups form fiber bundles and are referred on the dorsal aspect of the caudal portion of the
to as tracts or fasciculi. They extend between brain stem. Its developmental origin is from the
centers in the gray matter of the various portions dorsal part of the metencephalon.
of the central nervous system. For morphological information about the
The lumen of the neural tube inside the cen dog’s brain refer to Ackerknecht (1943), Ariens
tral nervous system persists in the form of cavi Kappers, Huber, and Crosby (1936), von Bech-
ties and connecting canals. These spaces develop terew (1899), Bourdelle and Bressou (1953),
into the central canal of the spinal cord and the Bradley and Grahame (1948), Bruni and Zim-
ventricular system of the brain (Fitzgerald 1961). merl (1951), Crosby, Humphrey, and Lauer
They contain cerebrospinal fluid. See Ullrich (1962), Dexler (1932), Edinger (1897, 1911),
(1928), Vuillaume (1935), Nigge (1944), Verwer Ellenberger and Baum (1891), Ellenberger and
(1952), Fankhauser (1962). Baum (1943), Flatau and Jacobsohn (1899),
The brain is customarily divided into five Ghe^ie, Riga, and Pastea (1956), Kuntz (1950),
480
T he B r a in 481
E
E
01
R ig h t
hemisphere
Longitudinal
fis s u r e
hemisphere
CEREBRUM CEREBELLUM
BRAIN STEM
F ig . 8-2. Gross subdivisions of th e brain.
482 C h apter 8. T he B r a in
Martin (1923), Miller (1958), Papez (1929), tex at the base of the brain is phylogenetically
Peele (1961), Ranson and Clark (1959), Sisson older and consists of the paleocortex or paleopal-
and Grossman (1953), Ziehen (1906). lium and archicortex or archipallium.
The atlas on the dog’s brain by Adrianov and In the course of development the cerebral
Mering (1959), the stereotaxic atlas by Lim, Liu, cortex becomes folded, and a system of grooves
and Moffitt (1960), and “The Brain of the Dog in and elevations is formed. The elevations or con
Section” by Singer (1962) contain histological volutions are known as gyri; the depressions be
information. For comparative considerations the tween the gyri are the sulci. The sulcal and gyral
most recent atlas on the cat’s brain (Snider and pattern is typical for the species, but individual
Lee 1961) may be added. variations are relatively frequent. A few partic
Weights and measurements of the dog’s brain ularly distinct sulci, however, are constant and
and its parts were reported by a number of in form a basic pattern. For some of these the terms
vestigators: Rudinger (1894a), Davison and “sulcus” and “fissure” are used interchangeably.
Kraus (1929), Michaels and Davison (1930), The following description of the sulcal and
Michaels and Kraus (1930), Latimer (1942,1946, gyral pattern of the cerebral hemispheres (Figs.
1954), Corder and Latimer (1947, 1949), Ste 8-3 through 8-9) will be limited to structures
phen (1954). essential for an understanding of normal condi
Clinical and pathological studies with related tions. Detailed information is available on cranio-
morphological information on the dog’s brain encephalic relations (Caradonna 1902; Mobilio
may be found in the works of Frauchiger and 1912; Lucas 1939), gyri and sulci (Wilder 1873a,
Fankhauser (1949), McGrath (1953, 1960), and 1873b, 1881; Krueg 1880; Langley 1883-84a;
Innes and Saunders (1962). Lesbre 1884; Ellenberger 1889; Ellenberger and
Baum 1891; Turner 1890; Flatau and Jacobsohn
1899; Holl 1899; Weinberg 1902; Balado 1925;
TELENCEPHALON Papez 1929; Cohn and Papez 1933; Ariens
Kappers, Huber, and Crosby 1936; Seiferle
The telencephalon forms the cerebrum. It 1957), variations of the sulcal and gyral pattern
consists of two cerebral hemisperes (Fig. 8-2) (Rudinger 1894b; Hoenig 1912; Filimonoff 1928;
which are separated along the midline by the Marburg 1934; Oboussier 1949, 1950; Starck
longitudinal fissure (fissura longitudinalis). 1954), and the evolution of the brain in fossil
Each of the two cerebral hemispheres is di carnivores (Pivetau 1951).
vided into four topographical areas: the frontal The rhinal sulcus or fissure (sulcus rhinalis) is
lobe (lobus frontalis), the parietal lobe (lobus one of the most constant grooves of the cerebral
parietalis), the occipital lobe (lobus occipitalis), cortex. It is located on the ventrolateral aspect
and the temporal lobe (lobus temporalis). The of the brain and extends along the entire length
boundaries of these lobes are more or less arbi of the cerebrum. It separates the phylogeneti
trary. The frontal lobe comprises the rostral por cally more recent neocortex or neopallium from
tion of the cerebral hemisphere. The occipital the olfactory cortex at the base of the brain. At
lobe is at the caudal end. The temporal lobe con about its middle it has a sharp flexure and is
sists of the ventrolateral areas, and the parietal joined obliquely by another constant furrow, the
lobe includes the remaining dorsolateral portion relatively short sylvian fissure or sulcus. This
of the cerebrum. junction divides the rhinal sulcus into the an
From a structural point of view the cerebrum terior and the posterior rhinal sulci (sulcus
consists of a peripheral layer of gray matter or rhinalis anterior et sulcus rhinalis posterior) re
cerebral cortex; white matter beneath the cortex; spectively.
centrally located basal nuclei or basal ganglia; From about the middle of the anterior rhinal
and the phylogenetically older olfactory por sulcus the presylvian sulcus (sulcus praesylvius)
tions. Inside each cerebral hemisphere there is extends rostrodorsally toward the midline. Its
a cavity, the lateral ventricle (ventriculus later dorsal end may unite with the inconstant prorean
alis) which developed from the lumen of the sulcus (sulcus proreus), which, if present, ap
telencephalic vesicle. pears as a sagittal sulcus and parallels the rostro-
The cerebral cortex is the outermost gray dorsal margin of the frontal lobe.
matter of the cerebrum which covers the hemi The olfactory sulcus (sulcus olfactorius) is the
spheres like a cloak or mantle. For this reason it rostral continuation of the anterior rhinal sulcus.
is also referred to as the pallium. Its dorsal part, It runs parallel to the presylvian sulcus and is
a phylogenetically recent acquisition, is called partly hidden by the olfactory bulb, which will
the neocortex or neopallium. The olfactory cor be described with the rhinencephalon. The pos
T E L E N C E PH A LO N 483
terior rhinal sulcus may bifurcate caudally and quently present than the precruciate sulcus.
form a lateral and a medial limb. On the medial surface the corpus callosum, a
The sylvian fissure (fissura lateralis Sylvii) is large fiber connection between the two hemi
identical with the lateral fissure as described by spheres, is bounded rostrally, dorsally, and
Sisson and Grossman (1953). For reasons of caudally by the callosal sulcus (sulcus corporis
homology, Ariens Kappers, Huber, and Crosby callosi). At the caudal end of the corpus callosum
(1936) suggest that pseudosylvian fissure would the callosal sulcus blends with the hippocam pal
be a better name for this groove because it is not sulcus (sulcus hippocampi), which runs ventro-
homologous with the sulcus lateralis (Sylvii) of rostrally to the temporal lobe. In doing so it
the human brain. From its junction with the describes a concave line and curves around the
rhinal sulcus it extends for a short distance in a brain stem.
caudodorsal direction. Peripherally, about halfway between the
In the caudal three-fourths of the lateral corpus callosum and the margin of the medial
aspect of the cerebrum the constant sylvian fis side of the cerebral hemisphere, the calloso-
sure is surrounded by three equally constant, marginal sulcus (sulcus callosomarginalis) de
concentric and almost semicircular grooves. The scribes an elongated semicircle. This sulcus may
first, and most central, is the ectosylvian sulcus be divided into three parts, rostral, middle, and
(sulcus ectosylvius). It is followed in a peripheral caudal.
direction by the suprasylvian and ectomarginal The rostral portion of the callosomarginal
sulci (sulcus suprasylvius et sulcus ectomargi- sulcus, the genual sulcus (sulcus genualis), is
nalis). separate from the other two portions. It is vari
The ectosylvian and suprasylvian sulci have able, but in general it parallels the course of the
three parts each: the anterior, middle, and pos callosal sulcus around the rostral end or genu of
terior ectosylvian and suprasylvian sulci (sulcus the corpus callosum. The genual sulcus is usually
ectosylvius et suprasylvius anterior, medius, et accompanied peripherally by the ectogenual
posterior) respectively. The ectomarginal sulcus sulcus (sulcus ectogenualis).
is also divided into three portions. The rostral The remaining middle and caudal portions of
part is the coronal sulcus (sulcus coronalis). The the callosomarginal sulcus sometimes are jointly
middle is referred to as the lateral sulcus (sulcus referred to as the splenial sulcus (sulcus spleni-
lateralis). The caudal part is the postlateral alis). Usually, however, this name is more logi
sulcus (sulcus postlateralis) and represents the cally reserved for the caudal part which curves
equivalent of sulcus medilateralis of some au around the caudal end or splenium of the corpus
thors. Occasionally two or all three of these may callosum. In this case the middle portion is called
not be connected, but they are usually in line the cingulate sulcus (sulcus cinguli). The lesser
and suggest the concept of the ectomarginal cruciate sulcus (sulcus cruciatus minor) is given
sulcus. off from about the middle of the cingulate sulcus
In most specimens a groove is found between and reaches the dorsal margin of the brain at the
the lateral sulcus and the middle and posterior level of the ansate sulcus.
suprasylvian sulci. Since this groove is peripheral In a substantial number of brains the splenial
to the lateral sulcus, it is appropriately named sulcus gives off a posterior horizontal ramus
ectolateral sulcus (sulcus ectolateralis). Less fre (ramus horizontalis posterior), which, according
quently a depression referred to as entolateral to Ariens Kappers, Huber, and Crosby (1936),
sulcus (sulcus entolateralis) is seen medial to the appears to be the forerunner of the posterior
lateral sulcus. The ansate sulcus (sulcus ansatus) part of the calcarine sulcus. See also Cohn and
is a short medial branch of the rostral part of the Papez (1933). The ventral end of the splenial
lateral sulcus. sulcus may connect with the previously de
The most conspicuous sulcus on the dorsal scribed medial limb of the posterior rhinal sulcus.
aspect of the cerebrum is the cruciate sulcus The suprasplenial and the postsplenial sulcus
(sulcus cruciatus). It is very deep and runs more (sulcus suprasplenialis et sulcus postsplenialis)
or less transversely. It meets the crucial sulcus of extend between the splenial sulcus and the
the other side at the midline and in this way a dorsocaudal border of the medial surface of the
crosslike design is formed by the intersecting of hemisphere. These two sulci correspond to the
the longitudinal fissure. Rostrally and caudally, ectosplenial sulcus (sulcus ectosplenialis) of
the precruciate and postcruciate sulci (sulcus the German authors. See Ackerknecht (1943).
praecruciatus et sulcus postcruciatus) may be Ventrocaudal to the ventral end of the splenial
found. The postcruciate sulcus is more fre sulcus the occipitotemporal sulcus (sulcus oc-
484 C h ap ter 8. T he B r a in
cipitotemporalis) may be found. In some speci the dorsal and caudal margin of the cerebral
mens it joins the lateral limb of the posterior hemisphere. It forms a keystone between the
rhinal fissure ventrally. Dorsally it may connect ectomarginal sulcus laterally and the callosomar-
with the postlateral or ectolateral sulci. ginal sulcus on the medial surface of the cere
The gyri of the cerebral cortex are separated brum. The name for this large cortical area is
from each other by the various sulci and with marginal gyrus (gyrus marginalis), but it is sel
few exceptions are named for adjacent sulci. dom used, except in comparative neuromorphol
On the lateral surface of the frontal lobe the ogy. It is merely introduced here to make the
cortical area rostral to the presylvian sulcus is names for the ectomarginal and callosomarginal
called the prorean gyrus (gyrus proreus). It has sulci more meaningful.
also been referred to as the prefrontal or orbital The components of the marginal gyrus on the
gyrus. dorsal and caudal surface of the cerebral hemi
In the caudal three-fourths of the lateral as sphere are, in rostrocaudal sequence, the sigmoid
pect of the cerebrum, four major concentric ar gyrus (gyrus sigmoideus), the lateral, and the
cuate convolutions are prominent. The most postlateral gyri (gyrus lateralis et gyrus postlat-
central one is called the sylvian gyrus (gyrus eralis). The sigmoid gyrus curves around the
Sylvii). It is followed in a peripheral direction by lateral end of the cruciate sulcus, by which it is
the ectosylvian, the suprasylvian and the mar divided into the anterior and posterior sigmoid
ginal gyri (gyrus ectosylvius, gyrus suprasylvius gyri (gyrus sigmoideus anterior et gyrus sig
et gyrus marginalis) respectively. moideus posterior). The area rostrodorsal to the
The sylvian gyrus curves around the sylvian coronal sulcus where the anterior and posterior
fissure and may be divided into an anterior and sigmoid gyri meet has been referred to as the
a posterior sylvian gyrus (gyrus sylvius anterior lateral sigmoid gyrus (Bartley and Newman
et gyrus sylvius posterior). These two parts cover 1931) and was labeled gyrus coronalis by Singer
the triangularly shaped insular area (regio in- (1962). The name “lateral sigmoid gyrus” offers
sularis), which consists of cerebral cortex in the no obvious advantage in terminology (Smith
depth of the ventral portion of the sylvian fissure. 1935a), and most authors, specifically Langley
The peripheral boundary of the sylvian gyrus is (1883-84a) and Papez (1929), use the term “co
formed by the ectosylvian sulcus. ronal gyrus” for the convolution caudolateral to
The ectosylvian gyrus likewise carries the the coronal sulcus. If the anterior and posterior
name of the sulcus which bounds its concave sigmoid gyri become indented by the precruci-
side. It lies between the ectosylvian and the ate and the postcruciate sulci respectively, the
suprasylvian sulci and is divided into the ante convolutions which are formed rostrally and
rior, middle, and posterior ectosylvian gyri (gyrus caudally are referred to as precruciate and post-
ectosylvius anterior, medius, et posterior). cruciate gyri (gyrus praecruciatus et gyrus post- >
The third convolution arches around the cruciatus).
suprasylvian sulcus. Its rostral portion lies on the The lateral and postlateral gyri (gyrus later
rostrodorsal side of the anterior suprasylvian sul alis et gyrus postlateralis) lie between the longi
cus and caudolateral to the coronal sulcus. It is tudinal fissure and the sulci with their respective
referred to as the coronal gyrus (gyrus coronalis) names. If the entolateral sulcus is present, it iso
or anterior suprasylvian gyrus (gyrus suprasyl lates the entolateral gyrus (gyrus entolateralis)
vius anterior). See Langley (1883-84a). The from the medial side of the lateral gyrus.
middle suprasylvian gyrus (gyrus suprasylvius The splenial gyrus (gyrus splenialis) repre
medius) is the middle part of the third arcuate sents the marginal gyrus on the medial aspect of
convolution. It lies between the middle supra the cerebrum. It extends from the cruciate sulcus
sylvian sulcus and the lateral sulcus. The caudal caudally between the dorsal border of the hemi
segment of the third convolution is divided by sphere and the cingular and splenial sulci. The
the ectolateral sulcus into two portions. The caudodorsal and the caudal portions are set apart
posterior suprasylvian gyrus (gyrus suprasylvius as suprasplenial and postsplenial gyri (gyrus
posterior) lies on the rostrolateral side of the suprasplenialis et gyrus postsplenialis) by the
ectolateral sulcus and is bounded laterally and suprasplenial and the postsplenial sulci respec
rostrally by the posterior suprasylvian sulcus. On tively.
the medial side of the ectolateral sulcus the ecto On the medial surface of the frontal lobe the
lateral gyrus (gyrus ectolateralis) parallels the genual gyrus (gyrus genualis) lies between the
lateral and postlateral sulci which form its me genual sulcus and the ectogenual sulcus, which
dial and caudal boundaries. in turn bounds the caudal margin of the ecto
The fourth arcuate convolution extends along genual gyrus (gyrus ectogenualis).
T E LE N C E PH A LO N 485
The cortical area ventral to the genu of the sented at the end of the second week post par-
corpus callosum is the paraterminal or subcal- tum. The sulcal and gyral pattern, as it is found
hsal gyrus (gyrus paraterminalis). It is bounded in the adult, is relatively well differentiated after
caudally by the lamina terminalis, which forms the first month, while the brain’s spherical shape
the rostral wall of the third ventricle. Ventrally, will persist well beyond this time. Details on the
the paraterminal gyrus reaches the basal border early morphogenesis of the dog’s cerebral cortex
of the cerebrum. It forms part of the subcallosal may be found in publications by Morawski
or precommissural area (area subcallosa), which (1912), Rheingans (1954), Herre and Stephan
will be described with the rhinencephalon. The (1955), Meyer (1957), and Schneebeli (1958).
dorsal end of the paraterminal gyrus is in contact For additional literature referring to the brain of
with the cingular gyrus. puppies see Bekhterieff (1886), Bottazzi (1893),
The cingular gyrus (gyrus cinguli) surrounds Bikeles (1894), Bary (1898), Dollken (1898),
the corpus callosum rostrally, dorsally and cau Vogt and Vogt (1902), Ziehen (1906), Bahrs
dally. It lies between the callosal sulcus and the (1927), Himwhich and Fazekas (1941), Becker
callosomarginal sulcus, which consists of the (1952), Meyer (1952), and Perkins (1961).
genual, cingular, and splenial sulci. The caudal Histological studies on the cerebral cortex
end of the cingular gyrus blends with the para- were reported by Loewenthal (1904), Campbell
hippocampal gyrus. (1904-05, 1905), Brodmann (1905-06), Klempin
The parahippocampal gyrus (gyrus parahip- (1921), Rawitz (1926), Gurewitsch andBychow-
pocampalis) runs ventrorostrally and slightly ski (1928), Sarkissow (1929), Maspes (1932),
laterad from the splenial region and continues Florio (1943-47), C. Schwill (1951), Rheingans
into the piriform area, which will be described (1954), Volkmer (1956), and Kreiner (1961).
with the rhinencephalon. It lies between the The early work on a decerebrate dog (Goltz
medial limb of the posterior rhinal sulcus and the 1884, 1888, 1892; Langley 1883-84b; Klein
hippocampal sulcus. At its dorsal end the para 1883-84; Fritsch 1884; Holmes 1901) was fol
hippocampal gyrus extends rostrally for a short lowed by a number of experimental decerebra
distance on the ventral side cf the splenium of tions or decortications in dogs and reports about
the corpus callosum. This part has been referred dogs lacking cerebral hemispheres due to patho
to as the callosal gyrus (gyrus callosus). See Dex- logical conditions (Zeliony 1913; Dresel 1924;
ler (1932) and Seiferle (1957). Rothmann 1924; Laughton 1925-26; Rade-
The dentate gyrus (gyrus dentatus) is partly maker 1926; Poltyrev and Zeliony 1929; Papez
involuted in the depth of the hippocampal sulcus and Rundles 1938; Papez 1938; Gantt 1948;
and extends from the temporal lobe to the sple Johnson and Browne 1954).
nium of the corpus callosum. Its free border is The dog was used for the classic stimulation
visible along the concave lateral side of the hip experiments by Fritsch and Hitzig (1870), and
pocampal sulcus. Ventral to the splenium it en Ferrier (1873, 1880), as well as for the localiza
circles the callosal gyrus rostral to which it be tion studies by Munk (1881), Hitzig (1901,1901-
comes prominent as the tubercle o f the dentate 02a, 1904), Luciani (1884-85) and Luciani and
gyrus (tuberculum gyri dentati). From the tu Seppilli (1885). See also James (1890).
bercle it continues caudad as the subsplenial Investigations on the dog’s motor cortex and
flexure o f the dentate gyrus (flexura subspleni- related areas were reported by Mayser (1878),
alis gyri dentati) and blends with the gyrus fas- Lowenthal (1883), Paneth (1885), Bekhterieff
ciolaris, a transitional area between the dentate (1886), Bianchi and d’Abundo (1886), Muratoff
gyrus and the indusium griseum. The indusium (1893a), .Mingazzini (1895), Marinesco (1895),
griseum consists of a thin layer of gray matter Schukowski (1897), Demoor (1899), Hitzig
hidden in the callosal sulcus. It encircles the cor (1902-03b), Katzenstein (1908), Feliciangeli
pus callosum and reaches the paraterminal gyrus (1910), Laughton (1924, 1928), Bellucci (1929),
in the subcallosal area. Simpson (1930), Bartley and Newman (1931),
The sulcal and gyral pattern is simple at birth, Delmas-Marsalet (1932a, 1932b), Smith (1933,
as is to be expected in an altricial species. The 1935a, 1935b), Steblow (1933), Woolsey (1933),
rhinal, suprasylvian, coronal, lateral, presylvian, Asratian (1935), Poltyreff and Alexejeff (1936),
cruciate, cingular, and splenial sulci can be rec Bailey and Haynes (1940), Delgado (1948), Kel
ognized. The sylvian fissure is an open triangular logg (1949), Chusid, De Guiterrez-Mahoney, and
groove in the depth of which the insular area Robinson (1949), Goldzbrand, Goldberg, and
may be seen. Clark (1951), Morin, Poursines, and Maffre
The time of appearance of the sulci is variable, (1951), and Dimic and Nonin (1954).
but in general all major sulci and gyri are repre Woolsey (1943), Hamuy, Bromiley, and
486 C h ap ter 8. T he B r a in
Woolsey (1950), and Iwama and Yamamoto dles which connect more distant cortical areas
(1961) reported work on the somatic sensory of the cerebrum.
areas of the dog. The intralobar fibers may be divided into in-
For additional information about the occipital tracortical and subcortical fibers (von Bechterew
lobe, including the visual cortex and related con 1899; Ariens Kappers, Huber, and Crosby 1936;
nections, the following authors may be con Kuntz 1950; Ranson and Clark 1959). Theintra-
sulted: Munk (1880,1890), Berger (1900), Hitzig cortical fibers (von Bechterew 1891; Kaes 1891)
(1900a, 1900b, 1901-02b, 1902-03a, 1903), run in the deeper part of the cerebral cortex and
Probst (1902), Buytendijk (1924), Oshinomi are not visible macroscopically. The subcortical
(1930), Marquis (1932a, 1932b), Culler andMet- fibers connect adjacent and close-by gyri by
tler (1934), Rosenzweig (1935a, 1935b, 1935c), curving around one or several intervening sulci.
Marquis and Hilgard (1936), Thauer and Stuke In doing so they form U-shaped loops referred
(1940), Wing and Smith (1942), and Iwai (1961). to as arcuate fibers (fibrae arcuatae cerebri). In
Details on the acoustic system are contained addition to the fibers which unite different gyri,
in papers by Kalischer (1907), Katz (1932), short association fibers can be found to connect
Buytendijk and Fischel (1933), Culler and Met- parts within certain gyri. These fibers may be
tler (1934), Girden (1938), Tunturi (1944,1945, called intragyral fibers (Meyer 1957). Arcuate
1948,1950,1952), Allen (1945), Mosidze(1960), and intragyral fibers can be demonstrated with
Chorazyna and Stepien (1961), and Sychowa out difficulty if the cerebral cortex is removed.
(1961a). Decortication and dissection of the fibers
For studies on the relation of the cerebral cor are relatively easy if the preparation method
tex to autonomic reactions see Kremer (1947, of Klingler (1935) is used. See also Ludwig
1948), Eliasson, Lindgren, and Uvnas (1952), (1935), Meyer (1954), and Ludwig and Klingler
Okinaka, Nakamura, Tsabaki, Kuroiwa, and (1956).
Toyokura (1953), and Brutkowski, Fonberg, and Among the interlobar fiber systems which are
Mempel (1961). described in the literature, the cingulum is the
No list of references on the dog’s cerebral cor only long fiber bundle whose existence and asso
tex would be complete without mentioning the ciative function have never been questioned. It
classic work on conditioned reflexes (Pavlov is the most distinct structure on the medial sur
1927; Pawlow 1952) and at least some of the face of the decorticated hemisphere (Fig. 8 - 8 ).
most recent papers in this interesting field of Its fibers extend from the genual and paratermi-
investigations (Stepien, Stepien, and Konorski nal gyri through the cingular gyrus to the callosal
1961; Zemicki 1961; Zernicki and Santibanez and parahippocampal gyri. Degeneration stud
1961; Sheiman 1961; Dumenko 1961). ies have shown that some of its fibers extend only
For literature on the blood vascular system re partway along the bundle, while others extend
lated to the cerebral cortex consult Andreyev along the entire length.
(1935a, 1935b), Gross (1939), and Billenstien The subcallosal bundle (fasciculus subcallo-
(1953). sus), which may also be referred to as superior
The white matter of the cerebrum basically fronto-occipital or superior occipito-frontal bun
consists of two types of fiber systems: the corti- dle, was first described in the dog (Muratoff
cocortical fibers and the projection fibers. 1893a, 1893b). It can be traced as a distinct en
The corticocortical fibers have both their ori tity along the peripheral wall of the lateral ven
gins and terminations in the cerebral cortex. tricle. Rostrally, it lies dorsal to the caudate nu
They are subdivided into association fibers and cleus. The connections of the subcallosal bundle
commissural fibers. The association fibers con are not exclusively between cortical areas along
nect different cortical areas in the same cerebral the course of the bundle. Fibers are also received
hemisphere. The commissural fibers extend be from the caudate nucleus, which would indicate
tween homologous areas on opposite sides of the that it is not a pure association system, but a
cerebrum. mixed fiber bundle (Figs. 8-10, 8-11).
The projection fibers either originate or ter The other long association bundles which are
minate in the cerebral cortex. Their respective referred to in the literature, the superior longi
other ends are found within lower centers of the tudinal bundle (fasciculus longitudinalis supe
basal nuclei, the brain stem, or the spinal cord. rior), the uncinate bundle (fasciculus uncinatus),
The association fibers (Figs. 8 - 8 through 8 - the inferior occipito-frontal bundle (fasciculus
11 ) are differentiated into short or intralobar occipito-frontalis inferior), and the inferior longi
fibers and long or interlobar fiber systems. The tudinal bundle (fasciculus longitudinalis infe
intralobar fibers unite closely related gyri. The rior) are of questionable nature in the dog.
interlobar fibers form more or less distinct bun A group of fibers dorsomedial to the insular
T E LE N C E PH A LO N 487
F ig . 8 -3 . Medial surface of the right cerebral hem isphere and lateral surface of the brain stem.
Ectogenual sulcus 12. Suprasplenial sulcus 25 . Accessory cuneate nucleus
Ectogenual gyrus 12'. Suprasplenial gyrus 26 . External arcuate fibers
Genual sulcus 13 . Postsplenial sulcus 27 . Cochlear nuclei
Genual gyrus 13'. Postsplenial gyrus 28 . Trapezoid body
Genu of corpus callosum 14 . Cut surface between cerebrum and brain 29 . Lateral lemniscus
Cingulate sulcus stem 30 . Transverse fibers of pons
Cingulate gyrus 15 . Lateral geniculate body 31 . Brachium of inferior colliculus
Callosal sulcus 1 6 . Superior colliculus 32 . Transverse peduncular tract
Cruciate sulcus 17 . Medial geniculate body 33 . Crus cerebri
Body of corpus callosum 1 8 . Inferior colliculus 34 . Left optic tract
Lesser cruciate sulcus 19 . Arbor vitae cerebelli 35. Optic chiasm
Splenium of corpus callosum 20 . Superior cerebellar peduncle 36 . Anterior commissure
Splenial sulcus 2 1 . Inferior cerebellar peduncle 37 . Paraterminal gyrus
Splenial gyrus 22 . Middle cerebellar peduncle 3 8 . Septum pellucidum
Posterior horizontal ramus of splenial 2 3 . Fasciculus cuneatus I I . Optic nerve
sulcus 24 . Spinal tract of trigeminal nerve I I I . Oculomotor nerve
I V . Trochlear nerve
area may be interpreted as the superior longi which, according to Ariens Kappers, Huber, and
tudinal bundle. This fiber group extends along Crosby (1936), can be demonstrated in the dog
the dorsal edge of the claustrum (described with by gross dissection, nor of the inferior longitu
the basal nuclei). The uncinate bundle which dinal bundle, could be found by Meyer (1957).
connects the frontal and temporal lobes may be For additional information on association fibers
represented by a group of fibers at the junction see Obersteiner and Redlich (1902), Piltz (1902),
of these lobes. There is no proof, however, that Poljak (1927), Poltyrew and Zeliony (1930),
they actually form corticocortical connections. Poltyreff (1936), and Meyer (1957).
No trace of the inferior occipito-frontal bundle, The commissural fibers of the cerebrum form
488 Chapter 8. T he B r a in
the corpus callosum, the anterior commissure, pus callosum (Fig. 8-14). See Edinger (1911).
and the hippocampal commissure. The hippocampus will be described with the
The corpus callosum (Figs. 8-3, 8 - 8 , 8-12, 8 - rhinencephalon.
16) is the largest commissure of the brain. It con The projection fibers of the neocortical areas
nects homologous neocortical areas of the two form the internal capsule (capsula interna). A
sides. Its rostral portion is referred to as the genu phylogenetically older projection system is that
(genu corporis callosi); its middle is called the of the fornix.
body (truncus corporis callosi); and its caudal The internal capsule (see Loewenthal 1886)
part is the splenium (splenium corporis callosi). consists of an anterior crus (crus anterius capsu-
The corpus callosum is separated from the cingu- lae internae), a posterior crus (crus posterius
lar gyrus by the callosal sulcus. In the center, capsulae internae), and the genu (genu capsulae
where all its fibers meet, the corpus callosum internae), where the two crura meet (Figs. 8-13,
forms the roof of the lateral ventricles. The lat 8-15; Plates 1, 10). The anterior crus passes
eral ventricles of the two sides are separated through the corpus striatum. The posterior crus
from each other along the midline by the septum extends along the lateral aspect of the dien
pellucidum (Figs. 8 - 8 , 8-14). The fibers of the cephalon. The fibers of the internal capsule
corpus callosum, which connect the lateral con either originate in the cerebral cortex and de
volutions of the cerebral hemispheres, are inter scend to lower centers or take origin in lower
sected by the fibers of the internal capsule. More centers and carry impulses to the cerebral cor
details on the corpus callosum are contained in tex. At the dorsolateral edge of the lateral ven
papers by von Koranyi (1890), Martin (1893, tricle they are intersected by the fibers of the
1894a, 1894b), Muratoff (1893b), Muratow corpus callosum. The resulting intermingled
(1893), Probst (1901b), Janischewski (1902), mass of white matter has the appearance of a
Bykoff (1925), Lindberg (1937), and Curtis crown of rays in which it is difficult to determine
(1940). the origin of individual fibers and fiber groups. It
The anterior commissure (commissura ante is called the corona radiata (Fig. 8-13).
rior) forms connections between olfactory areas The fornix connects the hippocampus with
of the cerebrum (Fig. 8-14; Plate 10). On me centers in the ventral part of the diencephalon.
dian sections (Figs. 8-12, 8-13) it may be seen It will be described in more detail with the struc
caudal to the paraterminal gyrus and rostral to tures of the rhinencephalon and the diencepha
the diencephalon, about halfway between the lon.
corpus callosum and the base of the brain. It The basal nuclei (Plate 1) consist of three
consists of a strong rostral portion and a consid nuclear masses, the corpus striatum, the claus-
erably smaller caudodorsal part. See Sander trum, and the amygdaloid body. These struc
(1866). The rostral part connects the two olfac tures are frequently referred to as a basal ganglia.
tory bulbs, which are located in the rostroven- The corpus striatum is the largest nuclear
tral portion of the rhinencephalon. On their way mass among the basal nuclei. It is subdivided by
rostrolaterad the fibers of the rostral part of the the anterior crus of the internal capsule into a
anterior commissure are embedded in the most medial portion, the caudate nucleus (nucleus
ventromedial portion of the caudate nucleus caudatus), and a lateral portion, the lentiform
(Fig. 8-16). As these fibers run rostrad to the ol nucleus (nucleus lentiformis). The lentiform nu
factory bulb they become ipterwoven with pro cleus in turn is further subdivided by the medul
jection fibers of the internal capsule. The caudal lary lamina (lamina medullaris) into the more
part of the anterior commissure extends between medially located globus pallidus and the more
the piriform areas of the two sides. The piriform lateral putamen.
area (Fig. 8-5) is an olfactory cortical region at The caudate nucleus protrudes from the lat
the temporal pole of the cerebrum. From the eral side into the rostral part of the lateral ven
midline the fibers of the caudal portion of the tricle (Figs. 8-10, 8-11, 8-16). Its large rostral
anterior commissure run slightly rostrad. Then portion is called the head o f the caudate nucleus
they curve laterad and become intermingled (caput nuclei caudati). The caudally tapering
with the fibers of the internal capsule. After leav part forms the tail (cauda nuclei caudati). The
ing the area of the internal capsule they turn middle portion may be referred to as the body o f
caudolaterad and ventrad to reach the piriform the caudate nucleus (corpus nuclei caudati).
area. Rostrally, across the fibers of the internal cap
The hippocam pal commissure (commissura sule, the gray matter of the caudate nucleus and
fornicis) unites the hippocampi of either side of the putamen is continuous. The head of the
and is located ventral to the splenium of the cor caudate nucleus is pierced ventrally by the ante-
T EL E N C E P H A L O N 489
nor commissure. The tail of the caudate nucleus anterior and posterior crura of the internal cap
ends ventral to the beginning of the caudal sule. Laterally, the lentiform nucleus is bounded
fourth of the corpus callosum. by a thin layer of white matter, the external cap
The globus pallidus, the smaller, medial por sule (capsula externa).
tion of the lentiform nucleus, lies in a more ven The claustrum is located lateral to the external
tral location. Its medial boundary is formed by capsule and extends along the dorsolateral mar
the genu of the internal capsule. Laterally, it is gin of the putamen. The very thin layer of white
separated from the putamen by the medullary matter that separates it from the cortex is re
lamina. ferred to as the extreme capsule (capsula ex
The putamen extends along the lateral side of trema).
the globus pallidus and is in contact with the The am ygdaloid body (Fig. 8-16) consists of
490 Chapter 8. T he B r a in
a number of nuclear masses which occupy the factory striae, the piriform area, and the optic
lateral wall and the rostral extremity of the ven tract (Fig. 8-5). The optic tract will be described
tral horn of the lateral ventricle. Ventrally, this with the diencephalon. The name “perforated
gray matter is continuous with the cerebral cor substance” is appropriate because a large num
tex of the piriform area at the temporal pole of ber of small blood vessels pass through the area
the cerebrum. The amygdaloid body is custom in order to reach the basal nuclei. Along the cau
arily listed with the basal nuclei for morpholog dal border of the anterior perforated substance
ical reasons. On the basis of its connections it a band of lighter color, the diagonal band (taenia
may be included among the structures of the diagonalis), extends to the medial side where it
rhinencephalon. continues into the paraterminal gyrus in the sep
For additional information on the basal nuclei tal area. The remaining part of the anterior per
the following authors may be consulted: Schuller forated substance is known as the olfactory tri
(1902), de Vries (1910), Berlucci (1927), Mittel- gone (trigonum olfactorium). The rostral portion
strass (1937), Mettler and Goss (1946), Fank- of the trigone may be prominent and is custom
hauser (1947), A. Schwill (1951), Tenerowicz arily referred to as the olfactory tubercle (tuber
(1960), and Kreiner and Marksymowicz (1962). culum olfactorium).
The rhinencephalon, or olfactory part of the The piriform area (area piriformis) is a con
brain, is separated from the remaining telen spicuous pear-shaped cortical region ventral to
cephalon by the anterior and posterior rhinal the middle of the rhinal sulcus at the temporal
fissures. It consists of the olfactory bulbs, the ol pole of the cerebrum (Fig. 8 - 6 ). It belongs to the
factory tracts and gyri, the anterior perforated phylogenetically older paleopallium. The lateral
substance, the piriform area, the amygdaloid olfactory gyrus unites with the rostrodorsal por
body, the hippocampus and fornix, the septal tion of the piriform area. Caudally, the piriform
area, and the anterior commissure. area blends with the parahippocampal gyrus.
The olfactory bulbs (bulbi olfactorii) are The gray matter of the piriform area is continu
rounded ventrorostral projections (Fig. 8-4) ous with the deeply located amygdaloid body.
which lie in the cribriform fossa of the ethmoid The am ygdaloid body was previously referred
bone. Through the openings in the cribriform to in the description of the basal nuclei. Several
plate they receive the fibers which constitute the nuclei of the amygdaloid complex receive fibers
first cranial or olfactory nerves. Inside the olfac from the olfactory bulbs and the anterior com
tory bulb there is a cavity which connects with missure. There are also connections with the
the lateral ventricle by way of an olfactory stem septal area by way of the diagonal band. The
(Fitzgerald 1961). This olfactory stem may be main efferent pathway is a clearly defined fiber
occluded. bundle, the stria terminalis (Fig. 8-16).
The olfactory tract (tractus olfactorius) con The stria terminalis arises from the rostrome-
tinues the olfactory bulb caudally (Fig. 8-5). dial part of the amygdaloid body and runs dorsad
After a short distance it bifurcates to form the and caudad along the optic tract to the tail of the
lateral and the medial olfactory striae (stria ol- caudate nucleus. From there the stria terminalis
factoria lateralis et medialis). The olfactory passes rostrad and ventrad in a groove between
tract and the lateral and the medial olfactory the caudate nucleus and the thalamus and ends
striae are accompanied by gray matter, the lat at the base of the brain in the preoptic and hypo
eral and the m edial olfactory gyri (gyrus olfac thalamic region.
torius lateralis et medialis). The hippocampus (Fig. 8-14; Plate 10) is a
The lateral olfactory gyrus lies between the long curved structure which projects into the
anterior rhinal fissure dorsally and the olfactory caudal part of the lateral ventricle. It consists of
tract and lateral olfactory stria ventromedially. the cortical region, which represents the archi-
Caudally, the lateral olfactory gyrus blends with pallium, the phylogenetically oldest part of the
the piriform area, which also receives the fibers cerebral cortex. It is involuted along the hippo
of the lateral olfactory stria. campal sulcus and contacts the lateral wall of
The medial olfactory gyrus extends along the the dentate gyrus. The ventricular surface of
medial side of the medial olfactory stria and joins the hippocampus is covered with a thin layer of
the septal area. The fibers of the medial olfactory white matter, the alveus, which consists of the
stria contribute to the anterior commissure and efferent and commissural fibers of the hippo
connect with the septal nuclei of the septal area. campus. The commissural fibers cross the mid
The anterior perforated substance (substantia line ventral to the splenium of the corpus callo
perforata anterior) is a more or less triangular sum. They constitute the hippocampal commis
area bounded by the lateral and the medial ol sure or the commissure of the fornix (Fig. 8-14).
T elen ceph a lo n 491
The remaining major part of the fibers of the reported by McCotter (1913). See also Ariens
alveus gathers along the concave edge of the Kappers, Huber, and Crosby (1936), Lazorthes
hippocampus and forms the fimbria o f the hip (1943), and Crosby, Humphrey, and Lauer
pocampus (fimbria hippocampi). (1962).
The fornix blends with the fimbria of the hip The septal nuclei are nuclear masses of appre
pocampus at the rostrodorsal end of the hippo ciable size which protrude from the medial wall
campus. It forms the rostral continuation of an into the lateral ventricle. They are in close spatial
efferent hippocampal fiber system, which con relation with the subcallosal area and have fiber
nects the hippocampus with the septal area, and connections with the hippocampus. The fibers
with the mammillary bodies in the hypothalamus received from the hippocampus were described
(Fig. 8-13). This efferent hippocampal fiber sys with the fornix. Those leaving the septal nuclei
tem parallels the course of the stria terminalis, for the hippocampus are the septal fibers which
from which it is separated by the lateral ventricle. are intermingled with the fibers of the body of
The part of the fornix rostral to the hippocam the fornix.
pus is referred to as the body o f the fornix (cor The septum pellucidum is a thin, small mem
pus fornicis). It contributes the major portion of branous area which is bounded by the concave
the longitudinal fibers along the ventral aspect side of the genu of the corpus callosum, the for
of the body of the corpus callosum. nix, and by the septal nuclei. In some species the
On its ventrorostral course the fornix forms septum consists of a double layer with a cavity,
two roundish bundles, the columns o f the fornix the cavum septi pellucidi, between. There is usu
(columnae fornicis), which become isolated and ally no cavity of the septum pellucidum in the
bypass the anterior commissure caudally (Fig. dog. According to Elliot Smith (1895-96) and
8 - 6 ). This portion of the fornix may be referred Ziehen (1906), it is rare or absent. Thompson
to as the postcommissural fornix. Under cover (1932) describes a cavum septi pellucidi which
of the gray matter of the ventral part of the di is open rostrally.
encephalon the columns of the fornix reach the The anterior commissure (Fig. 8-14) was de
mammillary bodies in the hypothalamus. scribed with the commissural fibers of the cere
The remaining fibers of the fornix enter the bral white matter. The pars anterior, which con
septal area, partly bypassing the anterior com nects the olfactory bulbs of the two sides, re
missure rostrally, as precommissural fornix ceives its fibers largely from the medial olfactory
(Fig. 8 - 12 ). stria. The fibers of the pars posterior reach the
The longitudinal fibers ventral to the body of piriform areas of the two sides by traversing the
the corpus callosum, which do not belong to the internal capsule.
fornix, are septal fibers and carry impulses from For additional information on the rhinenceph
the septal area to the hippocampus. alon see Sander (1866), Ganser (1879), Probst
The term septal area (area septalis) refers to (1901a), Read (1908), Allen (1937, 1944, 1948),
the basal gray matter bounded by the genu of the Schultz (1939), Kruger (1942), Fox (1943), Fox
corpus callosum, the anterior commissure, the and Schmitz (1943), Nilges (1944), Palionis
lamina terminalis, and the olfactory tubercle. (1950), Kaada (1951), Takahashi (1951), Becker
This area consists of the subcallosal area, includ (1952), De Groot (1958-59), Kalinina (1961).
ing the paraterminal gyrus, and the septal nuclei Details on the subfornical organ of the dog may
(nuclei septi). The septum pellucidum, which be found in papers by Pines (1926), Pines and
separates the two lateral ventricles, is not in Maiman (1928), Heidreich (1931), Cohrs (1936),
cluded in the septal area. For topographical and Akert, Potter, and Anderson (1961).
reasons, however, it is described with the struc The remaining part of the brain caudal to the
tures of the septal area. telencephalon, with the exception of the cere
The subcallosal or precommissural area com bellum, forms the brain stem (Fig. 8-17). It con
prises the medial part of the cerebrum ventral to sists of ascending and descending fiber tracts
the corpus callosum and rostral to the anterior and of nuclear masses which are connected
commissure. either with the tracts or with cranial nerves. A
The paraterminal gyrus was described with large area of the brain stem contains extended
the convolutions of the medial cerebral surface. gray matter which is diffusely intermingled with
It lies immediately rostral to the lamina termi fibers. It appears as a complicated network and
nalis and forms part of the subcallosal area. The is referred to as the reticular form ation (formatio
nervus terminalis enters the brain in this region. reticularis). See Brodal (1957) and Valverde
The first dissection of this nerve in the dog was (1961).
D ie n c e p h a l o n 493
II 10
F ig . 8-6. Ventral view of the cerebrum.
1. Olfactory bulb 8. Parahippocampal gyrus 15. Lateral limb of posterior rhinal sulcus
2. Olfactory trigone 9. Hippocampal sulcus 16. Medial limb of posterior rhinal sulcus
3. Diagonal band 10. Callosal gyrus 17. Posterior rhinal sulcus
4. Anterior commissure 11. Subsplenial flexure of dentate gyrus 18. Piriform area
5. Columns of fornix 12. Tubercle of dentate gyrus 19. Anterior rhinal sulcus
6. Cut surface between cerebrum and brain stem 13. Occipitotemporal sulcus 20. Olfactory tubercle
7. Fimbria of hippocampus 14. Dentate gyrus
dorsal midline of the diencephalon (Fig. 8-12). striae medullares of either side cross the midline
It consists of the stria medullaris thalami, the and form the habenular commissure.
habenular nuclei (nucleus habenulae), the ha- The thalamus, sometimes also referred to as
benular commissure (commissura habenu- the dorsal thalamus, is a wedge-shaped mass of
larum), and the pineal body (corpus pineale). gray matter which is located ventral and lateral
The pineal body (Fig. 8-12) is very small in to the epithalamus, between the posterior limb
the dog. See Venzke and Gilmore (1940). It is of the internal capsule and the midline (Fig. 8 -
about a millimeter long and projects caudad from 13).
the dorsocaudal end of the midline portion of Along the midline the thalami of the two sides
the diencephalon. Rostral to it the habenular are separated from each other by the third ven
nuclei may be seen as small, rounded nuclear tricle, except for a round area in the center of
masses. The stria medullaris thalami is a thin the ventricle, where the gray masses of the two
bundle of white matter which partly encircles sides adhere to each other. This area is called the
the diencephalon along the midline. It connects interthalamic adhesion (adhaesio interthalam-
olfactory centers at the base of the brain with ica).
the habenular nuclei. Some of the fibers of the The third ventricle (ventriculus tertius) lies in
D ie n c e p h a l o n 495
the median plane and encircles the interthalamic groove. The stria terminalis, which was described
adhesion (Fig. 8-12). It connects with the lateral with the rhinencephalon, partly encircles the
ventricles in the cerebral hemispheres by way thalamus along the bottom of this groove. On
of the interventricular foram ina (foramen inter- the concave side of the stria terminalis the thala
ventriculare) and opens caudally into the cere mus meets the internal capsule. This dose spatial
bral aqueduct (aqueductus cerebri) of the mes relation of the internal capsule and the thalamus
encephalon. (Fig. 8-13) has functional significance. The
Rostrally and dorsolaterally, the thalamus is fibers which leave or enter the thalamus on its
separated from the caudate nucleus by a deep lateral side form the thalamic radiation (fasciculi
496 Chapter 8. T he B r a in
subsidiary nuclei. Also included in the hypo 1900b, 1903a), Morgan (1927), Glorieux (1929),
thalamus is the neurohypophysis, which is con Rioch (1929a, 1929b, 1931a, 1931b), Oshinomi
tinuous with the tuber cinereum (Fig. 8-12). (1930), Hammouda (1933), Papez (1938), Papez
The optic chiasm belongs to the visual system, and Rundles (1938), and Sychowa (1961b). For
and its relation to the hypothalamus is largely comparative studies see Jasper and Ajmone-
topographical. It is formed by the convergence Marsan (1954).
and partial decussation of the two optic nerves, Numerous articles may be consulted on the
the second pair of cranial nerves, which carry hypothalamus (Ramirez-Corria 1927a, 1927b;
visual impulses from the retina to the level of the Griinthal 1929; Groschel 1930; Morgan 1930a,
optic chiasm. In the chiasm the optic nerve 1930b; Papez 1932; Roussy and Mosinger 1935;
fibers are redistributed to form the optic tracts, Collin and Grognot 1938; Rose 1939; Bonvallet,
which run to the lateral geniculate bodies. As a Dell, and Stutinsky 1949; Strom 1950; Jewell
result of the partial decussation each optic tract 1953; Knoche 1953, 1957; Frandson 1955;
receives fibers from the nasal half of the opposite Sloper 1955; Bleier 1961), the hypophysis (Basir
retina and from the temporal retinal portion of 1932; Otten 1943; Stutinsky, Bonvallet, and Dell
the same side. For details see Teljatnik (1897), 1949, 1950; Goldberg and Chaikoff 1952; Smith,
Arey, Bruesch, and Castanares (1942), Areyand Calhoun, and Reineke 1953; Knoche 1953;
Gore (1942), Seiferle (1949), and Bishop and Latimer 1954; Scharrer and Frandson 1954;
Clare (1955). Fujita 1957), and their relationship (Mogilnitzky
The supraoptic nucleus is one of the nuclei of 1928a, 1928b; Roussy and Mosinger 1933; Gagel
the supraoptic portion of the hypothalamus and and Mahoney 1933; Heinbecker and White
lies dorsal and slightly lateral to the optic chiasm. 1941; Pickford and Ritchie 1945; Stutinsky 1949,
The tuber cinereum is a gray mass between the 1958; Knoche 1952; O’Connor 1952; Scharrer
optic chiasm and the mammillary bodies. It ends and Wittenstein 1952; Scharrer 1954; Laqueur
ventrally in a tube-shaped process, the infundib- 1954; Hagen 1957).
ulum, by means of which the neurohypophysis
is attached to the tuberal portion. The mam
millary bodies (corpora mammillaria) are two MESENCEPHALON
spherical eminences between the cerebral
peduncles. They are fused at the midline, and The mesencephalon (Plate 3) is the portion of
lie rostral to the posterior perforated substance the brain stem between the diencephalon ros
in the interpeduncular space. The mammillo- trally and the pons and the cerebellum caudally.
thalamic tracts originate from the mammillary The dorsal part of the mesencephalon is referred
bodies and connect with the anterior nuclear to as the tectum (tectum mesencephali) and pro
groups of the thalamus. vides the roof of the mesencephalon. The ventral
The subthalamus (Plate 2) is a transitional portion of the mesencephalon is formed by the
zone between the thalamus and the midbrain. cerebral peduncles (pedunculi cerebri). The
It lies caudal and lateral to the hypothalamus. cerebral aqueduct (aqueductus cerebri), a tube
Laterally, it is bounded by the fibers of the like canal which is surrounded by the central
internal capsule, which continue caudad into gray matter (substantia grisea centralis), passes
the mesencephalon as the crus cerebri. The sub through the mesencephalon between the tectum
thalamus contains nuclear masses such as the and the cerebral peduncles (Fig. 8-12). It con
subthalamic nucleus (nucleus subthalamicus) nects the third ventricle in the diencephalon
and is traversed by efferent fiber systems from with the fourth ventricle in the metencephalon
the basal nuclei. These efferent fiber systems and the rostral portion of the myelencephalon.
connect the lentiform nucleus with nuclei of the For special structures on the surface of the
thalamus and subthalamus. They pass through aqueduct see Friede (1961).
and around the internal capsule. The fibers The tectum of the mesencephalon (Fig. 8-17)
which loop around the ventral side are referred consists of the tectal lamina (lamina tecti), which
to as the ansa lenticularis. Those intermingled is occupied by a rostral and a caudal pair of
with the internal capsule form the fasciculus rounded dorsal eminences. The rostral pair is
lenticularis. The fasciculus subthalamicus en called the superior (rostral) colliculi, the caudal
ters the subthalamic nucleus. pair, the inferior (caudal) colliculi. Jointly these
For more detailed information on the di structures are referred to as the quadrigeminal
encephalon of the dog the reader is referred to bodies (corpora quadrigemina). Their connec
the papers by Forel (1872), Probst (1900a, tions with the brain stem, the brachia of the
20 |9 |8 17
F ig . 8 - 9 . L a te ra l v iew o f a d e c o rtic a te d le ft ce re b ra l h em isp h ere.
6
F ig . 8-11. Lateral view of a deep dissection of the left cerebral hemisphere.
superior and inferior colliculi, are included in the connections with the habenular nucleus by way
tectum. of the habenulointerpeduncular tract or fascicu
Each superior colliculus (colliculus superior) lus retroflexus (Fig. 8-13).
forms a spherical elevation adjacent to the The fibers of the crus cerebri include cortico
superior colliculus of the other side. It receives spinal, corticonuclear (corticobulbar), and corti
fibers from the optic tract by way of the brachi copontine fibers. All these fibers originate from
um o f the superior colliculus (brachium colliculi the cerebral cortex, descend through the internal
superioris). The superior colliculi serve as centers capsule and the crus cerebri and continue to the
for visual reflexes. Within the substance of the pons or other lower levels. They form the corti
tectum the superior colliculi of the two sides are cospinal tract (tractus corticospinalis), the corti
connected with each other by the commissure conuclear tract (tractus corticonuclearis), and
of the superior colliculi. the corticopontine tract (tractus corticopon-
Each inferior colliculus (colliculus inferior) is tinus). The fibers of the corticospinal tract, after
a bulbous protuberance in the caudolateral part partial decussation, enter the gray matter of the
of the tectum. They are somewhat smaller than spinal cord at various levels. They send impulses
the superior colliculi and do not meet along the to the motor nerve cells in the ventral gray
midline. A band of white matter, the commissure column, apparently by way of interneurons
of the inferior colliculi, connects the two inferior situated in the intermediary zone and in the
colliculi. It is visible from the outside as it crosses ventral part of the dorsal column (Szentagothai-
the midline immediately caudal to the superior Schimmert 1941). The fibers of the corticonu
colliculi (Fig. 8-17). The inferior colliculi serve clear tract are distributed bilaterally to the motor
as centers for acoustic reflexes. Each inferior nuclei of the brain stem (Crosby, Humphrey,
colliculus receives acoustic impulses by way of and Lauer 1962). The corticospinal and the corti
the lateral lemniscus (lemniscus lateralis), which conuclear fibers constitute the tracts of the
originates from the cochlear nuclei and the pyramidal system.
superior olive in the myelencephalon and sweeps The substantia nigra is a wide platelike nu
rostrad and dorsad into the inferior colliculus clear mass which extends throughout the mes
(Fig. 8-15). In addition to serving as acoustic encephalon. Concerning its pigmentation see
reflex centers, the inferior colliculi convey Marsden (1961). It lies dorsomedial to the crus
acoustic impulses to the medial geniculate bodies cerebri and ventrolateral to the tegmentum. It
of the thalamus by way of the brachia of the has numerous connections with the globus pal
inferior colliculi. On each side the brachium o f lidus, the hypothalamus, the subthalamus, the
the inferior colliculus (brachium colliculi in- red nucleus, the interpeduncular nucleus, and
ferioris) extends rostroventrad from the lateral the reticular formation.
aspect of the inferior colliculus along the ventro The tegmentum is the dorsal portion of the
lateral edge of the superior colliculus to the cerebral peduncles. It contains a number of
medial side of the medial geniculate body. nuclei (Brown 1943b, 1944) and ascending as
The cerebral peduncles (pedunculi cerebri) well as descending fiber bundles. It also includes
form the part of the mesencephalon ventral to part of the reticular formation of the brain stem.
the cerebral aqueduct. See Brown (1943a). Each The red nucleus (nucleus ruber) is the most
cerebral peduncle consists of a dorsal part, the prominent of the nuclear masses of the tegmen
tegmentum, and a ventral part, the crus cerebri tum. It is rounded and lies in the rostral portion
(basis pedunculi, B.N.A.). The tegmentum and of the mesencephalon ventral to the superior
the crus cerebri are separated from one another colliculi. It receives fibers from the opposite
by a mass of gray matter, the substantia nigra. half of the cerebellum by way of the superior
The crus cerebri (Fig. 8-5) is a large group of cerebellar peduncle (brachium conjunctivum,
descending fibers at the base of the mesencepha B.N.A., or pedunculus cerebellaris superior,
lon. It becomes visible as it emerges from the P.N.A.), which also transmits cerebellar fibers to
caudal side of the optic tract halfway between the ventral lateral nucleus of the thalamus. The
the optic chiasm and the lateral geniculate body. fibers of the superior cerebellar peduncles cross
Rostrolaterally, it lies adjacent to the medial the midline in the decussation o f the superior
geniculate body (Fig. 8-3). Caudally, the crura cerebellar peduncles (decussatio pedunculorum
cerebri of the two sides converge toward the cerebellarium superiorum), which lies in the
midline and form the lateral boundaries of the caudal portion of the tegmentum of the mesen
interpeduncular space, which includes the pos cephalon ventromedial to the inferior colliculi.
terior perforated substance and the interpedunc For details see von Monakow (1909, 1910),
ular nucleus. The interpeduncular nucleus has Fuse (1919), and Yamagishi (1935).
M eten ceph a lo n 501
The nucleus of the oculomotor nerve or third Two major descending fiber systems originate
cranial nerve (nucleus nervi oculomotorii) is in the mesencephalon. One consists of the tecto
located ventral to the cerebral aqueduct at the spinal and tectonuclear tracts (tractus tecto-
level of the superior colliculi. The fibers of the spinalis et tractus tectonuclearis); the other of
oculomotor nerve sweep ventrad through the the rubrospinal tract (tractus rubrospinalis). See
tegmentum and emerge in the interpeduncular also Ogawa and Mitomo (1938).
space. The lateral fibers of the oculomotor nerve The tectospinal and tectonuclear tracts arise
traverse the medial portion of the red nucleus from the tectum and transmit impulses which
and the most medial part of the substantia nigra. involve motor neurons at lower levels in visual
The oculomotor nerve (Figs. 8-3, 8-5) sends and acoustic reflexes. Before descending through
motor fibers to all the extrinsic muscles of the the brain stem, the tectospinal and tectonuclear
eyeball except those innervated by the fourth tracts cross to the opposite side through the
and the sixth cranial nerves and supplies para dorsal tegmental decussation, which lies in the
sympathetic fibers to the intrinsic muscles of the dorsal portion of the mesencephalic raphe ven
eye. For additional information on the oculo tral to the medial longitudinal bundle and rostral
motor nucleus compare Frank (1930) and Szen- to the decussation of the superior cerebellar
tagothai (1942). peduncles.
The nucleus of the trochlear or fourth cranial The rubrospinal tract begins in the red
nerve (nucleus nervi trochlearis) lies close to the nucleus and sends fibers to motor neurons at
caudal extremity of the oculomotor nucleus at lower levels. See Fuse (1920b). It crosses the
the level of the inferior colliculi. The trochlear midline in the ventral tegmental decussation of
nerves of the two sides cross in the anterior the mesencephalon, at the level of the dorsal
medullary velum (Fig. 8-17), which forms the tegmental decussation, and descends through
roof over the caudal opening of the cerebral the tegmental portion of the pons and the reticu
aqueduct and the rostral part of the fourth lar formation of the medulla oblongata to the
ventricle. They emerge from the dorsal aspect of lateral funiculus of the spinal cord.
the mesencephalon immediately caudal to the The m esencephalic tract and nucleus o f the
inferior colliculi (Fig. 8-3) and supply motor trigeminal nerve (tractus mesencephalicus nervi
innervation to the dorsal oblique muscles of the trigemini et nucleus tractus mesencephalici
eyes. nervi trigemini) are associated with the trigem
The m edial longitudinal bundle (fasciculus inal nerve in the caudal portion of the pons, but
longitudinalis medialis), the phylogenetically are topographically located ventrolateral to the
oldest distinct fiber bundle of the tegmentum of central gray matter of the mesencephalon. For
the mesencephalon, runs along the midline im details see Thelander (1924), Schneider (1928),
mediately ventral to the central gray matter. It and Sheinin (1930). The mesencephalic tract
includes fibers which interrelate the vestibular carries proprioceptive impulses from the tri
nuclei in the rostral part of the myelencephalon geminal nerve to the mesencephalic nucleus.
with the nuclei of the cranial nerves which inner Additional information on the mesencephalon
vate the muscles of the eyeball. The vestibular may be obtained from the experimental work by
nuclei belong to the vestibular system, which is Probst (1903b). For the blood supply of the
responsible for the sensations of head position mesencephalon the reader is referred to Olivieri
and head movement. Among the cranial nerves (1946).
which supply motor fibers to the muscles of the
eye, the oculomotor and the trochlear nerves METENCEPHALON
have been covered. The abducens or sixth
cranial nerve, which innervates the lateral rectus The metencephalon lies between the mesen
and the retractor bulbi muscles, will be de cephalon and the myelencephalon or medulla
scribed with the myelencephalon. oblongata. It has two components. The ventral
Other ascending fiber systems of the mesen subdivision, the pons, belongs to the brain stem.
cephalic tegmentum include the m edial lemnis The dorsal metencephalon develops into the
cus and the trigeminal lemniscus, which were cerebellum.
mentioned in the description of the thalamus as The pons (Figs. 8-4, 8-5; Plate 4) consists of
entering the ventral posterolateral and the ven a ventral or basilar portion (pars basilaris pontis)
tral posteromedial nuclei respectively. In their and a dorsal or tegmental part (pars dorsalis
course through the mesencephalon these tracts pontis). The fifth cranial or trigeminal nerve
lie lateral to the decussation of the superior takes origin from the caudoventrolateral aspect
cerebellar peduncles and the red nuclei. of the pons.
502 Chapter 8. T he B r a in
31 30
F ig . 8-13. Medial view of hemisected brain with the internal capsule and thalamus dissected.
1. Olfactory bulb 13. Inferior colliculus 25. Pyramis
2. Corona radiata of frontal lobe 14. Fissura prima 26. Uvula
3. Anterior crus of internal capsule 15. Arbor vitae cerebelli 27. Medulla oblongata
4. Cruciate sulcus 16. Posterolateral fissure 28. Mammillotegmental tract
5. Stria medullaris thalami 17. Nodulus 29. Pons
6. Corona radiata of parietal lobe 18. Lingula 30. Habenulointerpeduncular tract
7. Posterior crus of internal capsule 19. Lobulus centralis 31. Location of interpeduncular nucleus
8. Dorsal aspect of thalamus 20. Lobulus ascendens 32. Mammillary body
9. Habenular nucleus 21. Culmen 33. Mammillothalamic tract
10. Posterior commissure 22. Declive 34. Optic chiasm
11. Superior colliculus 23. Folium 35. Column of fornix
12. Corona radiata of occipital lobe 24. Tuber 36. Anterior commissure
III. Oculomotor nerve
In the basilar part o f the pons (Fig. 8-19) nerve, from the muscles of mastication and pos
transverse fibers (fibrae pontis transversae) sibly from the extrinsic muscles of the eye, reach
bridge the ventral surface of the pons. Dorsal to the m esencephalic nucleus of the trigeminal
the transverse fibers longitudinal bundles nerve by way of the m esencephalic tract.
(fasciculi longitudinales) are located on either Among the other tracts in the dorsal portion
side. Each longitudinal bundle forms the caudal of the pons there are the ascending fiber bundles
continuation of the crus cerebri and consists of which, after passing through the pons, either
corticospinal, corticonuclear, and corticopontine continue through the midbrain or end in mesen
fibers. The corticospinal fibers (fibrae cortico- cephalic nuclei, as well as the descending fiber
spinales) pass caudad through the medulla systems which originate from the mesencepha
oblongata to the gray matter of the spinal cord. lon.
The corticonuclear fibers (fibrae corticonu- The m edial longitudinal bundle brings vestib
cleares) convey impulses to the lower motor ular impulses to the motor nuclei of the nerves
neurons in the brain stem. The corticopontine supplying the extrinsic muscles of the eye. As
fibers (fibrae corticopontinae) end in the pontine in the mesencephalon, it is located close to the
nuclei (nuclei pontis), which consist of the gray midline and ventral to the derivative of the
matter occupying the remainder of the basilar lumen of early developmental stages, which is
part of the pons. The pontine nuclei give rise to represented in the pons by the fourth ventricle.
the transverse fibers of the pons which cross the See Whitaker and Alexander (1932).
midline and ascend as the middle cerebellar The trigeminal lemniscus originates from the
peduncle (brachium pontis, B.N.A., or pedun- secondary sensory neurons of the main and
culus cerebellaris medius, P.N.A.) into the cere spinal trigeminal nuclei of the opposite side.
bellum (Figs. 8-3, 8-17). After crossing the midline its fibers accompany
The dorsal portion o f the pom contains motor the medial lemniscus through the pons and the
and sensory nuclei of the trigeminal nerve, the mesencephalon to the thalamus. See Russel
caudal part of the mesencephalic tract, and the (1954).
beginning of the spinal tract of the trigeminal The m edial lemniscus, with the lateral spino
nerve (tractus spinalis nervi trigemini). The re thalamic tract applied to its lateral border,
maining area is occupied by a number of ascend occupies a ventromedial position in the dorsal
ing and descending fiber bundles and by the part of the pons medial to the lateral lemniscus.
reticular substance of the pons. See Valverde The lateral lemniscus (Fig. 8-15), the link be
(1961). tween myelencephalic and mesencephalic struc
The trigeminal nerve (nervus trigeminus) car tures of the acoustic system, lies medial to the
ries afferent fibers from the head region and middle cerebellar peduncle in the caudal portion
efferent fibers to the muscles of mastication. It of the pons. Rostrally, it emerges from this posi
originates from the pons caudal to the middle tion to become superficial before entering the
cerebellar peduncle just ventral to the cerebel nucleus of the inferior colliculus (Fig. 8-3).
lum (Figs. 8-4, 8-5). Its sensory fibers enter the Along its path the nuclei of the lateral lemniscus
main sensory nucleus (nucleus sensorius superior (nuclei lemnisci lateralis) are inserted. See Fuse
nervi trigemini) and the spinal nucleus (nucleus (1920a, 1926).
tractus spinalis nervi trigemini). The main sen The superior cerebellar peduncles (Fig. 8-17)
sory nucleus lies immediately medial to the run dorsolaterally along the fourth ventricle
entering trigeminal fibers. The spinal nucleus is throughout their course through the pons except
located caudal to the main nucleus and medial for the very rostral part, where they converge
to the .spinal tract o f the trigeminal. The spinal ventrad to the decussation of the superior cere
tract is formed by trigeminal fibers which will bellar peduncles.
synapse with secondary sensory neurons at dif The tectospinal and the tectobulbar tracts
ferent levels of the spinal nucleus. Both the spinal descend close to the midline between the medial
tract and the nucleus of the spinal tract extend longitudinal bundle and the medial lemniscus.
caudad through the medulla oblongata to the The rubrospinal tract is in a lateral position
beginning of the spinal cord (Fig. 8-21). The medial to the lateral lemniscus.
motor fibers of the trigeminal nerve originate The fourth ventricle (ventriculus quartus)
from the motor nucleus of the trigeminal nerve represents the ventricular system in the rhomb
(nucleus motorius nervi trigemini), which lies encephalon (Figs. 8-12, 8-17). In the pons it is
medial to the main sensory nucleus (Plate 9). bounded ventrally by the dorsal free border of
The proprioceptive impulses of the trigeminal the dorsal part of the pons, and laterally by the
50 4 Chapter 8. T he B r a in
superior cerebellar peduncles. The roof is formed cerebellum. The corpus cerebelli becomes con
by the anterior medullary velum and the cere siderably enlarged and forms the bulk of the
bellum. Rostrally, it connects with the third cerebellum.
ventricle by way of the cerebral aqueduct in the The fissura prima, the next cerebellar fissure
mesencephalon. Caudally, the fourth ventricle to appear, divides the corpus cerebelli into the
continues into the myelencephalon. anterior and the posterior lobes. In the adult
The cerebellum is derived from the dorsal cerebellum the fissura prima runs in a dorsoven-
portion of the metencephalon. It lies caudal to tral direction and becomes deep. In spite of its
the mesencephalon and the occipital pole of the considerable depth it is not easily detectable
cerebrum and dorsal to the fourth ventricle in among the large number of similar folia unless
the region of the pons and the rostral portion of the cerebellum is hemisected (Figs. 8-13, 8-18).
the medulla oblongata. It is a deeply fissured, This subdivision of the cerebellum has func
more or less globular part of the brain and is con tional significance. The flocculonodular lobe is
nected to the brain stem by three pairs of cere associated with the vestibular system. The an
bellar peduncles and portions of the roof of the terior lobe receives fibers from the spinal cord.
fourth ventricle. The posterior lobe has largely corticoponto-
The middle portion of the ventral surface of cerebellar connections with the exception of the
the cerebellum forms the part of the roof of the paraflocculus laterally and the pyramis and
fourth ventricle between the anterior medullary uvula caudoventrally, which obtain spinal fibers
velum (velum medullare anterius) rostrally and similar to the anterior lobe.
the posterior medullary velum (velum medullare From a topographical point of view the cere
posterius) caudally. Both vela are attached to bellum is divided into a median portion, the
the cerebellum. Also connected to the ventral vermis, and two lateral hemispheres (Fig. 8-18).
surface of the cerebellum are the three pairs of The vermis and the hemispheres are subdivided
cerebellar peduncles (Figs. 8-3, 8-17; Plates 4 into a number of lobules which in turn are fur
to 7, 9). In rostrocaudal sequence they are the ther subdivided by secondary foliation into sub-
superior cerebellar peduncle (brachium con- i lobules.
junctivum, B.N.A., or pedunculus cerebellaris The classical names of the various vermian
superior), the middle cerebellar peduncle lobules are, in rostrocaudal sequence, lingula
(brachium pontis, B.N.A., or pedunculus cere cerebelli; lobulus centralis; culmen; declive;
bellaris medius), and the inferior cerebellar folium vermis; tuber vermis; pyramis vermis;
peduncle (corpus restiforme, B.N.A., or pedun uvula vermis; and nodulus. Dexler (1932),
culus cerebellaris inferior). Ackerknecht (1943), and Sisson and Grossman
Like the cerebrum, the cerebellum has a pe (1953) describe the lobus or lobulus ascendens
ripheral cortex, white matter underneath the between the lobulus centralis and the culmen.
cortex, and centrally located nuclear masses. The fissura prima is located between the culmen
The cortex consists of the cerebellar fo lia (folia and the declive. The declive represents the ver
cerebelli), which correspond to the gyri of the mian portion of the lobulus simplex (Fig. 8-13).
cerebral cortex. The white matter is called the Earlier workers based the terminology of the
corpus medullare (Fig. 8-19). It connects the vermian lobules on the medullary rays. More
cerebellar cortex and the cerebellar nuclei with recently cortical subdivisions, which develop
each other and with the brain stem by way of the long before the medullary rays, have been used
cerebellar peduncles. The arrangement of gray by Larsell (1953, 1954) and designated in rostro
and white matter results in a treelike appearance caudal sequence with roman numerals.
of the cerebellum, especially in sagittal sections, Vermian lobules I to V belong to the anterior
and is referred to as the arbor vitae (Fig. 8-13). lobe, VI to IX to the posterior lobe, and vermian
In early developmental stages the postero lobule X forms the nodulus which is the vermian
lateral fissure (fissura posterolateralis), the first part of the flocculonodular lobe. The pyramis
cerebellar fissure to appear in the embryo, di and the uvula, the vermian portions of the pos
vides the cerebellum into the beginnings of the terior lobe which receive spinal connections,
flocculonodular lobe caudally and the corpus correspond to lobules VIII and IX respectively.
cerebelli or body of the cerebellum rostrally The hemispheres are largely subdivided into
(Fig. 8-13). In the further development the the paraflocculus, the paramedian lobule, the
flocculonodular lobe stays comparatively small ansiform lobule, and the lateral part of the
and is almost entirely hidden on the ventral sur lobulus simplex (Figs. 8-4, 8-5, 8-18).
face of the caudal portion of the body of the The paraflocculus is located at the rostro-
M eten ceph alo n 505
16 14
F ig . 8-15. Lateral view of the brain dissected to show projection pathways.
1. Olfactory bulbs 12.
2. Left cerebral hemisphere olive)
3. Internal capsule (lateral aspect) 13. Location of olivary nucleus (inferior olive)
4. Crus cerebri 14. Pyramid
5. Acoustic radiation 15. Trapezoid body
6. Medial geniculate body 16. Transverse fibers of pons
7. Superior colliculus 17. Longitudinal fibers of pons
8. Brachium of inferior colliculus 18. Transverse peduncular tract
9. Inferior colliculus 19.
10. Lateral lemniscus 20.
11. Cerebellum 21.
II. Optic nerve
III. Oculomotor nerve
F ig . 8-16. Medial view of the right cerebral hemisphere with the hippocampus and paraterminal gyms removed.
1. Olfactory bulb
2. Caudate nucleus
3. Genu of corpus callosum
4. Body of corpus callosum
5. Splenium of corpus callosum
6. Posterior horizontal ramus of splenial sulcus
7. Cut surface between cerebrum and brain stem
8. Amygdaloid body
9. Piriform area
10. Stria terminalis
11. Anterior commissure
506 Chapter 8. T he B r a in
lateral edge of the hemisphere. It is divided into of the pons which convey corticopontocerebellar
dorsal and ventral limbs which consist of a U- impulses.
shaped series of concentrically arranged short The inferior cerebellar peduncle has the larg
folia. The ventral paraflocculus projects laterally est number of components. It consists of spino
into the cerebellar fossa of the temporal bone. cerebellar fibers, olivocerebellar fibers, external
Both limbs extend around the ventrolateral arcuate fibers, vestibulocerebellar fibers, and
margin of the hemisphere dorsal to the flocculus. fastigiobulbar fibers. The spinocerebellar fibers
The dorsal paraflocculus reaches the paramedian come from the dorsal spinocerebellar tract. The
lobule on the caudal aspect of the cerebellum. olivocerebellar fibers form the olivocerebellar
The paramedian lobule lies on the caudal tract. The external arcuate fibers are subdivided
aspect on either side of the vermis and consists into dorsal external arcuate fibers, derived from
of transversely running folia. It connects with the accessory cuneate nucleus, and into ventral
the ansiform lobule to form the ansoparamedian external arcuate fibers, which originate from the
lobule, which represents the bulk of the cere arcuate and lateral reticular nuclei and dissemi
bellar hemispheres and is bounded rostromedi- nated cells of the reticular formation. The vestib
ally by the lateral part of the lobulus simplex. ulocerebellar fibers originate from certain vestib
The flocculus (Fig. 8-19) is the most ventral ular nuclei and are joined by direct fibers from
portion of the cerebellum and forms the lateral the vestibular part of the vestibulocochlear
part of the flocculonodular lobe. It lies dorsal to nerve. The fastigiobulbar fibers originate from
the superficial origin of the trigeminal nerve and the fastigial nucleus and form the fastigiobulbar
rostral to the cochlear nuclei. tract (tractus fastigiobulbaris, I.N.A.). After par
The cerebellar nuclei (Plates 6 , 9) consist of tial crossing the fastigiobulbar tract occupies the
four paired nuclear masses: the dentate, the medial part of the inferior cerebellar peduncle
emboliform, the globose, and the fastigial nuclei and terminates in the lateral and inferior vestib
in lateromedial sequence. They are located along ular nuclei and in the reticular formation.
a transversely running line in the center of the For additional information about the cere
corpus cerebelli just dorsal to the roof of the bellum see Berkeley (1894), Mingazzini (1895),
fourth ventricle. They receive fibers from the Marburg (1904), Rademaker (1926), Nasedkin
cerebellar cortex and give rise to the efferent (1929), Miskolczy (1931), Bertrand, Medynski,
fibers of the cerebellum as well as to fibers which and Salles (1936), Dow (1940), Stella, Zatti, and
return impulses to the cerebellar cortex. Sperti (1955), Stam (1958-59), and Perkins
The dentate nucleus (nucleus dentatus) is the (1961).
largest of the group. It lies slightly more caudal
than the other nuclei. The emboliform and MYELENCEPHALON
globose nuclei (nucleus emboliformis et nucle
us globosus) are two smaller, less distinct nuclear The medulla oblongata (Plates 5 to 9) is the
masses and are jointly referred to as nucleus caudalmost portion of the brain stem and ex
interpositus. The fastigial nucleus (nucleus tends from the pons to the beginning of the
fastigii) is of an intermediate size. It appears spinal cord. It contains the remainder of the
roundish in cross section and lies adjacent to the cranial nerve nuclei, ascending and descending
fastigial nucleus of the other side. fiber bundles, the reticular formation of the
The efferent and afferent cerebellar fibers pass medulla oblongata, and the caudal part of the
through the cerebellar peduncles (Figs. 8-3, fourth ventricle. The term “bulb” is a synonym
8-17). The superior cerebellar peduncle is the for medulla oblongata or myelencephalon.
most medial of the three cerebellar peduncles. On the ventral aspect of the medulla oblon
It carries largely efferent fibers from the dentate, gata the two pyramids (Fig. 8-5) emerge from
emboliform, and globose nuclei to the opposite the pons. Separated from one another by the
red nucleus and thalamus as cerebellorubral and ventral m edian fissure (fissure mediana ven
cerebellothalamic tracts (tractus cerebelloru- tralis), they extend to the caudal part of the
bralis et tractus cerebellothalamicus). The medulla oblongata. They form the caudal con
ventral spinocerebellar tract reaches the superior tinuation of the longitudinal fibers of the pons
cerebellar peduncle and enters the cerebellum and carry corticospinal impulses to lower motor
rostrally. neurons in the spinal cord. The corticonuclear
The middle cerebellar peduncle lies more fibers do not continue into the pyramids but
laterally than the other two peduncles. It con enter the motor nuclei of both sides in the brain
sists of the continuation of the transverse fibers stem through the reticular formation. The corti
508 Chapter 8. T he B r a in
cospinal fibers of each pyramid continue into subdivisions of the facial nucleus see Yagita
the spinal cord as two unequal portions with dif (1910) and Papez (1927). The motor fibers course
ferent topographical locations. The larger parts dorsorostrad to the nucleus of the abducens
of both sides cross the midline by sweeping nerve as they form the internal genu of the facial
dorsolaterad and slightly caudad toward the nerve (genu nervi facialis). From the internal
lateral funiculus of the opposite side of the spinal genu the facial nerve extends obliquely ventro-
cord, where they form the lateral corticospinal laterad through the reticular formation and along
tract (tractus corticospinalis lateralis). The cross the lateral side of the superior olive to its super
ing of the fibers is referred to as the decussation ficial origin from among the fibers of the trape
o f the pyramids (decussatio pyramidum). The zoid body caudal to the medial edge of the super
smaller portions of the corticospinal fibers of the ficial origin of the trigeminal nerve (Fig. 8-5).
two sides continue caudad in the ventral funiculi The parasympathetic fibers come from the
of their respective sides as the ventral cortico superior salivatory nucleus (nucleus salivatorius
spinal tracts (tractus corticospinales ventrales). superior), which is an ill-defined nuclear mass
These fibers cross the midline farther caudally at within the reticular formation. See Yagita and
the various segmental levels of the spinal cord. A Hayama (1909). The taste fibers contribute to
small number, however, may remain on the same the tractus solitarius, a distinct fiber bundle,
side. which is essentially associated with the ninth
The sixth cranial or abducens nerve (nervus and the tenth cranial nerves in the caudal
abducens) supplies the lateral rectus and the medulla oblongata. The question of the proprio
retractor bulbi muscles with motor fibers. The ceptive innervation of the mimetic musculature
nucleus of the abducens nerve (nucleus nervi is still open (Brodal 1959).
abducentis) lies close to the dorsal midline of the The eighth cranial or vestibulocochlear nerve
medulla oblongata and the medial longitudinal (nervus vestibulocochlearis) consists of a vestib
fasciculus, from which it receives vestibular ular portion (pars vestibularis) and a cochlear
impulses. The fibers of the abducens run ven part (pars cochlearis) which enter the central
trally and slightly laterad through the rostral part nervous system together at the ventrolateral
of the reticular formation of the medulla ob border of the medulla oblongata directly caudal
longata and along the medial border of the to the superficial origin of the trigeminal nerve
superior olive. They emerge directly lateral to (Figs. 8-4, 8-5).
the pyramids among the fibers of the trapezoid The cochlear part conveys acoustic impulses
body, a group of transverse fibers belonging to from the inner ear to the dorsal and ventral
the acoustic system and located immediately cochlear nuclei. The dorsal cochlear nucleus
caudal to the pons (Fig. 8-5). (nucleus cochlearis dorsalis) lies in the dorso
The superior olive (nucleus olivaris superior, lateral portion of the medulla oblongata lateral
B.N.A., or nucleus dorsalis corporis trapezoidei, to the fibers of the inferior cerebellar peduncle.
P.N.A.) is another component of the acoustic The ventral cochlear nucleus lies at the lateral
system (Plate 5). It forms a rounded nuclear mass edge of the trapezoid body where the vestibulo
halfway between the midline and the lateral cochlear nerve enters the medulla oblongata.
margin of the rostral medulla oblongata just The cochlear nuclei give rise to the secondary
dorsal to the trapezoid body (Fig. 8-15). The acoustic fibers which enter the reticular forma
connections of both the trapezoid body and the tion and the superior olives and also contribute
superior olive will be described with the eighth to the lateral lemniscus.
cranial or vestibulocochlear nerve. The dorsal cochlear nucleus sends fibers
The seventh cranial or fa c ia l nerve (nervus through the reticular formation across the mid
facialis) carries motor fibers to muscles of the line to the opposite superior olive and lateral
face, parasympathetic fibers to the sublingual lemniscus. Some fibers of the dorsal cochlear
and mandibular salivary glands, taste fibers from nucleus terminate in the reticular formation of
the rostral two-thirds of the tongue by way of both sides.
the chorda tympani, and presumably proprio The ventral cochlear nucleus gives origin to
ceptive fibers from the muscles of facial expres the trapezoid body, which crosses the midline
sion. The motor nucleus of the facial nerve dorsal to the pyramids (Fig. 8-5). The trapezoid
(nucleus nervi facialis) lies caudal to the superior body carries fibers to the opposite lateral lemnis
olive and is marked on the ventral surface of the cus and the superior olives of both sides.
medulla oblongata by a slight elevation caudal to The superior olive functions as an acoustic
the trapezoid body (Plate 6 ). For details on the relay center by sending a considerable number
M yelen ceph a lon
F ig. 8-19. Lateral view of the brain with the left cerebral hemisphere, left transverse fibers of the
pons and left middle cerebellar peduncle removed.
1. Cut surface between left cerebral hemisphere 15. Longitudinal fibers of pons
and brain stem 16. Transverse fibers of pons
2. Medial surface of right cerebral hemisphere 17. Crus cerebri
3. Lateral geniculate body 18. Optic tract
4. Medial geniculate body II. Optic nerve
5. Superior colliculus III. Oculomotor nerve
6. Inferior colliculus IV. Trochlear nerve
7. Brachium of inferior colliculus V. Trigeminal nerve
8. Lateral lemniscus VI. Abducens nerve
9. Corpus medullare of cerebellum VII. Facial nerve
10. Vermis of cerebellum V III. Vestibulocochlear nerve
11. Flocculus IX . Glossopharyngeal nerve
12. External arcuate fibers X. Vagus nerve
13. Spinal tract of trigeminal nerve XI. Accessory nerve
14. Dorsal spinocerebellar tract X II. Hypoglossal nerve
510 Chapter 8. T he B r a in
F ig . 8-20. Lateral view of the brain with the left cerebral hemisphere removed and the inferior
cerebellar peduncle dissected.
1. Lateral lemniscus 9. Inferior cerebellar peduncle
2. Crus cerebri 10. Fasciculus gracilis
3. Lateral geniculate body 11. Fasciculus cuneatus
4. Medial geniculate body 12. Spinal tract of trigeminal nerve
5. Superior colliculus 13. External arcuate fibers
6. Inferior colliculus 14. Trapezoid body (transected)
7. Cerebellum 15. Location of main sensory nucleus of trigeminal nerve
8. Superior cerebellar peduncle V. Trigeminal nerve
F ig . 8-21. Lateral view of the brain with the left cerebral hemisphere removed and the spinal tract
of trigeminal nerve dissected.
1. Fasciculus cuneatus
2. Spinal tract of trigeminal nerve
3. Location of vestibular nuclei
4. Ventral spinocerebellar tract
5. Transverse fibers of pons (transected on midline)
6. Superior cerebellar peduncle
V. Trigeminal nerve
M yelen ceph a lon 511
of its fibers to the lateral lemniscus. These fibers origin and termination in the reticular formation
may carry acoustic impulses from either side, and emerge in linear fashion just caudal to the
since each superior olive receives secondary superficial origin of the vestibulocochlear nerve
acoustic fibers from both ventral cochlear nuclei. along the lateral border of the medulla oblongata
Most fibers of the lateral lemniscus terminate in (Figs. 8-4, 8-5).
the caudal colliculus. Some synapse in the nuclei The external arcuate fibers (fibrae arcuatae
of the lateral lemniscus, and some run directly to extemae) curve superficially around the lateral
the medial geniculate body. and dorsolateral aspect of the rostral medulla
In addition to relaying acoustic impulses to oblongata (Fig. 8-19). They may be separated
higher centers, the superior olive serves also as into dorsal and ventral external arcuate fibers.
an acoustic reflex center. Some of its fibers be The dorsal external arcuate fibers originate from
long to various acoustic reflex arcs which influ the accessory cuneate nucleus, which will be
ence cells in motor nuclei of cranial nerves either described with the dorsal aspect of the medulla
directly or by way of intercalated neurons in the oblongata. The ventral external arcuate fibers
reticular formation. originate largely from the arcuate and lateral
The fibers of the vestibular part of the vestib reticular nuclei and disseminated cells of the
ulocochlear nerve bypass the inferior cere reticular formation. The arcuate nuclei (nuclei
bellar peduncle ventrally and synapse in the arcuati) are located along the pyramids in the
vestibular nuclei which lie in the dorsolateral rostral part of the medulla oblongata and repre
portion of the rostral medulla oblongata and sent caudally displaced pontine nuclei. The
caudal pons in the floor of the fourth ventricle. lateral reticular nucleus will be described with
Some fibers of the vestibular part of the eighth the reticular formation. The ventral external
nerve enter the cerebellum directly, without arcuate fibers cover the rostral part of the spinal
synapsing in the vestibular nuclei, by way of the tract of the trigeminal nerve and contribute to
medial part of the inferior cerebellar peduncle. the inferior cerebellar peduncle together with
The vestibular nuclei form the sensory nuclei the dorsal external arcuate fibers (Fig. 8-20).
of the vestibular part of the vestibulocochlear The spinal tract and nucleus o f the trigeminal
nerve and consist of the superior, inferior, lateral, nerve were mentioned under the description of
and medial vestibular nuclei (nucleus vestibu the pons as extending from the superficial origin
laris superior, inferior, lateralis, et medialis). De of the trigeminal nerve to the beginning of the
tailed information about the secondary vestibu spinal cord, thus forming landmarks through the
lar connections is controversial. In a general entire length of the medulla oblongata. The
way, however, the vestibular nuclei convey im fibers which form the rostral portion of the spinal
pulses to the cerebellum, the spinal cord, the tract of the trigeminal nerve descend along the
motor nuclei of the nerves innervating the ex lateral margin of the medulla oblongata medial
trinsic muscles of the eye, and the cerebral cor to the ventral cochlear nucleus and the ventral
tex. See Whitaker and Alexander (1932). external arcuate fibers (Figs. 8-19, 8-20, 8-21).
The fibers to the cerebellum run in the medial The caudal part of the spinal tract of the tri
portion of the inferior cerebellar peduncle along geminal nerve becomes exposed at the dorso
with the primary vestibular fibers which enter lateral margin of the caudal portion of the me
the cerebellum without synapsing in the vestib dulla oblongata (Fig. 8-3).
ular nuclei. The secondary vestibular fibers to The nucleus of the spinal tract of the trigemi
the spinal cord form either the lateral vestibulo nal nerve accompanies the spinal tract medially
spinal tract (tractus vestibulospinalis lateralis), through the medulla oblongata. It contributes
or they enter the caudal continuation of the secondary sensory fibers to the trigeminal lem
medial longitudinal fasciculus, which becomes niscus and connects with brain stem motor
the ventral vestibulospinal tract (tractus vestib nuclei by way of the reticular formation.
ulospinalis ventralis). The secondary fibers to The fibers of the vagus group can be traced
the cranial nerves which innervate the extrinsic from their superficial origin among the ventral
muscles of the eye ascend in the rostral portion external arcuate fibers through the spinal tract
of the medial longitudinal fasciculus. and nucleus of the trigeminal nerve and the
The ninth cranial or glossopharyngeal nerve, reticular formation to their respective nuclei of
the tenth cranial or vagus nerve, and the origin and termination. See Okinaka and
eleventh cranial or accessory nerve (nervus glos- Kuroiwa (1952).
sopharyngeus, nervus vagus, nervus accessorius) The glossopharyngeal nerve carries sensory
are closely related and may be referred to as the and taste fibers from the caudal third of the
vagus group. They share some of their nuclei of tongue, parasympathetic fibers to the parotid
51 2 Chapter 8. T he B r a in
salivary gland, and motor fibers to the stylo tral to the nucleus gracilis, which forms a land
pharyngeus muscle. It carries sensory fibers from mark on the caudal medulla oblongata along the
the carotid sinus (Adams 1958), the pharynx, and dorsal midline together with the fasciculus
possibly from the ear. gracilis (Fig. 8-17).
The vagus nerve has its superficial origin The motor fibers of the vagus group which in
caudal to the superficial origin of the glosso nervate the striated musculature of the pharynx
pharyngeal nerve. It consists of parasympathetic and larynx arise from the nucleus ambiguus in
nerve fibers for the viscera in the neck, thorax, the center of the reticular formation. They run
and abdomen; motor fibers for the striated mus dorsad, then turn ventrolaterad and join the root
culature of the pharynx and larynx; sensory bundles of the glossopharyngeal, vagus, and
fibers from the pharynx, larynx, trachea, esopha accessory nerves. All three nerves receive motor
gus, and thoracic and abdominal viscera; taste fibers from the myelencephalon, but those of the
fibers from the epiglottis; and sensory fibers from accessory nerve join the vagus nerve by way of
the skin of the external ear. the internal ramus of the accessory nerve.
The accessory nerve carries only motor fibers The cutaneous fibers from the ear, which are
to striated musculature (Brodal 1959). It has carried by the glossopharyngeal and the vagus
spinal and cranial roots (radices spinales et nerves, synapse in the spinal nucleus of the tri
craniales). The spinal roots originate in the geminal nerve, which provides for the secondary
cervical spinal cord and are merely topographi connections to motor nuclei and higher levels.
cally related to the cranial roots. The cranial The twelfth cranial or hypoglossal nerve
roots leave the medulla oblongata caudal to the (nervus hypoglossus) supplies motor fibers to the
superficial origin of the vagus nerve. They join musculature of the tongue. Its cell bodies are
the fibers of the spinal roots for a short distance, located in the hypoglossal nucleus (nucleus
but enter the vagus nerve by means of the so- nervi hypoglossi), which lies along the midline
called internal ramus of the accessory nerve. In ventromedial to the tractus solitarius, the nucleus
this fashion they convey motor impulses to the of the tractus solitarius, and the dorsal nucleus
musculature of the pharynx and larynx by way of the vagus nerve. Its rostral portion protrudes
of the branches of the vagus nerve. into the fourth ventricle like the dorsal nucleus
The sensory and taste fibers of the glosso of the vagus nerve. Its caudal part is situated
pharyngeal and vagus nerves form the tractus directly lateral to the central canal. The fibers of
solitarius together with the taste fibers of the the hypoglossal nerve pass ventrally and slightly
facial nerve, which were already referred to. laterad through the reticular formation and
The tractus solitarius is a landmark of the caudo- emerge from the ventral surface of the caudal
dorsal part of the reticular formation of the medulla oblongata after traversing the inferior
medulla oblongata medial to the spinal nucleus olive (Figs. 8-4, 8-5).
of the trigeminal nerve. Its fibers enter the The inferior olive (nucleus olivaris inferior,
nucleus o f the tractus solitarius (nucleus tractus B.N.A., or nucleus olivaris, P.N.A.) is a nuclear
solitarii), which in turn conveys impulses to the mass in the caudoventral part of the medulla
motor nuclei of the cranial nerves by way of the oblongata directly dorsolateral to the pyramids
reticular formation. It also gives origin to the (Fig. 8-15; Plate 7). It receives fibers from a
secondary taste fibers, but the secondary path number of nuclear masses such as the globus
for taste impulses is still an open question. pallidus and the red nucleus which are involved
The parasympathetic fibers of the glosso in phylogenetically older motor systems. The
pharyngeal nerve have their cell bodies in the efferent fibers of the inferior olive form the
inferior salivatory nucleus (nucleus salivatorius olivocerebellar tract (tractus olivocerebellaris),
inferior), which lies caudal to the superior saliva which consists largely of crossed fibers. On its
tory nucleus of the facial nerve in the reticular way to the inferior cerebellar peduncle it tra
formation of the medulla oblongata. See Yagita verses the spinal tract of the trigeminal nerve,
(1909). which as a result becomes separated into several
The parasympathetic fibers of the vagus nerve bundles.
originate from the dorsal nucleus o f the vagus The dorsal spinocerebellar tract (tractus spino-
nerve (nucleus dorsalis nervi vagi). The dorsal cerebellaris dorsalis) is another source of fibers
nucleus of the vagus nerve lies medial to the for the inferior cerebellar peduncle. It carries
tractus solitarius and nucleus of the tractus proprioceptive impulses largely from the same
solitarius. Rostrally, it protrudes into the fourth side of the spinal cord to the cerebellum. In the
ventricle. Caudally, it becomes submerged ven (Text continued on page 523.)
a e nia Iis thal
Plexus chorio
Nuc rhomboideus thal
Nuc. paracentral is thal
Larr
2
Nuc anterovent thal.
PI
Ped. thal dorsolat r
Nuc anteromed thal pi
z
Nuc. centralis med thal n
pi
Nuc /ned dors thal
Adhaesio i reticularis thal s
>
r
Nuc veni ant thal c
Nuc reuniens thal z
Lamina meduiiaris ext. tjal.
Nuc ret culans thal
Genu capsulae int
Putamen
Capsulo externa ■ » .....
Fasc. mamillothalamic Nuc. interstitial is ped. *hal inf
S. suprasplenialis
nduseum griseum
Ventriculus lat
Fiss. chorioidea
Stria terminalis
Radiatio optica -
lat -
Nuc. habenularis ,
I med.__f
Ventriculus tert us
Nuc. cqrp. geniculati lat. dors.
Stria terminalis
G. ectosylvius post.
\
Capsula extrema —
Lamina medullaris ext thal
Plexus chorioid vent lat.-
Fiss chorioidea
Ventriculus la
optic
th c lj / ' j .
p o s te ^jm e ^y ia l. / /
Cru^neTt»R£L ' /
Nuc centrum me4»8fium th
Tr subthalamicotegmentalis
Nuc. subthalamicus
Tr cerebello-rubro-thol.
G. suprosplenialis
G. lateralis
S. entolaterolis
S. lateralis
514
G. suprasylvius med.
S. ectosylvius med.
Chapter
/
Nuc caudatus, cauda ,G sylvius
8.
jc tr. habenulointerpedunc, lat.
uc. suprageniculatus thol.
The
uc. post thai
B r a in
uc parafascic. thal.
^ X '^ Z o n a incejfta
Area tegmentans H
Hippocampus •
~?nyoquiomotoni \
ad opt
Ventriculus lot ■ Nuc. n. oculomotor!* ^ \
j Tr spinotecfon%P \
profundus mesen. \ *
dentatus uc tr. pnesen. n. ttigemini splnothilamicuX
Corpus jculatum r
^ / ■ \ \
-A k /- \
_____ (Icmniscus mod
Tr cercbrospinalis riformis
^ ' Subst. nigra camp
Tr ,cerebrobulbaris
„ Tr. frontopontinus N. oculomotorius
tC?*-'’ Dec. dors, tegmenti Fibrae n. oculomotorii
Area vent, tegmenti Tr. rubrospinalis
Tr. habenulointerpedunc’
Fibrae tr. rubrospinalis
Dec. vent, tegmenti
Plate 3. Transverse section of the brain. (Singer: The Brain of the Dog in Section.)
G. Iingual is Cuimen G. suprasylvius post
\ /
Fasc.
um meduMare ant
s. Tr mesen. n. trigemini
Nuc. parapeduncularis
Lemniscus lat.
Locus coeruleus
Nuc. dors, raphae
Nuc. laterodors tegmenti
Tr. tegmentalis cent. Nuc. dors tegmenti
Lemniscus lat Ped. cerebellaris med.
Tr. rubrospinalis Radix mot. n. trigemini
518
pnma
Nuc fastigii
-ae vestibulo-cerebellares
Ventriculus quartus
Chapter
Nuc vestibuloris lat.
L. onsiformis —
8.
Ped cerebellaris inf.
Nuc. vestibularis descend
The
et
Radix descend, n vestib
Flocculus
B r a in
Paraflocculus
Nuc vestibularis med
Nuc tr solitaru
^?-«^;Tuberculum acusticum
Tr. tegmentalis cent.
Tr. sol ifarius
Nuc. reticularis parvicell.
Nuc. cochlearis dors
Tr. rubrospinalis Nuc. cochlearis vent.
Radix sens, n, glossophar.
pars. lat.
cochlearis dors.
pars intermed
Nuc. n. facialis, pars ventrolat. N. glossopharyngeus
spinalis n. trigemini
pars med
pars ventromed Nuc. tr spin n trigemini, pars oralis
Fibrae vestibulares sec
Tr vestibulospinalis Fibrae n. facialis, ascend
Nuc. reticularis giganto Genu n facialis
Pyramis Tr cerebrospinalis
Plate 6. Transverse section of the hrain. (Singer: The Brain of the Dog in Section.)
Plate 7. Transverse section of the brain.
(Singer: The Brain of the D or in Section.)
519
Nuc. olivaris access.dors
Tc olivocerebellaris
Hilus nuc olivaris
Lemniscus med Nuc. olivaris
Nuc olivaris access, med
Pyramis
S. medianus Tr cerebraspinalis
520
Pyramis (vermis]
Chapter
Nuc. cuneatus
F asc gracilis
Fasc cuneatus
8.
Nuc. gracilis
T
Nuc tr spin. n. trigemini, pars caud.
he
Tr. cerebrospinalis lat.
Tr. spinalis n trigemini
B hain
Tr. rubrospinalis
Nuc. n, accessorii
Nuc. reticularis venf
Tr. spinothalamicus
Tr tectospmalis Tr spinocerebel laris vent.
S lateralis
L.ansiformis
M
Nuc“alfactorfus anr. pars lat.
yelen ceph a lo n
a j S i rum
Nuc interpositus
^externa
erebellaris vent
Nuc. olfactorius ant., pars vent Plj,arfien
Ped flocculi
Tuberculum olfactorium L. paramedianus
Globus pallidus 'ellaris sup.
Lamina medullaris inf pall
Tr. mesen n. trigemini
Nuc entopeduncularis
Tr. opticus' Radix n. intermedii
**** ?c==r=f e s n2adix sens, n. glossophar et n. vagi
Nuc amygdalae-|^0er^, ^ T r spinalis n. trigemini
Nuc. n accessorii
Nuc. subthalamicus
— —Tr spinothalamicus
Subst nigra-| rgt r. rubrospinalis
Tr. solitariu’- »—
Tr cerebrospinalis et bulbaris Radix n facialis, descend
Subst nigra lat
Nuc. olivaris sup.,pars lat.
Lemniscus / Nuc. preolivaris, pars med. Nuc. ambiguus^
Nuc vent caud. lemnisci lat Corpus trapezoideum Nuc reticularis lat
' Tr rubrospinalis
Tr. spinothalamicus' Oi
Nuc. mot. n. trigemini to
Plate 9. Sagittal section of the brain. (Singer: The Brain of the Dog in Section.)
F.b-ae tr tectaspinalis
A'vejs
Nuc dors raphae
Corpus gemculatum med
522
G dentatus Fibrae lemnisci lat.
Hippocampus Nuc. collicuii in*
Fimbna
locculus
Fiss rhmans post
S sylvius Veium medullare ant.
Ped, thai dorsolcf L -ansiforms
Lobus pyr-formis
Lingula
Fiss rhinaliS ant
Str a medullaris thal,
S presylvius Ventriculus quartus
Bulbus olfactorius
Chapter 8
[vermisj
The
Nuc, submed,!
B i.a in
Nut centrum medianum thal.
Putamen
Nuc vent lat t grisea cent
Globus paMi
Plate 10. Horizontal section of the brain. (Singer: T he Brain o f the Dog in S ection.)
M yelen ceph alon 523
caudal part of the medulla oblongata, before mid along the entire medulla oblongata rostral
joining the external arcuate fibers, the dorsal to the decussation of the medial lemniscus.
spinocerebellar tract runs along the ventrolateral Caudal to the inferior olive it receives the fibers
margin of the spinal tract of the trigeminal nerve of the ventral spinothalamic tract, and rostral to
(Fig. 8-19). The ventral spinocerebellar tract the inferior olive it is joined by the lateral spino
(tractus spinocerebellaris ventralis) carries homo thalamic tract, after which it carries all the
lateral as well as contralateral proprioceptive secondary sensory fibers from the opposite side
impulses. It ascends ventral to the dorsal spino of the body. The rostral course of the medial
cerebellar tract and the spinal tract of the tri lemniscus through the pons to the ventral pos
geminal nerve to the level of the pons, curves terolateral nucleus of the thalamus has been
around the superficial origin of the trigeminal described.
nerve, and enters the cerebellum by way of the The trigeminal lemniscus (lemniscus trigemi-
superior cerebellar peduncle (Fig. 8-21). nalis) originates from the spinal nucleus of the
On the dorsal aspect (Fig. 8-17) the rostral trigeminal nerve and the main sensory nucleus
part of the medulla oblongata forms the floor of in the pons. It carries the secondary fibers for all
the caudal portion of the fourth ventricle. This cutaneous sensations of the head region to the
floor is separated into two halves by the dorsal ventral posteromedial nucleus of the thalamus
median sulcus (sulcus medianus dorsalis). The and is topographically related to the medial
posterior medullary velum (velum medullare lemniscus.
posterius) forms the roof over this part of the Among the descending fiber tracts the corti
fourth ventricle. The fourth ventricle commu cospinal tracts form the pyramids, which were
nicates with the subarachnoid space by lateral described as landmarks of the ventral aspect of
apertures (aperturae laterales ventriculi quarti) the medulla oblongata.
or foramina of Luschka and the m edian aper The rubrospinal tract is located laterally in
ture (apertura mediana ventriculi quarti) or the reticular formation ventral to the spinal nu
foramen of Magendie. Caudally, the fourth ven cleus of the trigeminal nerve. The tectospinal
tricle continues as the central canal. For details tract runs along the midline dorsal to the medial
about the ventricular system see Fitzgerald lemniscus. Some vestibulospinal fibers, as men
(1961). tioned in the description of the vestibular nuclei,
In the caudal portion of the medulla oblongata form the lateral vestibulospinal tract; others
the fasciculus gracilis and the fasciculus cune- enter the caudal continuation of the medial
atus carry fibers for conscious proprioception longitudinal fasciculus. The lateral vestibulo
and, at least in the human being, tactile discrim spinal tract lies medial to the rubrospinal tract.
ination from the spinal cord to the nucleus grac The medial longitudinal fasciculus has a close
ilis and the nucleus cuneatus. The fasciculus spatial relation to the floor of the fourth ventricle
gracilis lies along the dorsal median sulcus and and the ventral side of the central canal. The
originates from lumbar and sacral spinal cord inferior olive gives rise to the olivospinal tract,
segments. The fasciculus cuneatus is located be which descends in the lateral funiculus of the
tween the fasciculus gracilis and the spinal tract spinal cord.
of the trigeminal nerve. It carries impulses from The inferior cerebellar peduncle (Fig. 8-20)
the thoracic and cervical part of the spinal cord has been referred to repeatedly. It connects the
(Fig. 8-17). medulla oblongata with the cerebellum. A sum
The fibers of the fasciculus gracilis terminate mary of its constituent fibers is given with the
in the nucleus gracilis, those of the fasciculus description of the cerebellum.
cuneatus in the nucleus cuneatus and accessory The remainder of the medulla oblongata is
cuneate nucleus (nucleus cuneatus accessorius), occupied by the reticular formation. It contains
which lies dorsolateral to the nucleus cuneatus a number of more or less well-defined reticular
(Fig. 8-3). The cell bodies of the accessory nuclei, including the lateral reticular nucleus
cuneate nucleus give rise to the dorsal external (nucleus reticularis lateralis), which was referred
arcuate fibers, which convey proprioceptive im to in connection with the ventral external arcu
pulses to the cerebellum by way of the inferior ate fibers.
cerebellar peduncle. The nucleus gracilis and The lateral reticular nucleus is a fairly well
the nucleus cuneatus give rise to internal arcuate circumscribed nuclear mass located lateral and
fibers which curve concentrically in a ventro caudal to the inferior olive. It receives fibers from
medial direction to the decussation of the medial the lateral funiculus of the spinal cord as well as
lemniscus (decussatio lemniscorum). descending fibers of yet to be established origin
The medial lemniscus lies dorsal to the pyra and contributes to the ventral external arcuate
524 Chapter 8. T he B r a in
fibers. The connections of the reticular nuclei ances of conditioned reflexes after ligation of arteries.
are the object of extensive studies. Arch. Neurol. Psychiat. (Chic.) 34: 699-713.
Arey, L. B., S. R. Bruesch, and S. Castanares. 1942. The rela
A number of cells in the reticular formation tion between eyeball size and number of optic nerve fi
aid in visceral functions such as cardiovascular bers in the dog. J. comp. Neurol. 76: 417-422.
and respiratory control. These physiological Arey, L. B., and M. Gore. 1942. The numerical relation be
centers are difficult to define morphologically. tween the ganglion cells of the retina and the fibers in the
optic nerve of the dog. J. comp. Neurol. 77: 609-617.
For comprehensive coverage of the dog’s
Ariens Kappers, C. U., G. C. Huber, and E. C. Crosby. 1936.
medulla oblongata see Hoffmann (1955) and The Comparative Anatomy of the Nervous System of
Bossy (1955). Compare also the publications by Vertebrates, Including Man. New York, Macmillan Co.
Biirgi and Bucher (1960) and Taber (1961). De Asratian, E. 1935. Motor defensive conditioned reflexes in dogs
tails on the pyramids and the pyramidal system with extirpated cortical motor areas of the cerebral hemi
spheres. C.R. Acad. Sci. (U.S.S.R.) 1: 159-164. (Quoted
may be found by consulting Starlinger (1895), from Biol Abstr. 1937/JJ; 3517.)
Lassek, Dowd, and Weil (1930), Szentagothai- Bahrs, A. M. 1927. Notes on reflexes of puppies in the first six
Schimmert (1941), Morin, Donnet, and Zwim weeks after birth. Amer. J. Physiol. 82: 51-55.
(1949), Morin, Poursines, and Donnet (1949), Bailey, P., and W. Haynes. 1940. Location of inhibitory respir
atory center in cerebral cortex of the dog. Proc. Soc. exp.
Morin, Poursines, and Maffre (1951), and Barone Biol. (N.Y.) 45: 686-687.
(1960). Investigations related to the cranial Balado, M. 1925. Anatomia externa del encephalo del perro.
nerves, in addition to those already cited, were Bol. Inst. Clin. quir. (B. Aires) 1: 128-168.
made by Parhon and Nadejde (1906), Kosaka Barnhard, J. W. 1940. The hypoglossal complex of vertebrates.
J. comp. Neurol. 72: 489-524.
(1909), Preziuso (1930), Sekita (1931), Barnhard
Barone, R. 1960. La substance blanche et ses courants de fibres
(1940), and Bossy (1955). Additional informa dans la moelle epiniere des mammiferes. Rev. Med. vet.
tion on structures in the medulla oblongata may I l l : 200-248.
be found in papers by Fuse (1920a), Phalen and Bartley, S. H., and E. B. Newman. 1931. Studies on the dog’s
Davenport (1937), Yoda (1941), and Lindgren cortex; I. The sensorimotor areas. Amer. J. Physiol. 99;
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and Borje (1953). Bary, A. 1898. Uber die Entwickelung der Rindencentren.
Arch. Anat. Physiol. Physiol. Abt. 1898: 341-360.
Basir, M. A. 1932. The vascular supply of the pituitary body in
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CHAPTER 9
TH E SPINAL CORD A N D M E N IN G E S
By ROBERT C. McCLURE
SPINAL CORD caudal to the last pair of spinal nerves, and others
the spinal cord caudal to the lumbar part. The
The spinal cord (medulla spinalis) is that part conus medullaris terminates as the filum termi-
of the central nervous system which lies in the nale.
vertebral canal from the level of the foramen
magnum to the junction of the sixth and seventh External Landmarks (Fig. 9-4)
lumbar vertebrae. The cross-sectional size and
shape of the spinal cord vary at different levels. The spinal cord, throughout its length, pre
sents a shallow longitudinal dorsal m edian sulcus
Spinal Cord Segments (Figs. 9 -1 , 9-2) (sulcus medianus dorsalis), which divides the
cord into two halves dorsally. A dorsal median
' The spinal cord may be divided into segments, septum of neuroglia extends ventrally from the
each segment being that portion of the spinal sulcus almost to the central canal. Lateral to
cord where fibers in the rootlets of a pair of the dorsal median sulcus, the dorsal rootlets of
spinal nerves enter and leave the cord. A spinal the spinal nerves enter the spinal cord along the
cord segment is identified by the same name and shallow dorsolateral sulcus (sulcus lateralis
number as the pair of spinal nerves which attach dorsalis). In the cervical part and cranial half of
to it (thus the second thoracic spinal cord seg the thoracic part of the spinal cord, the dorso-
ment [T2] is that part of the cord to which the intermediate sulcus (sulcus intermedius dorsalis)
rootlets of the second thoracic spinal nerves are is present between the dorsomedian and dorso
attached). The dog has thirty-six spinal cord seg lateral sulci. It is adjacent to the dorsolateral
ments: eight cervical, thirteen thoracic, seven sulcus at its origin in the mid-thoracic region.
lumbar, three sacral, and five coccygeal. The distance between the two increases as the
The spinal cord segments at the levels of the fasciculus cuneatus increases in size (ascending
brachial and lumbosacral plexuses are enlarged cranially in the spinal cord).
in diameter. The cervical enlargement (intumes- Ventrally, the spinal cord presents the longi
centia cervicalis) for the nerves of the brachial tudinal ventral median fissure (fissura mediana
plexus includes segments C5 to T2. The lumbar ventralis), which appears to divide the spinal
enlargement (intumescentia lumbalis) begins at cord into two halves ventrally. The ventral
segment L4 and terminates in segment S2, as the median fissure is about 3 mm. in depth. The
segment blends with the conus medullaris. ventral spinal artery is located in the superficial
For descriptive purposes the spinal cord is portion of the fissure.
divided into the cervical part (pars cervicalis),
the thoracic part (pars thoracica), the lumbar Roots of the Spinal Nerves
part (pars lumbalis), and the sacral and coccygeal
parts, or conus medullaris. The spinal nerves are all formed by two roots
The conus medullaris is a general term which from the spinal cord: ( 1 ) a ventral root, also
refers to the tapered, terminal end of the spinal called the motor root, is made up of fibers carry
cord. The term is interpreted differently by vari ing impulses from the spinal cord to the effector
ous authors. Some include only the spinal cord muscle and gland cells, and (2 ) a dorsal root, also
533
N. 1 C - -
(ventral br--
N. 2 C
(dorsal br.
Accessory n.-
Dorsal ro o t n .4 C -
Dorsal
r o o t g a n g l i o n n.8C
N. 1 0 T -
gag'/n
y*\ k\.h.
F 9-1. Dorsal roots of spinal nerves and spinal cord segments, cervical 1 to thoracic 11. Dorsal aspect. The dura mater has
ig .
been removed except on the extreme right side. (The figures on the right represent levels of the vertebral bodies.) (From Fletcher,
1964. Thesis, University of Minnesota.)
534
N. 5Co-
Coccygea/ l i g a m e n t - -J
Fig. 9-2. Dorsal roots of spinal nerves and spinal cord segments, thoracic 11 to coccygeal 5. Dorsal aspect. The dura mater has
been removed except on the extreme right side. (The figures on the right represent levels of the vertebral bodies.) (From Fletcher,
1964. Thesis, University of Minnesota.)
535
536 Chapter 9. The S p in a l C o r d and M e n in g e s
called the sensory root, is made up of fibers The dorsal and ventral columns are largest in
carrying impulses to the spinal cord. the cervical and lumbosacral areas because they
The ventral roots (radices ventrales) arise contain numerous nerves which go to and come
from the cord by a number of roodets (fila radi- from the limbs. The gray matter around the
cularia) at the junction of the lateral and ventral central canal is the central intermediate sub
funiculi. There is no definite ventrolateral sulcus. stance (substantia intermedia centralis). It be
They emerge in several irregular rows in a longi comes continuous laterally with the lateral
tudinal strip 2 to 3 mm. wide on the ventrolateral intermediate substance (substantia intermedia
surface of the cord. The fibers making up the lateralis), which is located between the dorsal
ventral roots are the axons of nerve cells in the and ventral gray columns.
ventral and lateral gray columns of the spinal The substantia gelatinosa is the cover or cap
cord. of the dorsal gray column. In the thoracic and
The dorsal roots (radices dorsales) attach to cranial lumbar portions of the cord the thoracic
the spinal cord in the dorsolateral sulcus. They nucleus (nucleus thoracicus), formerly called the
contain the central processes of the nerve cells nucleus dorsalis or Clarke’s column, is located
in the spinal ganglion. The peripheral processes at the junction of the dorsal column and the gray
of these cells are distributed to the sensory end commissure. The cells in it receive impulses from
ings in the muscles, skin, viscera, and other body fibers in the dorsal funiculus and transmit im
structures. pulses via their axons, which form the dorsal
spinocerebellar tract, to the cerebellar cortex.
Cauda Equina (Fig. 9-3) The lateral column (columna lateralis) is the
lateral protuberance from the lateral intermedi
Cauda equina is the term applied to the caudal ate substance. It is most prominent in the tho-
portion of the spinal cord and the spinal nerves. raco-lumbar portion of the spinal cord and is
This bundle of parallel nerves results from the absent in the other areas.
differential growth in length between the verte White matter (substantia alba) (Fig. 9-4).
bral column on one hand and the spinal cord on The white matter is composed of fibers, myeli
the other. Early in the development of the em nated and unmyelinated, in a network of neu
bryo, the spinal nerves leave the spinal cord at roglia cells. The white matter is divided into six
right angles and pass through the intervertebral funiculi, three on each side. The two dorsal
foramina. But as the fetus grows in size the verte funiculi (funiculi dorsales) lie on each side of the
bral column becomes longer than the spinal median plane between the dorsolateral sulci.
cord, and the spinal nerves have to course The lateral funiculi (funiculi laterales) lie be
obliquely caudally in the vertebral canal before tween the roots of the spinal nerves. The ventral
entering the intervertebral foramina. This is funiculi (funiculi ventrales) are located on each
most noted in the caudal lumbar, sacral, and side of the ventral median fissure and between
coccygeal areas of the spinal cord. In man this the sites of emergence of the ventral roots of the
differential is much greater than in the dog, spinal nerves.
since the spinal cord ends in the area of the first The funiculi at different levels of the spinal
lumbar vertebra. cord vary in shape and size because of additions
and terminations of the fibers in the ascending
Internal Structure of the Spinal Cord and descending fiber tracts, and because of vari
ations in the size and shape of the gray matter.
In contrast to the brain, the white matter of Many of the fibers in the spinal cord originate
the spinal cord is found superficially, and the and terminate in the spinal cord as the fasciculi
gray matter deeply. The remains of the cavity of proprii, which surround the gray matter and
the neural tube is the centrally located central provide for coordinated activity. The majority
canal (canalis centralis). of the fibers in the funiculi are gathered into
Gray matter (substantia grisea) (Fig. 9-4). functional units called tracts, which are com
The gray matter resembles the letter H in cross posed of longitudinally coursing fibers connect
section. The dorsal protuberances are called the ing the dorsal and ventral roots of the spinal
dorsal columns (columna dorsalis), and the ven nerves with other parts of the spinal cord and
tral protuberances the ventral columns (columna with the brain. Kitchell and Stromberg (1958)
ventralis). They are often referred to as the dorsal state that many of the tracts have not been dem
gray horn and the ventral gray horn, respec onstrated as discrete pathways in domestic ani
tively. mals.
S p in a l C ord 537
[d orsal br.-\^~'~% f V
[ventral br -
D e n t i c u l a t e /ig■--
A*
Lumbosacral-fi
e nl a r g e me n t
D ur a ma t e r ( r ef l e c t e d )
Termi nati on of
D e n t i c u l a t e lig.
D o r s a l r o o t IS -
S e g me n t 5Co-
Dors al root
F i l u m t e r m in
D o r s a l r o o t gang I i
Do r s a l r o o t IS —
C auda e y u i n a -
F ig . 9-3. Distal end of spinal cord showing spinal cord segments. Dorsal aspect. (The dura mater has been cut mid-dorsally
and reflected. The dorsal roots have been cut on the left side to expose the ventral roots and the denticulate hgament.) (From
Fletcher, 1964. Thesis, University of Minnesota.)
Fasciculus g ra c ilis Dorsal median sulcus septum
F asciculus cuneatus Dorsal funiculus
D o r s o la t e r a l sulcus
D orsolateral tr a c t
Dorsal g r a y colum n
Lateral corticospinal tract
Substantia ge lo tino sa
D o r s a l spi nocer ebel l a r t r a c t
L a te r a l reticulospinal t r a c t Nucleus dorsalis
Ftubrospinal t r a c t
- L a t e r a l g ra y column
F a sciculi pro prii
L a te ra l funiculus
Vent, s p i n a c e r e b e l l a r t n
Lat. s p i n o t h a l a m i c tr.
S ubsta ntia intermedia
Vestibulospinal t r a c t lateralis
Spi notec ta I t r a c t S ub sta nt ia intermedia
S pino-alivory tract centralis
V e n tr a I s p in o th a la m ic t r a c t V e n t r a l g r a y c ol umn
Central canal
V e n tra l reticulospinal tra ct
Tectospinal tra ct Ventral funiculus
V entral corticospinal tract Ventral m edian f i s s u r e
B TI2 SI
F ig . 9-4. Schema of the spinal cord in cross section.
A. Spinal tracts (the distribution of the tracts is theoretical)
B. Cross sections at selected levels
538
M e n in g e s 539
D o r s a l F u n ic u l u s . The dorsal funiculus is spinal cord of the dog: (a) ventral corticospinal
composed of two large ascending fiber tracts tract, (b ) vestibulospinal tract, (c) ventral spino
called the fasciculus cuneatus and fasciculus thalamic, (d) ventral fasciculus proprius, (e) ven
gracilis. The existence of the semilunar fasciculus tral reticulospinal tract, (f) tectospinal tract,
(fasciculus semilunaris or f. interfascicularis) has and (g) spino-olivary tract.
not been determined in the dog. The fasciculi
proprii are thin and adjacent to the gray sub MENINGES
stance.
The fasciculus gracilis lies on either side of The meninges are the fibrous membranes
the dorsal median septum and extends the entire which surround and protect the spinal cord and
length of the spinal cord. Fibers from the dorsal the brain. They are composed of three mem
spinal nerve roots are added to it laterally as it branes: the dura mater, the arachnoid, and the
ascends in the sacral, lumbar and thoracic areas pia mater. The dura mater is sometimes referred
caudal to the mid-thoracic level. The fibers from to as the pachymeninx, because of its tough,
the lower levels remain medial in position, and, fibrous nature. The combined arachnoid and pia
by a laminating process, fibers from higher mater is called the leptomeninx because of its
levels assume a more lateral position. Many of thinness.
the fibers probably give off collaterals which end
in relation to cells in the nucleus thoracicus, Cranial Meninges
nucleus proprius and ventral horn. Some fibers
may end in these nuclei. The fibers which give Cranial dura mater (dura mater encephali).
off the collaterals mentioned above continue The dura mater of the cranial cavity serves a
craniad with those that do not give off collaterals dual function and consists of two layers, an in
to end in the nucleus gracilis of the medulla ob ternal or meningeal layer and an external or en
longata. dosteal layer. They are closely united except
The fasciculus cuneatus begins in the mid- where venous sinuses are located between them.
thoracic area and is located lateral to the fascicu The outer layer is closely adherent to the bones
lus gracilis in the cranial thoracic and cervical forming the cranial cavity. At the foramina
areas. It increases in size as it ascends to the through which the cranial nerves leave the cra
brain stem. It is formed in a similar manner to nial cavity the dura mater contributes to the
the fasciculus gracilis, but from the fibers of the sheath of the nerves. The meningeal layer is con
cranial thoracic and cervical spinal nerve roots. tinuous with the dura mater of the spinal cord at
The fibers give off collaterals similar to those of the foramen magnum. The meningeal layer of
the fasciculus gracilis, but those that reach the the cranial dura mater in the dog forms three
brain stem end in the internal cuneate nucleus. internal processes between parts of the brain:
L a t e r a l F u n ic u l u s . The lateral funiculus of (a) the falx cerebri, (b) the tentorium cerebelli,
the spinal cord of the dog has not been studied (c) the diaphragma sellae.
thoroughly. The following tracts (Fig. 9-4, A) are F a l x C e r e b r i . The falx cerebri is the mid-
thought to be present: sagittal sickle-shaped fold of dura mater extend
1. Ascending: (a) ventral spinocerebellar tract, ing ventrally between the cerebral hemispheres
(b) dorsal spinocerebellar tract, (c) lateral of the brain. It is fused with the tentorium cere
spinothalamic tract, (d) spinotectal tract, and belli caudally and to the crista galli rostrally.
(e) dorsolateral tract. The straight venous sinus is contained in the
2. Descending: (a) lateral corticospinal tract, junction of the falx cerebri with the tentorium
(b) rubrospinal tract, (c) lateral reticulospinal cerebelli. The dorsal sagittal sinus is located in
tract, (d) tectospinal tract, (e) lateral vestibu the dorsal convex border of the falx cerebri
lospinal tract, and ( f ) olivospinal tract. where the meningeal layer is separated from the
V e n t r a l F u n ic u l u s . The ventral funiculus endosteal layer. The ventral border is 1 to 2 cm.
is composed of the white substance lateral to the dorsal to the corpus callosum of the brain and
ventral median fissure and medial to the emer does not contain an inferior sagittal sinus as in
gence of the ventral roots of the spinal nerves. man. It is in close relation with the genu of the
The two ventral funiculi are joined by the white corpus callosum rostrally and attaches to the
commissure (commissura alba) ventral to the ethmoid and presphenoid bones on the mid line.
central canal. T e n t o r iu m C e r e b e l l i . The tentorium cere
The following tracts are found in many spe belli is the transverse partition between the cere
cies, but have not been clearly defined in the bellum and the occipital poles of the cerebral
540 Chapter 9. The S p in a l C o r d and M e n in g e s
hemispheres. In the dog the dorsocaudal half obtaining a sample of cerebrospinal fluid in the
contains the osseous tentorium cerebelli, a pro dog. Other cisterns present are the chiasmatic
jection of the parietal bone. The deep half is (cisterna chiasmatis), interpeduncular (cistema
composed only of the meningeal layer of the interpeduncularis), and the cistern o f the lateral
dura mater. The internal concave border is free fossa (cisterna fossae lateralis cerebri), which
and forms the border of the tentorial notch (in- are all relatively small.
cisura tentoria), which is occupied primarily by The cranial subarachnoid cavity is continuous
the midbrain. Ventrolaterally, the tentorium with the spinal subarachnoid cavity at the fora
cerebelli attaches to the upper ridge of the men magnum. It communicates with the
petrous temporal bone, and contains a portion of ventricular system of the brain via the lateral
the dorsal petrosal venous sinus. Ventrally, the apertures or foramina of Luschka of the fourth
tentorium blends with the diaphragma sellae at ventricle. The cranial subarachnoid space has ex
the dorsum sellae, forming the roof of the cavern tensions around the optic and olfactory nerves,
ous sinuses. The transverse venous sinus is pri the perineural spaces. The subarachnoid space
marily located in the transverse canal of the does not invest the hypophysis in the dog, ac
osseous tentorium. The trochlear nerve is im cording to Schwartz (1936). Rather, the cisterna
bedded in the free border of the tentorium cere chiasmatis extends only as far caudally as the
belli for a short distance. proximal end of the pars distalis. At this point
D ia p h r a g m a S e l l a e . The diaphragma sellae the subarachnoid cavity courses around the pars
is a circular horizontal diaphragm that bridges tuberalis caudally, joining the rostral boundary
the sella turcica. It is continuous with the ten of the cistema interpeduncularis. The dura ex
torium cerebelli and the posterior clinoid proc tends throughout the diaphragma sellae and
esses. It attaches rostrally to the anterior clinoid blends imperceptibly with the hypophyseal cap
processes. It has a round foramen through which sule, leaving no subdural space.
the infundibulum of the pituitary gland passes. The arachnoid granulations (granulationes
The pituitary gland lies in the sella turcica, par arachnoideales), also known as pacchionian
tially covered by the diaphragma sellae. granulations, are enlargements of the arachnoid
The dura mater passes out through the optic villi. They project through the meningeal layer
canal and forms a tube around the optic nerve in of the dura mater into the dorsal sagittal and ,
its extracranial course. It also surrounds the ol other venous sinuses. At birth arachnoid granula
factory nerves as they pass through the foramina tions are imperceptible in man, but by the age of
of the ethmoid bone. eighteen months they are visible on close inspec
Cranial arachnoid (arachnoidea encephali). tion and at three years are widespread (Davson
The cranial arachnoid is a delicate avascular 1956).
membrane which loosely surrounds the brain. It Weed (1914) describes the arachnoid villi in
is connected to the pia mater by many thin con the dog as being microscopic in size and com
nective tissue trabeculae, which pass through posed of small tufts of arachnoid cells that project
the subarachnoid cavity located between the pia into the venous sinuses, particularly the cavern
mater and the arachnoid. The outer surface of ous and dorsal sagittal venous sinuses. Fank-
the arachnoid is covered by a layer of flat meso- hauser (1962a) did not find arachnoid granula
thelial cells, as is the inner surface of the menin tions in the dog and concluded that they existed
geal layer of the dura mater. A narrow space, the only in large animals, especially horses.
subdural cavity, is located between the dura The arachnoid villi are the principal route
mater and arachnoid and contains a clear yellow through which the cerebrospinal fluid leaves the
fluid. In the live animal the pressure of the cere subarachnoid cavity to enter the venous system
brospinal fluid in the subarachnoid space pushes (Weed 1923).
the arachnoid against the dura mater, making Cranial pia mater (pia mater encephali). The
the subdural cavity almost nonexistent. pia mater is a thin connective tissue membrane
The subarachnoid cavity (cavum subarach- that is closely adherent to the brain. It is highly
noideale) is enlarged by separation of the arach vascularized and extends deep into the sulci of
noid and pia mater. Subarachnoid cisterns the cerebral hemispheres and between the folia
(cisternae subarachnoideales) are present. The of the cerebellum. It receives the trabeculae of
largest is the cerebellomedullary cistern (cistema the arachnoid and forms the internal wall of the
cerebellomedullaris), or cisterna magna, located subarachnoid cavity.
in the angle between the cerebellum and the The pia mater and arteries combine to form
medulla oblongata. This is a common site for the tela choroidea of the choroid plexuses of the
M e n in g e s 541
ventricles. The choroid plexuses are composed The spinal dura mater is in the form of a long
of the tela choroidea and the layer of modified tube surrounding the spinal cord. It has lateral
ependyma covering them. The tissue between tubular extensions which cover the spinal nerve
the blood in the vessels of the tela choroidea and roots and accompany them to the intervertebral
the cerebrospinal fluid in the ventricles may be foramina. As the dorsal and ventral roots join to
spoken of as the “blood-cerebrospinal fluid bar form the spinal nerve, the dura mater blends to
rier” (Gardner et al. 1960). form a single sheath which continues as the epi-
The choroid plexuses of the lateral ventricles neurium of the spinal nerve.
project through the choroid fissure and are con The capillary space between the dura mater
tinuous with the choroid plexus of the third ven and the arachnoid is the subdural cavity (cavum
tricle at the interventricular foramina. subdurale), which contains a small amount of
The choroid plexus of the fourth ventricle is fluid.
relatively large and is an invagination of the roof Caudally, the spinal dura mater tapers to a
of the fourth ventricle. It has two L-shaped parts point and forms a part of the filum terminale
which extend caudally from the junction of the (filum durae matris spinalis). The dura surrounds
pons and medulla, through the lateral apertures the filum terminale of the spinal pia mater, which
of the fourth ventricle, into the subarachnoid fuses to it, and then extends caudally to attach
cavity, to the eighth cranial nerve. to the periosteum of the spinal canal at the sev
The choroid plexuses produce the major por enth or eighth coccygeal vertebra. It serves to
tion of the cerebrospinal fluid. attach the dural sac and spinal cord caudally.
Spinal arachnoid (arachnoidea spinalis). The
Spinal Meninges (Fig. 9-5) spinal arachnoid is a thin, almost transparent
tube which envelopes the spinal cord and has,
Spinal dura mater (dura mater spinalis). The like the spinal dura mater, tubular extensions
spinal dura mater consists of only one layer, the surrounding the dorsal and ventral spinal nerve
meningeal layer. It is separated from the perios roots. It is continuous with the cranial arachnoid
teum of the vertebrae by the epidural cavity at the foramen magnum and ends caudally in a
(cavum epidurale). The epidural cavity is filled cone-shaped sac. It forms part of the filum ter
by a semifluid fat (at body temperature) and by minale.
the vertebral venous sinuses. The spinal dura The spinal arachnoid is connected to the spinal
mater is continuous with the meningeal layer pia mater by connective tissue trabeculae which
of the cranial dura mater at the foramen mag pass through the subarachnoid cavity.
num. The subarachnoid cavity (cavum subarach-
Pia m ater
Subarachnoid c a v ity
Dorsa/ root
Arachnoid membrane
Subdural cavity
P o r s a / ✓ v ent r al Dura m a t e r
roots in adherent Do r s a l + v e n t r a l r o o t s
m e n i n g e a l tubes in separate
me n i n g e a l tubes
V entra l root
Cl to C 5 ij C6 Caudad
F ig . 9 -5 . Schem a o f spinal meninges.
542 Chapter 9. The S p in a l C o r d and M e n in g e s
noideale) is the space between the spinal pia tribute to the formation of the cerebrospinal
mater and the arachnoid membrane. It is filled fluid.
with cerebrospinal fluid which pushes the arach The fluid produced by the choroid plexus
noid peripherally and holds it in contact with (plexus choroideus ventriculi lateralis) of each
the spinal dura mater. The subarachnoid cavity lateral ventricle drains into the third ventricle
extends along the spinal nerve roots for variable through the interventricular foram en. The cho
distances and blends with the epineurium of the roid plexus o f the third ventricle (plexus cho
nerves, prior to fusing with the dura. The cere roideus ventriculi tertii) adds fluid to that from
brospinal fluid serves to cushion and protect the the lateral ventricles. It then passes through the
spinal cord. cerebral aqueduct (aqueductus cerebri) of the
The lumbar cistern of the spinal subarachnoid mesencephalon to the fourth ventricle. The cho
cavity envelopes the spinal nerves of the cauda roid plexus o f the fourth ventricle (plexus choroi
equina. The cistern is narrow at the level of the deus ventriculi quarti) adds a relatively large
lumbosacral foramen, gradually tapers to a amount to the cerebrospinal fluid, which leaves
point, and ends at the level of the first sacral ver the fourth ventricle by way of the lateral aper
tebrae. Fankhauser (1962) states that one can tures o f the fourth ventricle (apertura lateralis
obtain a few drops of cerebrospinal fluid by ventriculi quarti) to enter the subarachnoid cav
lumbar puncture, especially in larger dogs. How ity. A small amount also enters the central canal
ever, only puncture of cerebellomedullary cis of the medulla oblongata and spinal cord. The
tern furnishes a quantity of spinal fluid sufficient median aperture o f the fourth ventricle (apertura
for extensive analysis. mediana ventriculi quarti) or foramen of Magen-
Spinal pia mater (pia mater spinalis). The die is absent in the dog and other animals below
spinal pia mater is similar to the cranial pia the anthropoid apes (Blake 1900, and Schalten
mater, with which it is continuous at the fora brand and Putnam 1927).
men magnum. It is a tough, highly vascularized The cerebrospinal fluid system has been de
membrane that intimately adheres to the spinal scribed as the lymphatic system of the central
cord and roots of the spinal nerves, forming nervous tissue, since no true lymphatics are pres
parts of the epineural sheaths. It is continuous ent. It serves to protect the central nervous sys
with the neuroglial framework of the spinal cord tem from shock and vibration. It also acts as a,
at the sulci. It dips deeply into the ventral fissure fluid buffer and, according to Gardner et al.
and contains all the blood vessels immediately (1960), “compensates for changes in blood vol
peripheral to the spinal cord. ume within the cranium, allowing the cranial
The denticulate ligament (ligamentum den- contents to remain at a fairly constant volume.”
ticulatum) is a condensation of the pia mater on The cerebrospinal fluid is absorbed primarily
the lateral sides of the spinal cord (Fig. 9-3). The by the blood of the cranial venous sinuses
lateral edge of the ligament is free except for the through the arachnoid villi. It is also absorbed by
toothlike serrations which attach to the arach the lymphatics in the olfactory mucosa and in
noid and dura mater. The most cranial attach the epidural tissue of cranial and spinal nerves,
ment is at the foramen magnum. The remaining as they leave their dural and arachnoid sheaths
serrations are attached to the arachnoid and dura (Brierly and Field 1948). Additional sites of ab
mater between successive spinal nerves. The sorption may be located at the larger perivascu
most caudal attachment is at the interspace be lar spaces around blood vessels (Davson 1956).
tween the roots of the fourth and fifth lumbar Studies of the cerebrospinal fluid in the dog
nerves. The denticulate ligament anchors the were made by Davson (1956) and Fankhauser
spinal cord in the center of the subarachnoid (1962). The ventricular system has been de
cavity. scribed in detail by Fitzgerald (1961) and Lim
et al. (1960).
CEREBROSPINAL FLUID
BIBLIOGRAPHY
The cerebrospinal fluid fills the ventricles of Blake, J. A. 1900. The roof and lateral recesses of the fourth
the brain, the central canal of the spinal cord, ventricle considered morphologically and embryologi-
and the subarachnoid cavity. cally. J. comp. Neurol. 10: 79-108.
Brierley, J. B., and E. J. Field. 1948. The connexions of the
The cerebrospinal fluid is produced chiefly
spinal subarachnoid space with the lymphatic system. J.
by the choroid plexuses located in the ventricles. Anat. 82: 153-166.
Schaltenbrand and Putman (1927) demonstrated Davson, H. 1956. Physiology of the Ocular and Cerebrospinal
that the blood vessels in the pia mater also con Fluids. Boston, Little, Brown and Company.
B ib l io g r a p h y 543
Fankhauser, R. 1962. The Cerebrospinal Fluid, Chapter III Atlas of the Dog’s Brain. Springfield, 111., Charles C
in Comparative Neuropathology by J. R. M. Innes and L. Thomas.
Z. Saunders. New York, Academic Press. Schaltenbrand, G., and T. Putman. 1927. Untersuchungen zum
--------------- 1962a. Untersuchungen uber die arachnoidalen Kreislauf der Liquor cerebrospinalis mit Hilfe intravenose
Zotten und Granulationen bei Tieren. Schweiz. Arch, Fluorescineinspritzungen. Dtsch. Z. Nervenheilk. 96:
wiss. prakt. Tierheilk. 104: 13-34. 123-32.
Fitzgerald, T. C. 1961. Anatomy of the cerebral ventricles of Schwartz, H. G. 1936. The meningeal relations of the hypoph
domestic animals. Vet. Med. 56: 38-45. ysis cerebri. Anat. Rec. 67: 35-51.
Gardner, E., D. J. Gray, and R. O’Rahilly. 1960. Anatomy; A Weed, L. H. 1914. Studies on cerebro-spinal fluid. No. III. The
Regional Study of Human Structure. Philadelphia, W. B. pathways of escape from the subarachnoid spaces with
Saunders. particular reference to the arachnoid villi. J. med. Res.
Kitchell, R. L., and M. Stromberg. 1958. Lecture outlines: 31: 51-91.
Fundamentals of neurology. (Mimeographed) College of --------------- 1923. The absorption of cerebrospinal fluid into
Veterinary Medicine, University of Minnesota. the venous system. Amer. J. Anat. 31: 191-221.
Lim, R. K. S., C. Liu, and R. L. Moffitt. I960. A Stereotaxic
CHAPTER 10
General Considerations
fibers in the spinal and frequently in the cranial
The nerves which arise from the brain are re nerves, the efferent and afferent.
ferred to as cranial nerves. The twelve pairs of Cranial nerves can be further subdivided ac
cranial nerves are I, olfactory; II, optic; III, ocu cording to the types of structures which they
lomotor; IV, trochlear; V, trigeminal; VI, abdu supply. Somatic fibers innervate skeletal muscles,
cens; VII, facial; VIII, vestibulocochlear; IX, tendons, joints, and ligaments. Also in this group
glossopharyngeal; X, vagus; XI, accessory; and are most of the sensory fibers from nerve endings
XII, hypoglossal. located in the skin, the somatic afferents. Those
The olfactory and optic nerves are often con fibers which innervate the internal organs,
sidered to be tracts of the central nervous system smooth muscles, glands, and vessels are referred
rather than peripheral nerves. to as visceral fibers. There are four categories
The anatomical organization of cranial nerves of fibers: somatic afferent, somatic efferent, vis
is similar in many ways to that of spinal nerves ceral afferent, and visceral efferent. These prin
(see Chap. 9). Each spinal nerve is attached to ciples apply to the cranial as well as to the spinal
the spinal cord by a ventral and a dorsal root. nerves.
The fibers making up the ventral root emerge Somatic afferent fibers transmit impulses from
as a series of filaments along the ventrolateral the skin muscles and mucous membranes in the
surface of the spinal cord, while those forming cranial area to the brain stem. As in the spinal
the dorsal root emerge dorsolaterally from the cord, the somatic afferent fibers have their cell
spinal cord. The fibers making up the rootlets bodies in sensory ganglia (for the most part),
which unite to form the spinal nerve are derived outside the central nervous system. These are
from a portion of the spinal cord which is re associated with the roots of the cranial nerves.
ferred to as a spinal cord segment. The cells in the ganglia are either bipolar or
Fibers which conduct impulses away from the pseudounipolar, similar to those in the dorsal
spinal cord make up the ventral roots. Their cell root ganglia of the spinal nerves, and give off
bodies are located in the gray matter of the spinal one branch to the brain stem and the other to
cord and are called efferent because they con the periphery.
duct impulses away from the spinal cord. The The visceral afferent fibers in the cranial
fibers which conduct impulses from the periph nerves conduct sensory impulses from the in
ery of the body to the spinal cord form the dorsal ternal organs. For example, the vagus nerve con
roots and are referred to as afferent fibers. They tains visceral afferent fibers from the heart,
have their cell bodies in the spinal or dorsal root lungs, and stomach. Cell bodies of these vagal
ganglia. The cell bodies are considered to be fibers are found in the nodose (inferior) gan
unipolar or pseudounipolar in that they have a glion, separate from those containing the somatic
single fiber from the cell body which divides in afferent cell bodies, which are located in the
the form of a T, one going centrally to the spinal jugular (superior) ganglion. In the spinal nerves,
cord and the other peripherally to a neurosen- both the somatic and visceral afferent fibers have
sory organ. The central branch ends by synaps- their cell bodies in the dorsal root ganglia.
ing with nerve cells in the gray matter of the The visceral efferent fibers of the cranial
spinal cord. Thus there are two main groups of nerves are those fibers which have their cell
bodies aggregated to form particular nuclei in
544
G en era l C o n s id e r a t io n s 545
E t h m o i d a l n. , F r o n t a l bone
V ome r o n a s a l n e r v e s -M u c o s a o f e t h m o t u r b i n o t e s
N a s a l bone -Mucosa of fr o n ta l sinus
N a s a l mucasa
•E t h m a i d a l n.
Dorsal p a r i e ta l ,
nasal c a r t C rib rifo rm plate of ethm oid
bone w i t h c u t e nd s af
o l f a c t o r y nn., I
- -P resphenoid
~ - 01 f a c t o r y n n. , I
' -Vome r
Incisive '-N a s a l pharynx
Nasopalatine duct ' Br. o f c a u d a l n a s a l n.
V o m e r o n a s a l Organ ' P a l a t i n e bone
Maxilla'i ' S e p t a l br. o f c a u d a l n a s a l n.
Fir,. 10-1. Sagittal section of the nose with the cartilaginous and bony nasal septum removed to show distribution of nerves on
the septal mucosa.
Nasolacrim al duct
,M a x i l l a r y br. o f V
, Mucosa of m a x i I l a r y s i n u s
i E t h m a i d a l n.
Location o f d o r s a l e t h m o id a l c r e s t
! Location a f maxi I la tu rb in a te c re s t
i P a r i e t a l nasal
cartilages
Infraorbital n
Pterygopalatine 1
g a n g I i on ' Nasal mucosa
M i n o r p a l a t i n e n. , 1Max i l i a
C a u d a l n a s a l n. i ' B r a n c h e s to m a x i l l o t u r b i n o t e
1
M a j o r p a l a t i n e n, ' B r a n c h e s to m a x i 11ary s i n u s
A c c e s s a r y p a l a t i n e nJ Septal br.
the brain stem. These fibers are preganglionic. branchial musculature develop into the ventro
The nerve cells in these nuclei are of the same lateral muscles of the neck and the trapezius
type as those in the intermediolateral column of muscle, which are supplied by cranial nerve XI
the spinal cord, being medium-sized and multi or the spinal accessory nerve.
polar. The largest of the visceral efferent cranial
nerve nuclei is the dorsal motor nucleus of the OLFACTORY NERVES
vagus (parasympathetic). All the visceral effer
ent fibers leaving the brain in the cranial nerves The olfactory nerves (nn. olfactorii) consist of
belong to the craniosacral (parasympathetic) many small bundles of nerve fibers which pass
division of the autonomic nervous system. They from the olfactory mucosa to the olfactory bulb
terminate on nerve cell bodies which are in (Fig. 10-1). They are classified as special vis
groups or scattered collections forming the vis ceral afferent fibers. The cell bodies of the nerve
ceral (autonomic) ganglia. The visceral ganglia fibers are located in the olfactory nasal mucosa.
are in or on the walls of the organs innervated. The central processes of the neurons travel cau
The fibers of the nerve cells in the visceral gan dally in the submucosa to the cribriform plate of
glia are postganglionic. the ethmoid bone. The fibers pass through the
The somatic efferent fibers arise from nuclei many foramina of the cribriform plate to end in
composed of large multipolar nerve cell bodies the olfactory bulb. The bundles of fibers are en
which usually contain large amounts of chromo- closed by the dura mater, arachnoid, and pia
phil (Nissl) substance. mater, as they pass through the foramina.
In addition to the four functional types just Read (1908) described the extent of the ol
enumerated, there are three additional types of factory mucosa in the dog adjacent to the cribri
fibers found in cranial nerves that are not found form plate. It occupied the surface of about half
in the spinal nerves: (1 ) the special visceral a f of the numerous ethmoturbinates, the caudal
ferent, (2) the special som atic afferent, and (3) third to half of the nasal septum, one ethmo-
the special visceral efferent fibers. turbinate scroll which extended into the anterior
The special visceral afferent fibers are those portion of the frontal sinus, and a portion of the
which come from the visceral sensory organs medial wall of the sinus. The olfactory mucosa
(taste and olfaction). The special visceral affer can be distinguished in the fresh specimen by its
ent fibers are incorporated in cranial nerves I, yellow to brown color, due to the pigment of the
V, VII, IX, and X, which convey impulses from sustentacular cells.
the olfactory mucosa and the mucosa of the The vomeronasal nerves (nn. vomeronasales)
tongue. arise from the dorsomedial surface of the vomer
The .special somatic afferent fibers come from onasal organ and unite to form six to eight nerve
the special sensory organs which are considered bundles (Fig. 10-1). The bundles further unite,
to be derived from the ectodermal layer of the as they pass caudally in the submucosa of the
embryo (the eye and the ear). nasal septum, to form one or two nerve trunks
The special visceral efferent or branchial mo which pass through foramina of the ethmoid
tor fibers are found in cranial nerves V, VII, IX, bone (cribriform plate). The nerve fibers pass
X, and XI. These nerves supply motor fibers to caudally on the medial surface of the olfactory
striated muscle of branchial arch origin. The bulb to end on the accessory olfactory bulb,
first branchial arch is supplied by the trigeminal which is located on the medial surface of the ol
(V) nerve. The musculature of the first branchial factory tract at the caudal edge of the olfactory
arch gives rise to the muscles of mastication, bulb. The vomeronasal nerves are the axis cylin
which are supplied by the mandibular division der processes of the neurosensory cells in the
of the trigeminal nerve. The musculature of the mucosa of the vomeronasal organ. McCotter
second branchial arch develops into the muscles (1912) has provided a detailed description of
of facial expression and several other muscles the vomeronasal nerves and their connections.
(stapedius and caudal portion of the digastricus) The terminal nerve (n. terminalis) of the dog
which are supplied by the facial (VII) nerve. was described by McCotter (1913) as “a gan-
The remaining branchial arches, three through glionated nerve connected with the vomeronasal
six, and sometimes seven, are supplied by cranial nerves on one hand and apparently with the fore
nerves IX and X. The musculature of the latter- brain on the other, having thereby the same
named branchial arches differentiates into the morphological relations in these mammals (dog
muscles of the pharynx, larynx, and cranial por and cat) as described for the terminal nerve of
tion of the esophagus. Some portions of the lower forms.” The terminal nerve is formed by
T roch lear N erve 547
several small bundles which arise from the scopic structure of the oculomotor nerve in the
vomeronasal nerves as they terminate on the dog and found both large and small myelinated
accessory olfactory bulb. The small bundles unite fibers.
to form a single trunk which runs caudoventrally The oculomotor nerve leaves the brain stem
on the medial surface of the olfactory tract and at the medioventral surface of the mesencepha
appears to enter the brain substance 1 or 2 cm. lon, just medial to the cerebral peduncle. It
caudal to the olfactory bulb. courses rostrolaterally and enters the wall of the
cavernous sinus. The oculomotor nerve emerges
OPTIC NERVE through the wall of the cavernous sinus and
leaves the cranial cavity by way of the orbital
The second cranial or optic nerve (n. opticus) fissure (Figs. 10-3, 10-4, 10-6). Immediately
is composed of the central processes of the gan upon emergence from the orbital fissure it di
glion cells of the retina, which converge at the vides into dorsal and ventral rami.
optic disc and pass through the choroid and scle The dorsal ramus, after a short rostrodorsal
ral layers, lateroventral to the caudal pole of the course, divides into several branches which en
eyeball. The fibers of the optic nerve are classi ter the muscular portion of the dorsal rectus
fied as special somatic afferents. The optic nerve muscle at its caudal end (Figs. 10-4, 10-6). After
is surrounded by extensions of the cranial menin coursing through the muscle to approximately
ges throughout its slightly sinuous course from the junction of the distal and middle thirds, a
the eyeball to the optic canal, where it enters small branch becomes superficial and enters the
the cranial cavity. ventral surface of the levator palpebrae muscle,
The optic nerve (and its meninges) is within a in its distal portion.
cone formed by the retractor bulbi muscles. It The ventral ramus, much the larger of the two
is related to the orbital fat and to the ciliary rami, travels rostrally and gives off branches to
iierves and vessels in the rostral portion of their the medial rectus, ventral rectus, and ventral
course (Figs. 10-3, 10-4). The ciliary ganglion oblique muscles of the eye. The parasympa
is bounded ventrolaterally by the lateral rectus thetic or general visceral efferent fibers leave
muscle and laterally by the ventral portion of the ventral ramus to enter the ciliary ganglion.
the retractor bulbi muscle and medially by the The ventral ramus continues rostrally for ap
optic nerve. Dorsally, the ophthalmic arteries proximately 3 to 4 cm. (Figs. 10-3, 10-4). It
and the nasociliary nerve are adjacent to the divides into two branches. One passes medio-
meningeal coverings in the caudal portion of dorsally and ventral to the optic nerve to enter
the orbit. The optic nerves join at the optic the medial rectus muscle. The other travels ros
chiasm shortly after entering the cranial cavity, trolaterally and divides into two branches which
where fibers from the medial portions of both supply the ventral rectus and ventral oblique
nerves cross to the optic tract of the opposite muscles. The branch to the ventral rectus enters
side. its dorsal surface at the junction of the proximal
Bruesch and Arey (1942) state that the optic and middle thirds. The branch to the ventral
nerve of the dog contains 154,000 fibers, all of oblique passes laterad, ventral to the ventrolat
which are myelinated. The fibers have no neuri eral portion of the retractor bulbi muscle and
lemma and become myelinated as soon as they then rostrally to enter the middle of the caudo-
leave the eyeball. The optic nerve may be con dorsal surface of the ventral oblique muscle. The
sidered to be a tract of the brain. Developmen- ciliary ganglion is located at the point where the
tally, it is the connection between the dienceph ventral ramus divides into branches which sup
alon and the retina, a derivative of the brain ply the medial rectus, the ventral rectus, and
(Gardner et al. 1960.) the ventral oblique muscles. The oculomotor
root of the ciliary ganglion is very short and
OCULOMOTOR NERVE leaves the ventral ramus of the oculomotor nerve
in the angle formed by the divergence of the
The oculomotor nerve (n. oculomotorius) is two previously mentioned branches. The oculo
the principal nerve to the muscles of the orbit. motor root of the ciliary ganglion contains pre
It contains two types of fibers: (1) general so ganglionic parasympathetic nerve fibers.
matic efferent, which are motor to the striated
ocular muscles, and (2 ) general visceral efferent TROCHLEAR NERVE
(parasympathetic), which are destined for the
ciliary ganglion. Koch (1916) studied the micro The fourth cranial or trochlear nerve (n. troch-
548 Chapter 10. The C r a n ia l N erv es
F r o n t a l n.fbr. o f o p h t h a l m i c ) s
I n f r o tro c h I ea r
/ L a c rim a l g la n d
S h o r t c i I i a r y nn. n
Z y g o m a t i c o t e m p o r a l n.
C i I i a r y aa. /
' ,B r. to l a c r i m a l g l a n d
E x t. e t h m o i d a l a. „
^ Z y g o m a t i c o f a c i a l n.
C r i b r i fo r m p ia te v v ^ x
T r o c h I e a r n. to ^ v If T / j ' To m• r ? c f us v e n t r a l is
J U 'I f _ - T o m. o b i i g u u s v e n t r a l i s
m. o b i i q u u s d o rs.
" " "•
I nt. o p h t h a l m i c a- " « 7a lr~ ~ C i l i a r y g o n g I i o n
E th m a id a I n - fv 7y ~ -To m. r e c t u s m e d i a l i s
Z-Onc^ c i I i a r y n. ^ ^ v - ~ X’,
Z '- To m. r e c t u s l a t e r a l i s
To mm. r e c t u s d o rs . — Nn- to m. r e t r a c t o r b u lb i
v le v a to r p a lp e b ra e - _ _ - A b d u c e n s n.
N a s o c i l i a r y n. - O c u lo m o t o r n.
(b r.o f o p h t h a lm ic )
" Ex t. oph t h a l m ic a.
A n t. c e r e b r a l a . ' " " ^ O r b i t a l a.
s.
O p t i c nn., I I - " v ' • L a c r i m a l n. (br. o f f r o n t a l )
In t. o p h t h a l m i c a . ' O p h th a l m i c br. o f V
Long c i l i a r y n.\
M. o b i ic^uus d o r s .
To m. rec tus med I rtf r a t r o c h l e a r n.
E th m o id a l n, sr M. r e c t u s dors, ( cut)
e x t . e t h m o i d a l a.
n (cut)
Troch I e a r n.(cut)^
M r e t r a c t o r bulbi^
( dors , l at .v do r s . med.) v
Opt i c n.x
6 'V
To r ect us do r s , v
l e v a t o r pa I p e b r a e
La c r i m a l n L a c rim a l g la n d
O c u l o m o t o r n.~ _
T ro c h le a r n — $'\ '•M. r e c t u s l a t ( c u t )
*
N asoc i h o r y n<
F r o n t a t n.-
IL o b h q u u s vent.
M a x i 11a r t ) a. ■ / i j \To m . o b l i q u u s v e n t
2 iqomatic n ' J ' ':!^ - ^ J ^ T o jT ^ r e c tu s v e n t .
O r b i t a l a .' 1 \ S h o r t c i l i a r y nn.
M a x i H a r y br. o f V ' j / ' C ilia r y a a n q lio n
A b d u c e n s nJ j \M. r e t r a c t o i b u l b i ( v e n t l a t ? v e n t, m ed.)
To m. r e t r a c t o r b u l b i ' 1 M. r e c t u s v e n t r a l i s
I
M. r e c t u s l a t ( c u t ) l O c u l o m o t o r n.
learis) is the smallest of the twelve cranial nerves. nerve, arises rostrally and leaves the cranial
It is unusual in several respects. The trochlear cavity through the orbital fissure. The second di
nerve is the only cranial nerve to emerge from vision, the maxillary nerve, arises from the rostro-
the dorsal surface of the brain stem. The right lateral side and leaves via the round foramen to
and left trochlear nerves decussate (decussatio enter the alar canal. The third division, the man
nervorum trochlearium) in the rostral medullary dibular nerve, arises from the lateral side of the
velum caudal to the caudal opening of the cere ganglion, caudal to the maxillary nerve, and
bral aqueduct. Shortly after emerging from the emerges through the oval foramen. Koch (1916),
brain stem the trochlear nerve pierces the dura in studying the intracranial portion of the trigem
mater and runs in the ventrolateral extension of inal nerve in the dog, found both large and small
the tentorium cerebelli along the dorsal ridge of myelinated fibers and small numbers of un
the spine of the petrous temporal bone and myelinated fibers, which are largely associated
passes dorsally to the semilunar ganglion (Figs. with the sensory portion of the nerve.
10-3, 10-4). The trochlear nerve remains in the
dura until it passes through the orbital fissure O p h t h a l m ic N e r v e
between the ophthalmic branch of the trigem
inal nerve and the oculomotor nerve. Upon The ophthalmic nerve (n. ophthalmicus) (Fig.
emergence from the fissure it turns dorsomediad 10 - 6 ) is the principal sensory nerve of the orbit,
and enters the dorsomedial surface of the dorsal the skin on the dorsum of the nose, and a portion
oblique muscle (Figs. 10-4, 10-6). This is the of the mucous membranes of the nasal cavity
only muscle supplied by the trochlear nerve. In and paranasal sinuses. It is the smallest division
its course from the orbital fissure to the dorsal of the trigeminal nerve. It arises from the trigem
oblique muscle, the nerve is related ventrally to inal ganglion and passes rostrally in the lateral
the dorsal rectus and levator palpebrae muscles. wall of the cavernous sinus, ventral to the troch
Koch (1916) found both large and small myeli lear nerve. It receives filaments from the
nated fibers in the trochlear nerve of the dog. cavernous plexus of sympathetic nerves and
usually connects with the oculomotor, trochlear
TRIGEMINAL NERVE and abducens nerves. The three primary
branches, frontal, lacrimal, and nasociliary
The fifth cranial or trigeminal nerve (n. tri nerves, arise in or near the orbital fissure.
geminus) is the largest of the cranial nerves. Its The frontal (or supraorbital) nerve (n. fron
sensory fibers (somatic afferents) receive im talis) (Figs. 1 0 -3 ,1 0 -4 ,1 0 -5 ,1 0 -6 ) is the sensory
pulses from the cutaneous muscles of the head, nerve to the middle portion of the upper eyelid
the nasal and oral cavities, and the muscles of and adjacent skin. It arises from the ophthalmic
mastication. The motor fibers (special visceral nerve, in the orbital fissure, in common with the
efferents) supply the muscles which are derived lacrimal nerve. It passes rostrodorsally deep to
from the first branchial arch of the embryo, the periorbita and dorsal to the dorsal rectus and
principally the muscles of mastication. levator palpebrae muscles. In the rostrodorsal
The trigeminal nerve is attached to the brain portion of the orbit it lies between the tendon
stem at the junction of the pons and trapezoid of the dorsal oblique muscle and the supraorbital
body. Its two roots, a small motor (radix motoria) process of the frontal bone, lateral to the troch
and a large sensory (radix sensoria), are usually lea for the tendon of the dorsal oblique muscle.
not separable. The motor and sensory roots, in The lacrimal nerve (n. lacrimalis) (Fig. 10-6)
their common sheath, pass through the canal of is a very small branch which arises from the
the petrosus temporal bone and expand into the ophthalmic division of the trigeminal nerve as
semilunar-shaped trigeminal ganglion (ganglion stated above. The lacrimal nerve travels rostro
trigeminale), which is located in the cavum tri- dorsally along the lateral edge of the dorsal
geminale of the dura mater lateral to the caver rectus muscle to end in the lacrimal gland.
nous sinus at the apex of the petrous temporal The nasociliary nerve (n. nasociliaris) (Fig.
bone. 10-3) is the continuation of the ophthalmic divi
The trigeminal ganglion contains most of the sion into the orbit. It passes ventral to the troch
unipolar cell bodies of the general somatic lear nerve and between the dorsal and ventral
afferent fibers which are distributed by the rami of the oculomotor nerve to reach the dorsal
branches of the trigeminal nerve. The three di part of the optic nerve. There it gives off the long
visions of the trigeminal nerve arise at the tri ciliary nerve, which travels rostrally parallel to
geminal ganglion. The first, the ophthalmic the optic nerve. It joins the short ciliary nerves
(Text continued on page 554.)
T r ig e m in a l N erve 551
Z y g o m a t i c n.
Moxi l l a r y b r a n c h
.O phthalm ic b ra n c h
peep temporal branch
i Zygomaticotemporal n.
( { Z i j t j o m a t i c o f a c i a I n.
I / Su p r a o r b i t a I n. ( F r o n t a I )
I I / i n f r a t r o c h I e a r n. ( c u t )
Rostral i n te r n a I P t e r y g o p a l a t i ne n.
a u r i c u l a r n. M i n o r p a l a t i n e n.
Dorsal a l v e o la r branches
/
/
B r a n c h e s to s k i
I n f r a o r b i t a I n.
Anast omoses wi th
r o s t r a l a u r i c u l a r ____
X Branches to ~
v i b r i ssae r ski n
Ma n d i b u I a r b r a n c h -
B r a n c h e s to p a r o t i d g l a n d
A u r i c u l o t e m p o r a ! n.- "
Common t r u n k of /
m a s s e t e r i c v d e e p t e m p o r a l nn- /
/ Rostra I
Chorda t y mpa n i ' / / m e n t a l n.
M y l o h y o i d n. Anast. w i t h
L i n g u a l n. vent, b u c c a l n.
Se c r e t o r y b r a n c h 1 1M i d d l e m e n t a l n.
To m . d i g a s t r i c u s ' / t C a u d a l m e n t a l n.
A n a s t o m o s e s w i t h v e n t r a l t > u c c a l n. 1 / To m. m y l o h y o i d e u s
N. b u c c a I i s ^ To vi b r i s s a e •? s k i n
M a n d i b u l a r a l v e o l a r n. To m u c o s a v s k i n
T r o c h le a fo r t e n d o n o f m .o b l iq^uus d o r s . v
M. o b i i g u u s d o r s . v
I n f r a t r o c h l e a r n.x ^
M. r e t r a c t o r b u l b i ^ ^ L a c rim a l g la n d
M. r e c t u s m e d i a I i s ^ - O r b it a l lig . (c u t)
To m. le v a t o r p a l p e b r a e - _ j _ Z y g o m a t i c o f a c i a l n.
I _ - Z y g o m a tic a rc h (c u t)
Cmn+nl r, ''rJM
— Br. to l a c r i m a l g l a n d
■ i In f.___ Z y g o m a t i c o t e m p o r a l n.
E th m o id a l a - "
£
%r— M. r e c t u s d o r s a l i s
£■— M. r e c t u s l a t e r a l i s
M. levator palpebrae ■
-Tom . r e c tu s la t e r a lis
Troc h I ea r n.- "■ ' 4 -T o m- r e t r a c t o r b u l b i
7o m m r e ct us dorsal is) _ " L a c r i m a I n.
r l evator palpebrae] / x.
A b d u c e n s n.
Optic n n - " 'P e r io r b ita I fa s c ia
Orbital fissu re- ■V,'!
Abducens n.,VI " . .
O p h th a lm ic b r of V
Oculomotor 'jjg i
• Z y g o m a t i c b r. o f V
B r a n c h e s tc p e r i o r b i t a x
Ai.of p te ry g o id co n a l
P t e r y g o p a la t in e n
Z y g o m a tic b ra n c h y - M . p t e r icjoideus med.
Foram en fo r
M a x illa ry b r o f V %
" m, o b lig u u s v e n t r a l i s
M. p t e r y g o i d e u s l a t
- - I n f r a o r b i t a l n.
M . p t e r y g o i d e u s med.
- D o r s a l a l v e o l a r n.
- - C a u d a l n a s a l n.
M a j o r p a l a t i n e n.
Pterygopalatine ganglion A c c e s s o r y p a l a t i n e n,
M i n o r p a l a t i n e n.'
Fic.. 10- 7. The pterygopalatine ganglion Lateral aspect.
L a c r im a l g la n d
Z y g o m a t i c o t e m p o r a l n.\
/Z y g o m a t i c o f a c i a l n
B r a n c h e s to p e r , o r b i t a \
P e r io rb ita v
N. o f p te r y g o id ca n a l
P t e r y g o p a l a t 'r ie n.
Z y g o m a tic b ra n c h
- M .o b h g u u s v e n t r a l i s
M a x i l l a r y br. o f V
- In fra o rb ita l n
- C audal n a s a l n
M a j o r p a l a t i n e n.
'-A c c e s s o ry p a l a t i n e n.
M- pterygoideus lat.' n e n.
M. p t e r y g o i d e u s med. D o rs a l a l v e o l a r n.
Pteri.:i c jo p a la tin e q a r g i i a n
F it. 10-8. Maxillary branch of the trigeminal nerve. Lateral aspect.
55 4 Chapter 10. The C r a n ia l N erv es
from the ciliary ganglion and enters the eyeball. in the alar canal on its dorsal side, it may lie in
The fibers in the long ciliary nerve are primarily the zygomatic groove on the dorsal wall of the
sensory to the structures of the eyeball. In addi alar canal, or it may course in the zygomatic
tion, it contains some postganglionic sympathetic canal of the sphenoid bone and emerge from the
fibers from the cavernous plexus. zygomatic foramen (Fig. 10-7) (McClure 1960).
A connection between the nasociliary nerve The zygomatic nerve enters the apex of the
and the ciliary ganglion may be present. The periorbita soon after emerging from the cranial
nasociliary nerve, after giving off the long ciliary cavity. It divides into two rami, the zygomatico
nerve, divides into the infratrochlear and temporal and zygomaticofacial (Figs. 10-3, 10-
ethm oidal nerves. The infratrochlear nerve (n. 5, 10-8). This division may occur before the
infratrochlearis) (Figs. 10-3, 10-4, 10-6) passes zygomatic nerve leaves the bony canal or after
rostrodorsally along the lateral edge of the dorsal it enters the periorbita.
oblique muscle. It passes ventral to the trochlea The zygomaticotemporal nerve (n. zygomati-
for the tendon of the dorsal oblique muscle and cotemporalis) is the most dorsal of the two
ramifies in the medial portion of the upper eye branches. It courses rostrad and dorsad im
lid and adjacent skin. The ethm oidal nerve (n. mediately under the periorbita and penetrates
ethmoidalis) (Figs. 10-3, 10-4) may be regarded the caudal edge of the orbital ligament to ramify
as the continuation of the nasociliary nerve. It in the lateral portion of the upper eyelid and
is sometimes referred to as the external eth skin of the rostral temporal area. In the rostro
moidal nerve because it accompanies the ex dorsal portion of the periorbita the nerve is
ternal ethmoidal artery in a portion of its course. related to the dorsolateral aspect of the lacrimal
The ethmoidal nerve runs rostrally and medially, gland, to which it gives off some branches. These
passing between the dorsal oblique and medial are believed to be the parasympathetic secretory
rectus muscles to enter the ventral ethmoid fibers to the gland because of the communica
foramen. From the ventral ethmoid foramen it tions between the pterygopalatine ganglion and
enters the cranial cavity and courses dorsally the trigeminal nerve.
and rostrally in a small groove on the lateral wall. The zygomaticofacial nerve (n. zygomatico-
It leaves the cranial cavity through a foramen in facialis) parallels the zygomaticotemporal ramus
the dorsomedial aspect of the cribriform plate. in the periorbita (Figs. 10-3, 10-6, 10-8). In the
It contributes sensory branches (n. nasales rostral portion of the periorbita it deviates ven
intemi) to the nasal turbinates as it courses trally from the other ramus and emerges ventral
rostrally. The nerve terminates rostrally as the to the lateral canthus of the eye. It ramifies in the
external nasal nerve (n. nasalis externus), which lateral portion of the lower eyelid and adjacent
is distributed to the skin of the muzzle rostral to area of skin (Fig. 10-10).
the nasal bone (Fig. 10-2). The next branches of the maxillary nerve are
the pterygopalatine nerves (nn. pterygopalatini),
M a x il l a r y N erve numbering from one to three small branches,
which communicate with the pterygopalatine
The maxillary nerve (n. maxillaris) is the larg ganglion (Figs. 10-7, 10-8). They are composed
est of the trigeminal divisions. It is the sensory primarily of sensory fibers which pass through
nerve to the skin of the cheek, side of the nose, the ganglion to enter the maxillary nerve.
muzzle, mucous membrane of the nasopharynx, The next three nerves usually leave the ven
maxillary sinus, soft and hard palates, and the tral aspect of the maxillary nerve as a common
teeth and gingivae of the upper jaw. The nerve trunk, as it courses rostrally between the medial
leaves the trigeminal ganglion and passes ros pterygoid muscle and the ventral surface of the
trally in the dura mater of the lateral wall of the periorbita. They receive many branches from
cavernous sinus to the round foramen. It leaves the pterygopalatine ganglion (Fig. 10-7).
the cranial cavity by way of the round foramen The minor palatine nerve (n. palatinus minor)
to enter the alar canal. It turns rostrally in the accompanies the minor palatine artery between
alar canal and is related to the maxillary artery, the maxillary bone and medial pterygoid muscle
which it accompanies across the pterygopalatine to reach the soft palate (Figs. 10-2, 10-8). It
fossa, after leaving the anterior alar foramen. contains sensory fibers from the mucosa as well
The zygomatic nerve (n. zygomaticus) is the as motor fibers which go from the pterygopala
first branch of the maxillary division (Figs. 10-3, tine ganglion to the gland cells in the mucosa.
10-4). It leaves the parent trunk near the round The major palatine nerve (n. palatinus major)
foramen. It may accompany the parent nerve accompanies the major palatine artery and
T r ig e m in a l Nerve 555
The deep temporal nerve (n. temporalis pro the lateral side of the mandible dorsal to the
fundus) is usually composed of two parts. One digastricus muscle and deep to the masseteric
part arises in common with the buccal nerve and muscle to anastomose with the ventral buccal
the other in common with the masseteric nerve nerve (Figs. 10-5, 10-10).
(Fig. 10-9). The deep temporal nerve enters the The mylohyoid nerve passes rostrally on the
temporalis muscle. ventral surface of the mylohyoid muscle and
The masseteric nerve (n. massetericus) is the supplies it with many fine motor rami. It also
lateral portion of the dorsal trunk of the man supplies small rami to the skin of the interman-
dibular nerve. After giving origin to one of the dibular area. A larger ramus goes to the sinus or
deep temporal nerve branches, it passes laterad, tactile hairs (vibrissae).
caudal to the temporalis muscle and through the The mandibular alveolar nerve (n. alveolaris
mandibular notch of the mandible to enter the mandibularis) leaves the ventral lateral trunk of
masseter muscle (Figs. 10-9, 10-11). The nerve the mandibular division of the trigeminal
ramifies in the muscle and is its sole motor nerve nerve and enters the mandibular canal through
supply. the mandibular foramen (Figs. 10-5, 10-9).
The auriculotemporal nerve (n. auriculo- The mandibular alveolar nerve accompanies the
temporalis) arises from the ventrolateral trunk mandibular alveolar artery and gives off sensory
of the mandibular nerve (Fig. 10-9). The fibers branches to the teeth of the mandible. Several
of the auriculotemporal nerve are sensory and branches (mental nerves) leave the nerve rostral
are distributed to the skin in conjunction with ly and pass out through the mental foramina
branches of the facial nerve. It passes laterally, (Figs. 10-5, 10-10). The mental nerves are dis
ventrally, and caudally to the glenoid process tributed to the skin ventral to the lower incisor
of the squamous temporal bone. It emerges be teeth.
tween the caudodorsal border of the masseteric The lingual nerve (n. lingualis) passes rostrally
muscle and the external auditory canal (Fig. 10- on the dorsal surface of the medial pterygoid
11). The auricular branches supply the skin on muscle and turns ventrally over its rostral border.
the lateral surface of the external auditory canal Near its origin from the mandibular nerve, it
and the rostral internal surface of the auricular receives the chorda tympani branch of the facial
cartilage (rostral internal auricular nerve). There nerve (Figs. 10-5, 10-9, 10-11, 10-16). The
are many fine anastomoses which with the lingual nerve gives off one or two small branches
auricular branches anastomose with the to the mucosa of the isthmus of the fauces.
branches of the facial nerve and are distributed The sublingual nerve (n. sublingualis) leaves
to the skin of the temporal and zygomatic areas the rostral surface of the lingual nerve and is
(Fig. 10-10). One large branch goes directly to distributed to the oral mucosa between the man
the sinus or tactile hairs (vibrissae) on the caudo dible and the tongue (Fig. 10-11). The ramus
lateral surface of the cheek. The parotid salivary communicans to the mandibular ganglion leaves
gland also receives many small branches from the caudal surface of the lingual nerve and
the auriculotemporal nerve. courses alongside the mandibular duct to the
The fibers in the parotid nerves arise from the mandibular ganglion located in the hilus of the
otic ganglion. They join the auriculotemporal mandibular salivary gland. The fibers in this
nerve (Fig. 10-9) shortly after its origin from ramus are parasympathetics derived from the
the mandibular nerve and leave it where it is chorda tympani nerve. Some branches are dis
related to the rostral deep aspect of the parotid tributed to the sublingual salivary gland. Usu
gland. They supply parasympathetic fibers to ally there is a small ganglion (sublingual gan
the gland. glion) situated in the angle between the ramus
The m ylohyoid nerve (n. mylohyoideus) con communicans and the lingual nerve. It contains
ducts motor and sensory impulses. It arises from the nerve cell bodies which are secretory to the
the ventral lateral trunk of the mandibular nerve, sublingual salivary gland.
rostral to the origin of the auriculotemporal The main trunk of the lingual nerve continues
nerve (Figs. 10-9, 10-11). The nerve courses into the musculature of the tongue, where it has
ventrally over the rostral border of the medial several anastomoses with the hypoglossal nerve
pterygoid muscle to the ventral border of the (Fig. 10-13) (FitzGerald and Law 1958). The
mandible. At the ventral border of the mandible fibers are distributed to the dorsal mucosa of the
it gives off a branch which supplies motor fibers tongue rostral to the circumvallate papillae. It
to the anterior belly of the digastricus muscle conducts somatic afferent impulses to the brain
(Fig. 10-5). Another branch passes rostrally on stem via the trigeminal nerve. Special visceral
T r ig e m in a l N erve 557
Deep t e m p o r a l b r a n c h e s \ M. p te ry g o id e u s la t.
B u c c a l i s n.x M p t e r y c j o i d eu s med.
To m . p t e r y g o i d e u s /a; j n g u a l n.
To m. te n s o r v e l i p a l a t i n i tym pa ni
- A u r i c u l o t e m p o r a l n.
M a n d ib u la r b ra n c h -
Fac C horda t y m p a n i
/ - - -
V e s tib u la r n /
C o c h le a r n /
G e n ic u la te g a n g I i o n 1 nne«i«o*
F ig. 10-9. The temporal bone sculptured to show the otic ganglion in relation to the trigeminal nerve. Dorsal aspect.
558 Chapter 10. The C r a n ia l N erv es
afferent fibers from the taste buds leave the visceral efferent fibers (preganglionic parasym
lingual nerve, via the chorda tympani nerve, pathetic) are motor to the postganglionic nerve
and enter the brain stem with the facial nerve. cells supplying the glandular cells of the nasal
cavity, the mandibular and sublingual salivary
ABDUCENS NERVE glands and the lacrimal gland. The cell bodies of
the general visceral efferent fibers are located in
The sixth cranial or abducens nerve (n. ab the rostral salivatory nucleus, near the facial
ducens) leaves the ventral surface of the medulla nucleus in the rostral portion of the medulla
oblongata just caudal to the pons. The abducens oblongata. The fibers are distributed by the
nerve conveys motor impulses to the retractor major petrosal and chorda tympani nerve
bulbi and the lateral rectus muscles. Koch (1916) branches to the pterygopalatine, mandibular,
states that both the large and small myelinated and sublingual ganglia.
fibers of the abducens nerve are joined by a large The sensory nerve fibers, both visceral af
bundle of unmyelinated fibers in the cavernous ferent and special visceral afferent, have their
sinus. cell bodies in the geniculate ganglion. The
The nerve courses rostrally in the subarach peripheral processes of the sensory nerves are
noid space and enters the wall of the cavernous distributed via the chorda tympani and major
sinus. It passes rostrally, medial to the trigeminal petrosal nerves, primarily to the taste buds in
ganglion of the fifth cranial nerve (Figs. 10-3, the rostral two-thirds of the tongue and to other
10-4, 10-6), and leaves both the cavernous sinus visceral receptors in the epithelium covering the
wall and the cranial cavity by way of the orbital soft palate, nasopharynx, and nasal cavity.
fissure. It is the most ventral and medial of the
cranial nerves which emerge from the orbital Course of the Facial Nerve
fissure. It is related laterally to the ophthalmic
division of the fifth cranial nerve and to the The central course of the facial nerve fibers
anastomotic branch of the orbital artery. within the brain stem is described in Chapter 8 .
Approximately 1 to 2 cm. rostral to the orbital The facial nerve emerges from the brain stem
fissure, the abducens nerve gives off a branch to at the rostral edge of the trapezoid body lateral
the retractor bulbi muscle. This short branch to the origin of the abducens nerve. After a short
divides into smaller branches which supply both course laterad it becomes closely associated with
the dorsal and ventral parts of the muscle. the vestibulocochlear nerve and accompanies it
into the internal auditory meatus. The facial
FACIAL NERVE nerve diverges from the vestibulocochlear nerve
and enters the facial canal. Upon reaching the
General Considerations genu of the facial canal it turns sharply (about
90 degrees), forming the genu o f the facia l nerve
The facial nerve (n. facialis) is a mixed nerve (geniculum n. facialis), and courses caudally.
containing special visceral efferent (branchial The geniculate ganglion (ganglion geniculi)
motor) and afferent fibers and visceral efferent (Fig. 10-15) is located on the dorsorostral
(parasympathetic) and afferent fibers. border of the facial nerve at the genu (Fig. 10-
The cell bodies of the special visceral efferent 14). The cell bodies of the visceral and special
fibers are located in the fa c ia l nucleus (nucleus visceral afferent fibers of the facial nerve are
n. facialis) in the rostroventral portion of the located in the ganglion.
medulla oblongata. The special visceral efferent The major petrosal nerve (n. petrosus major)
fibers make up the larger portion of the facial arises from the geniculate ganglion and facial
nerve. They are distributed peripherally to the nerve at the genu (Figs. 10-9, 10-15). The nerve
auricular, facial and other musculature derived passes rostroventrad in the petrous temporal
from the second branchial arch, via the caudal bone, emerges, and becomes associated with the
auricular, digastric, stylohyoid, cutaneous colli, auditory tube (Fig. 10-16). The deep petrosal
auriculopalpebral, stapedial, and dorsal and ven nerve (sympathetic fibers) joins with the major
tral buccal nerves. petrosal nerve to form the nerve o f the pterygoid
The special visceral afferent and visceral canal. The nerve of the pterygoid canal enters
efferent and afferent fibers make up a part of the caudal end of the pterygoid canal and
the facial nerve which is often referred to as emerges from the canal in the pterygopalatine
the nervus intermedins. It is not grossly separa fossa and ends in the pterygopalatine ganglion
ble from the rest of the nerve in the dog. The (Figs. 10-7, 10-8, 10-15).
F a c ia l N erve 559
The major petrosal nerve and the nerve of the course caudally parallel to each other and to the
pterygoid canal contain visceral efferent (pre mid line. Ramus I then gives off branches to the
ganglionic parasympathetic) fibers destined for platysma muscle, the cervico-auricularis pro
the pterygopalatine ganglion. The nerve cells in fundus muscle, and the musculi obliquii and
the pterygopalatine ganglion give rise to the transversi auriculae.
postganglionic fibers to the nasal glands and The other caudal auricular nerve (ramus II)
lacrimal gland. Some visceral afferents are con may arise in common with ramus I or separately
tained in the nerves. Their distribution and func from the facial nerve at the stylomastoid fora
tion are uncertain; they may subserve general men. It is larger than ramus I and courses
sensation from the nasal mucosa. dorsocaudally for 3 to 5 cm. on the caudoventral
The nerve to the stapedial muscle (n. sta surface of the temporalis muscle. It turns dor
pedius) (Fig. 10-15) leaves the dorsal medial sally and divides into branches which end in the
surface of the facial nerve just before the latter following muscles: cervico-auricularis profundus
turns ventrolaterad in the facial canal. The facial (medial portion of the major part), cervico-
nerve also receives the auricular branch of the auricularis superficialis, cervico-scutularis, inter-
vagus nerve and gives rise to the chorda tympani parieto-auricularis, interparieto-scutularis, oc
nerve, which enters the cavity of the middle ear. cipitalis, interscutularis, scutulo-auricularis
In its ventrolaterad course the facial nerve re superficialis accessorius, scutulo-auricularis
ceives the auricular branch of the vagus, prior to superficialis medius, scutulo-auricularis profun
emerging from the facial canal at the stylomas dus major and minor, helicis, and mandibulo-
toid foramen (Fig. 10-15). The fibers in the auricularis. The branch to the mandibulo-
branch are somatic afferent, which are dis auricularis muscle courses dorsorostrally around
tributed by the internal auricular nerves to the the medial surface of the external ear canal to
skin on the concave surface of the auricular enter the dorsal portion of the muscle.
cartilage of the ear. Miller and Witter (1942) The digastric nerve (n. digastricus) innervates
state that the cell bodies of the fibers in the the caudal belly of the digastricus muscle. It
auricular branch of the vagus are probably lo leaves the caudoventral surface of the facial
cated in the jugular (superior) ganglion of the nerve immediately after the latter emerges from
vagus nerve. the stylomastoid foramen.
The chorda tympani new e arises from the The caudal internal auricular and the lateral
cranial surface of the facial nerve opposite the internal auricular nerves (Fig. 10-10) arise
junction of the auricular branch of the vagus from the dorsal surface of the facial nerve and
with the facial (Fig. 10-15). The chorda tympani pass through the cartilage to ramify in the skin
nerve enters the cavity of the middle ear. of the ear canal and in the auricular cartilage of
Just prior to emerging from the stylomastoid the ear.
foramen, two or three auricular branches are The stylohyoid nerve (n. stylohyoideus) is a
given off the caudal surface. These branches small nerve which leaves the facial trunk and
accompany the main portion of the facial nerve supplies the stylohyoid muscle (Fig. 10-10).
to emerge from the stylomastoid foramen (Huber
1923). Terminal Branches of the Facial Nerve
The caudal auricular nerves (nn. auriculares
caudales) (Fig. 10-10) are paired and supply the The facial trunk terminates as the auriculo-
caudal or retroauricular musculature. They leave palpebral, the dorsal buccal, and the ventral
the facial nerve as it emerges from the stylo buccal nerve.
mastoid foramen. They travel caudodorsally The auriculopalpebral nerve (n. auriculopal-
from the stylomastoid foramen in company with pebralis) (Fig. 10-10) is distributed to the rostral
the great auricular artery. The most caudal of auricular muscles and the muscles of the eyelids.
the two branches (ramus I) (Fig. 10-10) gives Auricular and palpebral branches arise approxi
off a branch to the cervical portion of the pla mately 1 to 2 cm. from the origin of the parent
tysma muscle. It then passes caudodorsally, trunk. The auricular branches enter the follow
parallel and deep to the muscle 2 or 3 cm. from ing muscles: scutulo-auricularis superficialis
the mid line. The branch to the platysma gradu dorsalis, scutulo-auricularis profundus major,
ally diminishes in size as it gives off fine twigs to and frontalis. The rostral auricular nerve has
the muscle. Grossly, it cannot be traced much several anastomoses with the auricular branches
farther caudally than the middle of the neck. of the auriculotemporal nerve of the trigeminal
Occasionally this branch divides at the cranial nerve. The palpebral branch of the auriculo
edge of the muscle. The two resulting branches palpebral nerve passes dorsorostrally and enters
(Text continued on page 563.)
560 Chapter 10. T h e C r a n ia l N erv es
To m. s c u t u l o a u r i c u l a r i s s u p e r f d o r s .
To m. s c u t u l o a u r i c u l a r i s p r o f. m a j o r
1 , R o s t r a l a u r i c u l a r n.
I ] j P a l p e b r o l n.
R o s t r a l a u r i c u l a r p le x u s
Bn to platysma
( r e t r o a u r i c u l a r n.)
G rea t a u r i c u l a r n
I I C , v e n t r a I br.
B r a n c h e s to s k in
Au r i cu lo r I r a m u s I
b ra n c h e s J ra m u s 2 '
F a c i a l r>.,Vll
Ext. j u g u l a r v.
v M e n t a l nn.
To m . d i g a s t r i c u s
' B u c c a l i s n.
^ ' B r a n c h e s of m y l o h y o i d n.
To m . d e p r e s s o r a u r i c u t o e 1 , ,1 ' D o r s o l b u c c a l n.
To m. s t y l o h y o i d e u S 1 I \ ' Nn. to v i b r i s s a e
L a t. i n t a u r i c u l a r n. J ' P a r o t i d d u c t ( c u t)
C e rv i c a l br.' ' A u r i c u l o t e m p o r a l n.
A u r i c u l o p a l p e b r a I n.1 V e n t r a l b u c c a l n.
Fic. 10-10 Superficial branches ol the facial and trigeminal nerves. Lateral aspect.
F a c ia l N erve 561
M a s s e t e r i c n. 'M ylohyoid n
M. p t e r y g o i d e u s lat. Deep te m poral br.
C h o rd a t y m p a n i Buccal is n.
M a n d ib u la r br. o f V M o n d ib u la r a lv e o la r n.
A u r ic u lo te m p o r a l n. , M.pterygoideus med.
M. s ty lo p h a ry n g e u s ' - U n g u a l n.
S t y lo h y o i d bone ^ M. te m p o r a l is
F o c ia l n., To b u c c a l m ucosa
M. d ig a s tric u s , Z y g o m o tic g l a n d
M a n d ib u la r
g la n d
' Subl in g u a l g la n d ,
p o ly s to m a tic p a r t
i ' M. geniog lossus
, ' M a jo r su bl in g u a l d u c t
'M a n d ib u la r d u c t
M th y ro h y o i d c u *
' M. g e n ia h y o id e u s
Cran. l a r y n g e a l n!
' M. s ty lo g lo s s u s
S u b lin g u a l gland,1
m o n o s to m a tic p a r t 1To mm. s t y l o g l o s s u s v h y o g lo s s u s
M. h yo p h a ry n g e u s ' To m. g e n ia h y o id e u s
H y p o g lo s s a l n 'M. hyoglossus
F ig. 10-11 Nerve distribution medial to the mandible. (The digastricus muscles, the mandible, and structures lateral to it
have been removed.^
562 Chapter 10. The C r a n ia l N erv es
M a n d ib u la r d u c t
M. m y l o h y o i d e u s
S u b lin g u a l d u ct N
.M . g e n i o h y o i d e u s
S u b I i n g u a l n .•?
g a n g lio n
M.genioylossus
L i n g u a l n.
M.s ty I oylossus
R a m u s communi cans
to mandi bul ar g angl i on
,M. hy o y l o s s u s
S u b l i n g u a l salivary Cjland _
- N y p o y l o s s a l n.
M. m y lo h y o id e u s ------ , ~ L i n y u a l a.
M. d i y a s t r i c u s — M. s t yl oh yo i de u s
M o n o s t o m a t i c p a r t of M. i h y r o h y o i d e u s
su b I i n y u a I y l a n d
M. c r i c o t h y r o i d e u s
M a n d ib u la r s a liv a ry y ia n d
C ra n ia l l a r y n y e a l n.'' M. s t e r n o h y o i d e u s
F i r s t c e r v i c a l n. 'M.sternothyroi d e u s
Medial retropharyngeal
lymph n o d e
Common c a r o t id a
Vagosympathetic trunk
M. s t e r n o c e p h a l i c u s
F ig. 10-12. Nerves of the ventral surface of the head and neck.
G lo sso ph a ryn g ea l N erve 563
into the formation of the extensive rostral auric of the cochlea and is concerned with the sense
ular plexus. It anastomoses with the most rostral of hearing. Their origins are described in the
branch of the rostral auricular nerve. The pal section on the vestibulocochlear organ.
pebral nerve supplies the orbicularis oculi and The two parts of the vestibulocochlear nerve
corrugator supercilii muscles and terminates in are enclosed in a common dural sheath (with the
the levator nasolabialis and maxillonasolabialis facial nerve) as they pass through the internal
muscles. acoustic meatus (Figs. 10-9, 10-15). They enter
The dorsal buccal nerve (n. buccalis dorsalis) the brain stem caudal to the pons at the lateral
(Fig. 10-10), the second terminal branch of the end of the trapezoid body.
facial nerve, also has several anastomoses with The cell bodies of the fibers in the vestibular
branches of the auriculotemporal nerve and the nerve are bipolar and located in the vestibular
trigeminal nerve (source of sensory fibers). The ganglion in the vestibular portion of the inner
dorsal buccal nerve passes rostrodorsally, form ear. The cell bodies of the fibers in the cochlear
ing an arc which roughly parallels the zygomatic nerve are bipolar and located in the spiral gan
arch, and anastomoses with the ventral buccal glion in the cochlear portion of the inner ear.
nerve caudodorsal to the commissure of the
mouth. It passes through the orbicularis oris GLOSSOPHARYNGEAL NERVE
muscle and terminates in the maxillonasolabi
alis muscle. In its course through the orbicularis The ninth cranial or glossopharyngeal nerve
oris muscle it also anastomoses with the infra (n. glossopharyngeus) is a mixed nerve. It is
orbital and buccal branches of the trigeminal motor to the stylopharyngeus muscle and to the
nerve. parotid and zygomatic salivary glands, and sen
The ventral buccal nerve (n. buccalis ven sory to the pharynx, a portion of the tongue, and
tralis) (Fig. 10-10) immediately passes ventrally the carotid sinus. The nerve cells of origin for the
and slightly rostrally from the termination of the fibers (special visceral efferents) to the stylo
facial nerve. After coursing 1 to 2 cm., it gives pharyngeus muscle are located in the nucleus
off, caudoventrally, the cervical branch, which ambiguus of the medulla. The nerve cells of
innervates the musculi depressor auriculae and origin for the preganglionic parasympathetic
the sphincter colli primitivus. There are several fibers (general visceral efferents) for the glands
branches which join the cervical branches of the are located in the caudal salivatory nucleus. The
second cervical nerve. The ventral buccal nerve sensory cell bodies are in the petrosal ganglion.
then passes rostrally on the lateral surface of the Their peripheral processes are distributed to the
masseter muscle. Approximately 2 cm. caudal to pharyngeal and lingual mucosa and to the carotid
the commissure of the mouth it divides into sinus. Their central processes end in the nucleus
numerous branches which enter the ventral por of the solitary tract in the cat (Kerr 1962). Kozma
tion of the orbicularis oris muscle. The ventral and Gellert (1959) report that there are numer
buccal nerve, at this point, also receives anasto ous nerve cells in the terminal trunk and the
mosing branches from the mylohyoid nerve of lingual ramus.
the trigeminal. The terminal branches of the The glossopharyngeal nerve arises from the
ventral buccal nerve also join the mental nerves rostral part of the medulla oblongata by several
which are terminal branches of the mandibular small rootlets and leaves the cranial cavity
alveolar nerve. through the jugular foramen (Figs. 10-15, 10-
16). The petrosal (superior) ganglion (g. petro-
VESTIBULOCOCHLEAR NERVE sum) is located in the jugular foramen. It is small
and produces only a slight enlargement of the
The eighth cranial or vestibulocochlear nerve glossopharyngeal nerve (Fig. 10-16). Grossly, no
(n. vestibulocochlearis), formerly referred to as inferior ganglion is present in the dog.
the auditory, acoustic or statoacoustic nerve, is The glossopharyngeal nerve gives rise to the
composed of afferent fibers from the internal following nerves.
ear. The fibers are arranged in two bundles, the The tympanic nerve (n. tympanicus) leaves
vestibular and cochlear parts (nerves) (Figs. 10- the glossopharyngeal nerve at the level of the
9,10-15). The vestibular nerve (pars vestibularis) petrosal ganglion and enters the cavity of the
is involved with the sense of balance and is dis middle ear. It divides into several branches to
tributed to the cristae ampullares of the semi form the tympanic plexus on the promontory of
circular canals and the maculae of the saccule the petrous temporal bone. The minor petrosal
and utricle. The cochlear nerve (pars cochlearis) nerve (n. petrosus minor) arises from the tym-
is distributed to the hair cells in the spiral organ (Text continued on page 567.)
5 64 Chapter 10. The C r a n ia l N erv es
M. l e v a t o r v e li p a l a t i n i te n s o r v e li p a l a t i n i
Esophagus
, ' M. g e n io g lo s s u s
, 'M. g e n io h y o id e u s
, ' L in g u a l n . ( c u t r r e f l e c t e d )
'/W. s ty lo g lo s s u s ( c u t r r e f l e c t e d )
M. th y ro p h a ry n g e u s ' m . m ylo hyo ide us (c u t «■ r e f l e c t e d )
M. h y o p h a ry n g e u s 1 , 'H y p o g lo s s a l n. ( c u t)
L i n g u a l br. o f I X ' 'L o c a tio n of p a la tin e to n s il
E p ih y o id b one1 [M. h y o g lo s su s
10-13. Deep dissection of the pharyngeal region and tongue showing distribution of the glossopharyngeal, hypoglossal,
F ig .
and lingual nerves.
G lo sso ph a ryn g ea l N erve 565
V a gu s n., X
H y p o g l o s s a l n., X I I
N o d o s e g an glion 4
Int. c a r o t i d a.
O c c i p it a l a.
G lo s s o p h a r y n g e a l n „ I X
A c c e s s o ry n, X I v
, Cran. c e r v i c a l g a n g l i o n
To mm. sternomosToideus C lX
sM. s t y l o p h a r y n g e u s
r d eidom astoideus
- P h aryn ge al b r
M. s te rnom ostoideus
. _ L i n g u a l br.
M. d e id o m a s to id e u s ^
v P h a ry n g o e s o p h a g e a l br.
Br. fro m cran. cerv. g a n g l i o n „
-To c a r o t i d s i n u s
(
Ext. c a r o t i d a.
To m. tr a p e z iu s
_ L i n g u a l a.
Vagosympathetic tr u n k In te rn a l
Cran. th y r o i d O. ^ c a r o t i d a.
M. omotransverso ri us
I n.
so c e r v i c a l is
l a r y n g e a l n.*a.
Common
c a r o t i d ay ' M. m y lo h y o id e u s
Esophagus '/W. h yo g lo s s u s
Caudal l a r y n g e a l nJ M. h y o p h a ry n g e u s
T h yro id g la n d \M. s t y l o h y o id e u s
M .s t e r n a t h y r a i d e u s ' , M. t h y r o h y o i d e u s
M .s te rn o h y o id eus 1 'i n t e r n a l br.
M. c ric o p h a ry n g e u s 1 E x t e r n a l br.
M. c r ic o t h y r o id e u s M. t h y r o p h a r y n g e u s
F ig. 10-14. Nerves of the pharyngeal region. Lateral aspect. The digastricus muscle and superficial structures have been
removed.)
566 Chapter 10. The C r a n ia l N erv es
M. t e n s o r t y m p a n i
M a jo r petrosal n
Chorda ty m p a n i
T r i g e m i n a l n.
M i n o r p e t r o s a I n.
A b d u c e n s n.
- Head o f m alleus
Gen i c u l a t e g a n g l ion - J
Retroglenoid canal
r ' - S h o r t cr u s o f incus
F a cia l n
.Y — S t a p e s i n o v a l w i n d o w
V e s t i b u l a r n.~
- - V estibule
C o c h l e a r n.
L o c a t i o n o f m- s t a p e d i u s
- Chorda t y m p a n i
G l o s s o p h a r y n g e a I n.~ N. to m. s t a p e d i u s
J u g u l a r g a n g l i o n ----- W A u r i c u l a r b ra n c h e s o f VII
Vagus n.~~ A u ricular branch o fX
A c c e s s o ry n." .
H y p o g l o s s a l n."
C o n d y lo id canal
F ig . 10-15. The petrous temporal bone sculptured to show the path of the facial nerve. Dorsal aspect.
V agus N erv e 567
panic plexus and passes dorsally to gain the muscle and glands of other thoracic and ab
dorsal aspect of the tensor tympani muscle (Figs. dominal viscera.
10-15, 10-16). The nerve then passes rostrally Special Visceral Efferent. These supply motor
on the dorsolateral aspect of the auditory tube to innervation to the musculature derived from the
the otic ganglion (Fig. 10-9). The preganglionic last three branchial arches of the embryo and to
parasympathetic fibers in the nerve synapse with the esophageal musculature. These include the
the postganglionic cells in the otic ganglion muscles of the pharynx (except the stylopharyn
which are distributed to the parotid and zygo geus) and the larynx.
matic salivary glands. General Visceral Afferent. These visceral sen
The pharyngeal ramus (r. pharyngeus) of the sory nerves transmit impulses from the base of
glossopharyngeal nerve leaves the parent nerve the tongue, pharynx, esophagus, stomach, intes
ventral to the petro-occipital fissure at the ven tines, larynx, trachea, bronchi, lungs, heart, and
tral border of the tympanic bulla (Fig. 10-12). other viscera.
The glossopharyngeal nerve and the pharyngeal Special Visceral Afferent. A few sensory fibers
ramus are lateral to the cranial cervical ganglion come from the small number of taste buds on the
(Fig. 10-14). The pharyngeal ramus passes epiglottis and pharyngeal wall.
rostrally, medial to the stylopharyngeus muscle General Somatic Afferent. These are the fibers
and the stylohyoid bone, on the dorsolateral which come from the skin of the external ear
aspect of the pterygopharyngeus muscle. The canal. They form the auricular branch of the
ramus courses around the rostral edge of the vagus, which joins the facial.
levator veli palatini muscle and ramifies in the The vagus nerve arises from the medulla ob
mucosa of the dorsal aspect of the pharynx (Fig. longata of the brain stem by a series of fine root
10-13). The pharyngeal ramus frequently re lets. It passes through the middle portion of the
ceives a branch from the pharyngeal ramus of the jugular foramen (Fig. 10-15). The jugular (su
vagus nerve and sometimes a branch from the perior) ganglion (g. jugulare), which contains the
cranial cervical ganglion. unipolar nerve cells of the general somatic affer
The lingual ramrn (r. lingualis) leaves the ent fibers of the vagus nerve, lies in the jugular
glossopharyngeal nerve a short distance distal to foramen.
the pharyngeal ramus (Fig. 10-17). It passes The vagus nerve leaves the jugular foramen
through the stylopharyngeus muscle, to which and enters the petro-occipital fissure. It lies
motor branches are given off (special visceral caudal to the glossopharyngeal nerve and the
efferents), and ramifies in the mucosa of the internal carotid artery and rostral to the acces
lateral pharyngeal wall and the palatine tonsil sory nerve (Figs. 10-14, 10-16). Ventral and
(Fig. 10-13). medial to the tympanic bulla, it presents the
The ramus to the carotid sinus (r. sinus ca nodose (inferior) ganglion (g. nodosum), which
rotid) leaves the glossopharyngeal nerve and is prominent. Distal to the nodose ganglion it is
parallels the internal carotid artery to the carotid joined by the sympathetic trunk and becomes
sinus. The ramus is joined by a branch from the the vagal part of the vagosympathetic trunk,
cranial cervical ganglion and often forms a which travels in the carotid sheath (with the
plexus of several nerves which end at the carotid common carotid artery) to the thoracic inlet.
sinus and around the arteries at the termination Branches in the H ead and Cranial Cervical
of the common carotid artery. The glossopharyn Region. The auricular ramus (r. auricularis)
geal nerve occasionally sends a branch to the leaves the vagus nerve at the level of the jugular
pharyngoesophageal nerve of the vagus (Fig. ganglion, and travels laterad through the caudal
10-13). edge of the petrous temporal bone to the facial
canal (Figs. 10-15, 10-16). The fibers of the
VAGUS NERVE auricular ramus join with the facial nerve and
are distributed to the external acoustic meatus
The tenth cranial or vagus nerve (n. vagus) is by way of the auricular branches of the facial
the longest of the cranial nerves. It traverses the nerve (Fig. 10-10).
neck, thorax, and abdomen. A listing of its com The pharyngeal ramus (r. pharyngeus) leaves
ponents indicates its distribution and functions. the vagus nerve at the proximal end of the no
General Visceral Efferent. These are the pre dose ganglion. It sends a branch rostrally to join
ganglionic parasympathetic nerve fibers that with the pharyngeal ramus of the glossopharyn
supply the muscle of the heart and the smooth geal nerve. The main trunk of the pharyngeal
568 C h ap ter 10. T h e C r a n ia l N erv es
/ , Lingual n
M a n d i b u l a r a l v e o l a r n.
I
M y l o h y o i d n. ^
B u c c a l i s n.
Mass sTo m p t e r y g o i d e u s l o t
s ' ^To m. p t e r y g o i d e u s med.
AuriCulOtempora I -To m t e n s o r i/ e l i p a l a t i n i
Otic gang!ion
M a n d i b u l a r b r of V
■Osseous a u d i t o r y t u b e
To m t e n s o r t y m p a m
To a u d i t o r y t u b e w a l l
M i n o r p e t r o s a l n. _
- C a r o t id foramen
Refroqlenoid f o r a m e n _
V. C a r o t i d ca n a l
M tensor tyrnpani _
Ext acoustic Ty mp a n i c b u l l a (cut)
meatus, d o rs a l w a l I - - Foromen f o r
x - v e n t r a l p e t r o s a l sinus
M a l l e u s j - --------
M in o r petrosaln. - ' Int. c a r o t i d nn.
Incus Petrobasi la r canol
Stapes ~ j-
" T y m p a n i c plexus
Chorda
P ro m o n to ry
C ra n. c e r v ic a l
g a n g lio n
To nn. I X r X
N
.
> N Glossopharyngealn.JX
F a c i a l n., V I I
/
Round w in d o w ' , \ \ Hypoglossal n„ XU
A u r i c u l a r b r of vagus n ! , ' i ,\ Jugular foramen
Tym panic n! I Petrosal ganglion
V a g u s n., X ■A c c e s s o r y n „ X I
Fic. 10-16, Nerves in the region of the middle ear. Ventral aspect. 'The tympaim bulla is removed.)
V agus Nerve 569
To l a r y n g e a l m u c o s a
To t r a c h e a ^
To m. c r ic o a r y t e n o i d e u s la t ."
- -T h y ro id c a r tila g e
To m. v o c a l is " cut r re fle c te d
To rr. t h y r o a r y t e n o i d e u s
P e tro s o l g a n g lio n To n u c le u s o f V II
C ra n ia l p o rt of XI \
\ I To t r a c t u s s o l i t a r i u s
i VII ;
S p in a l p a r t o f XI
' To n u c le u s s a l i v a t o r i u s S u p e r i o r
J u g u l a r g a n g lio n _
x T ym p a n ic n.
N od ose g a n g l io n /
V a g o s y m p a th e tic t r u n k __, T ym p a nic plexus
, ' " „ - M in o r p e t r o s a l n.
Cron, la ry n g e a l n - - ' O t i c g a n g lio n
To p a r o tid v z y g o m a tic
Cran. c e rv ic a l g a n g lio n
g la n d s
,M a jo r p e t r o s a l n
To pharyngeal p le x u s "
N. of p te r y g o id
To c a r o t i d p le x u s
canal
Int. c a r o t i d n n .' /
A u r i c u l a r br. o f X '
’ P t e r y g o p a l a t in e g a n g lio n
C h o rd a t y m p a n i < ( b r a n c h e s to l a c r i m a l ,
G e n ic u la te g a n g lio n / j n a s a l , p a l a t in e g la n d s )
M a n d ib u la r g a n g ! io n ’ j Max i I l a r y br. o f V
C o m m u n ic a tin g ra m u s 1 L i n g u a l n.
S u b lin g u a l g a n g lio n 1
F ig . 10-18. Schema of eranial nerves VII. IX. \, and XI. and autonomic interconnections.
570 Chapter 10. The C r a n ia l N erv es
ramus is short and divides into many smaller esophagus at its cranial end. The caudal laryn
branches to form the pharyngeal plexus (plexus geal nerve is the motor nerve to all the intrinsic
pharyngeus) (Figs. 10-13, 10-14). The cranial muscles of the larynx, except the cricothyroid
cervical ganglion also contributes several muscle.
branches to the pharyngeal plexus. A larger The remaining branches of the vagus nerve
branch in the plexus, the pharyngoesophageal are described in Chapter 12.
branch (n. pharyngoesophageus), usually re
ceives branches from the glossopharyngeal nerve ACCESSORY NERVE
and the cranial cervical ganglion and is distrib
uted to the caudal pharyngeal muscles (crico- The eleventh cranial or accessory nerve (n.
pharyngeus and thyropharyngeus) and to the accessorius) has a spinal origin in addition to
esophagus. Hwang et al. (1948) report that the several small rootlets of origin (radices craniales)
pharyngoesophageal branch supplies the cranial from the medulla oblongata of the brain stem.
two-thirds of the cervical esophagus and, in some The spinal origin (radices spinales) arises from
dogs, the entire cervical portion of the esopha cervical segments one to seven of the spinal cord
gus. (Fig. 9-1). The spinal rootlets of origin are lo
The cranial laryngeal nerve (n. laryngeus cated between the dorsal and ventral roots of the
cranialis) leaves the vagus nerve at the nodose cervical spinal nerves and form the spinal root,
ganglion (Figs. 10-13, 10-14). It passes ven which is located on the dorsal side of the denticu
trally, medial to the occipital and common ca late ligament. The spinal root enters the cranial
rotid arteries, and divides into an external branch cavity through the foramen magnum and is
(r. externus) and an internal branch (r. internus) joined by several rootlets from the medulla
on the cranial portion of the thyropharyngeus oblongata and enters the jugular foramen caudal
muscle (Fig. 10-14). to the vagus nerve (Fig. 10-15). A branch (r.
The external branch travels caudally on the internus) leaves the accessory nerve in the
ventral portion of the thyropharyngeus muscle. jugular foramen and joins the vagus nerve distal
A branch leaves the external branch ventrally to the jugular ganglion via common epineural
and passes deep to the insertion of the sterno- sheaths. DuBois and Foley (1936) found in the
thyroideus muscle, to supply the cricothyroideus cat that the branch from the accessory to the
muscle. The external branch continues caudad vagus nerve contained fibers from the medulla
and ends in the area of the thyroid gland (Fig. oblongata and that these conveyed motor im
10-14). pulses in the caudal laryngeal nerve.
The internal branch passes between the thy The nerve cell bodies for the spinal root are
ropharyngeus and hyopharyngeus muscles located in the dorsolateral part of the ventral
cranial to the edge of the thyroid cartilage to gray column in the spinal cord. The fibers of the
enter the larynx. It ramifies in the mucosal lining cranial portion originate from nerve cells in the
of the larynx. Before entering the cavity of the caudal portion of the nucleus ambiguus.
larynx, the internal branch gives off a branch The trunk of the accessory nerve (ramus ex
caudally which anastomoses with the caudal ternus) leaves the jugular foramen and turns
laryngeal nerve (n. laryngeus caudalis) (Fig. 10- caudally in the petro-occipital fissure (Fig. 10-
17). 16). It passes lateral to the hypoglossal nerve and
The recurrent laryngeal nerve (n. laryngeus enters the cranial portion of the combined
recurrens) arises from the vagus nerve at the cleidomastoideus and sternomastoideus muscles.
thoracic inlet. The right recurrent nerve leaves The nerve then divides into two parts, a ventral
the right vagus nerve and passes caudal to the ramus and a dorsal ramus.
right subclavian artery and then cranially on the The ventral ramus supplies the cleidomas
right dorsolateral aspect of the trachea. The left toideus and sternomastoideus muscles. It gives
recurrent nerve leaves the left vagus nerve and off many branches as it passes caudally and ven
passes caudally around the aortic arch and then trally on the medial side of the muscle. It is fre
cranially between the esophagus and the trachea. quently embedded in the muscle tissue (Fig.
Both recurrent laryngeal nerves give off rami to 10-14).
the esophagus (r. esophagei) and the trachea (r. The dorsal ramus penetrates the dorsal aspect
tracheales) as they run cranially. of the cleidomastoideus muscle under the wing
The recurrent laryngeal nerve terminates as of the atlas and passes caudally deep to the
the caudal laryngeal nerve (Fig. 10-17). It also cleidocervicalis muscle. It gives off several
anastomoses with the nerves supplying the branches to the muscle.
H ypo g lo ssa l N er v e 571
The dorsal ramus has a number of anastomoses the hypoglossal nerve and the rami of the lingual
with the second, third, and fourth cervical nerve (Fig. 10-13) (FitzGerald and Law 1958).
nerves. The dorsal ramus parallels the dorsal
border of the omotransversarius muscle. At the BIBLIOGRAPHY
shoulder it travels between the supraspinatus
muscle and the cranial portion of the trapezius Bruesh, S. R., and L. B. Arey. 1942. The number of myelinated
muscles. It gives off branches to the cranial por and unmyelinated fibers in the optic nerve of vertebrates.
tion of the trapezius muscle and terminates in its J. comp. Neurol. 77: 631-665.
DuBois, F. S., and J. O. Foley. 1936. Experimental studies on
caudal portion. These branches to the trapezius the vagus and spinal accessory nerves in the cat. Anat.
muscle constitute its sole motor supply. Rec. 64: 285-307.
FitzGerald, M. J. T., and M. E. Law. 1958. The peripheral
HYPOGLOSSAL NERVE connexions between the lingual and hypoglossal nerves.
J. Anat. (Lond.) 92; 178-188.
Gardner, E., D. J. Gray, and R. O’Rahilly. 1960. Anatomy: A
The twelfth cranial or hypoglossal nerve (n. Regional Study of Human Structure. Philadelphia, W. B.
hypoglossus) arises as a series of rootlets from the Saunders.
ventrolateral sulcus of the medulla oblongata. Huber, E. 1923. Uber das Muskelgebiet des Nervus facialis
The nerve leaves the cranial cavity through the beim Hund, nebst allgemeinen Beotrachtungen uber die
Facialis-muskulatur. Morph. Jb. II Teil. 52: 353-414.
hypoglossal foramen (Figs. 10-15, 10-16). It Hwang, K., M. I. Grossman, and A. C. Ivy. 1948. Nervous con
passes medial to the accessory nerve and lateral trol of the cervical portion of the esophagus. Amer. J.
to the vagus nerve, sympathetic trunk, and in Physiol. 154: 343-357.
ternal carotid artery (Fig. 10-14). Kerr, F. W. L. 1962. Facial, vagal and glossopharyngeal nerves
in the cat. Arch. Neurol. (Chic.) 6: 264-281.
The hypoglossal nerve passes ventrally and Koch, S. L. 1916. Structure of the third, fourth, fifth, sixth,
gives rise to a descending branch which anasto ninth, eleventh and twelfth cranial nerves. J. comp.
moses with the ventral branch of the first cervical Neurol. 26; 541-552.
nerve, to form a part of the ansa cervicalis (Fig. Kozma, A., and A. Gellert. 1959. Vergleichende histologische
Untersuchungen uber die mikroskopischen Ganglien und
10-14). Another descending branch leaves the
Nervenzellen des Nervus glossopharyngeus. Anat. Anz.
caudal side of the hypoglossal nerve about 2 cm. 106: 38-49.
distal to the first and joins the ventral branch of McClure, R. C. 1960. Occurrence of the zygomatic groove and
the first cervical nerve on the dorsal aspect of canal in the sphenoid bone of the dog skull (Canis fa
the stemohyoideus muscle. miliaris). Abstract, Anat. Rec. 138: 366.
McCotter, R. E. 1912. The connection of the vomeronasal
The hypoglossal nerve curves rostrally on the nerves with the accessory olfactory bulb in the opossum
lateral surface of the hyopharyngeus and hyo and other mammals. Anat. Rec. 6: 299-317.
glossus muscles (Figs. 10-12, 10-14). Deep to --------------- 1913. The nervus terminalis in the adult dog and
the mylohyoideus it gives rise to rami which sup cat. J. comp. Neurol. 23: 145-152.
ply the styloglossus, hyoglossus, genioglossus, Miller, M. E., and R. E. Witter. 1942. Applied anatomy of the
external ear of the dog. Cornell Vet. 32: 65-86.
and geniohyoideus muscles as well as the intrin Read, E. A. 1908. A contribution to the knowledge of the ol
sic muscle fibers of the tongue (Fig. 10-13). factory apparatus in dog, cat and man. Amer. J. Anat.
Several anastomoses occur between the rami of 8: 17-48 (Plates 1-17).
CHAPTER 11
TH E SPINAL NERVES
The spinal nerves (nervi spinales) (Figs. 9-1, 9- the last several lumbar, the sacral, and the coc
2 ) usually number thirty-six pairs in the dog. cygeal nerves have to pass increasingly longer
They have retained their embryological seg distances before they reach the correspond
mental characteristics, although the spinal cord ing intervertebral foramina. Therefore lumbar,
from which they arise and the structures which sacral, and coccygeal nerves leave the caudal
they supply have largely lost this feature. Each part of the spinal cord and lie within the dural
spinal nerve is attached to its corresponding and arachnoid membranes. Since the caudal
segment of the spinal cord by the dorsal and part of the spinal cord and the nerves which
ventral rootlets or root filam ents (fila radicularia). leave it resembles a horse’s tail, it is called the
The dorsal filaments carry incoming or afferent cauda equina.
impulses; the ventral filaments carry outgoing The spinal ganglia (ganglia spinale) are ag
or efferent impulses. The afferent filaments col gregations of unipolar nerve cells which are lo
lectively are called the dorsal root (radix dor cated in the dorsal root within (rarely external)
salis), and the efferent filaments collectively are the corresponding intervertebral foramen. The
called the ventral root (radix ventralis) of the axons of the unipolar cells divide into central and
spinal nerve. The afferent filaments enter the peripheral processes. The central processes form
dorsolateral sulcus of the spinal cord. The effer the dorsal root filaments, whereas the peripheral
ent rootlets leave the ventrolateral funiculus processes intermingle with the axons of the ven
over an area which is about 2 mm. wide. Neither tral root filaments in forming the mixed (sensory
the dorsal nor the ventral root filaments are com and motor) spinal nerve.
pact units. They consist of loosely united bundles
of nerve fibers which are difficult to differentiate Branches of a Typical Spinal Nerve
from each other because of the transparency of
the covering arachnoid. The number of dorsal Just peripheral to the intervertebral foramen
root filaments agrees closely with the number of each spinal nerve trifurcates into dorsal, ventral,
ventral root filaments for each spinal nerve. The and visceral branches (Fig. 11-1).
number of dorsal and ventral root filaments aver The dorsal branch (ramus dorsalis) extends
ages six each for the first five cervical nerves. dorsad and usually subsequently divides into
They increase in size and in number to an aver medial and lateral parts.
age of seven dorsal and seven ventral filaments The ventral branch (ramus ventralis) supplies
from the fifth cervical segment as far caudad as all hypaxial structures, including the limbs. It
the second thoracic segment. From the second divides into medial and lateral parts except
thoracic segment through the thirteenth thoracic where it is specialized to supply the extremities
segments there are two dorsal and two ventral or the tail.
filaments which form each thoracic nerve root. The visceral branch (ramus visceralis or ramus
The filaments which merge to compose the communicans) differs from the dorsal and ven
nerves of the lumbosacral plexus are large and tral branches in that it carries only motor and
constantly double. There are usually two dorsal sensory fibers to and from visceral structures
and two ventral root filaments for each of the (gland tissue and smooth muscle). Some visceral
three pairs of sacral and five pairs of coccygeal branches also contain fibers which come from
spinal nerves. and go to the heart. Consult Chapter 12 for
Because the vertebral column continues to further information. The visceral branch is con
grow after the spinal cord has ceased growing, nected to the ventral branch of the spinal nerve.
572
C e r v ic a l N erves 573
-D o rsal root
D o rs o l b ra n c h - - - - D o rsa l ro o t g a n g lio n
Ve ntra l branch - V e n tra l r o o t
V is c e r a l bronch ----
S y m p a th e tic t r u n k -----
C u ta n e o u s b ra n c h e s
^ i Ansa c e r v i c a l i s
T r a n s v e r s e c e r v i c a l n.
Long
t h o r a c i c n.
— To m. b r a c h i o c e p h a l i c u s
- - S u p r a s c a p u la r n.
Phreni
- S u b s c a p u l a r n.
I n t e r c o s to
b ra c h ia l „ ■v" " M u s c u l o c u t a n e o u s n.
■v
" 'T o m. p e c t o r a l is s u p e rf.
2nd. i n t e r s.
N (
co sta I 'v7o m. c o r a c o b r a c h i a l i s
' A x i l l a r y n.
1st. in t e r c o s ta l
NR a d i a l n.
L at. t h o r a c i c n
To m. p e c to r a l i s p ro f. • ' Me d i a n n.
i
1U l n a r n
T h o r a c o d o r s a I n.1
F k ;. 11 -2. Schema of the cervical nerves and brachial plexus.
5 74 Chapter 11. S p in a l N e rv e s
The spinal nerves usually leave the vertebral part of the large obliquus capitis caudalis mus
canal by means of spaces between adjacent cle. It arborizes in the muscles of the cranial
vertebrae, the intervertebral foram ina. Certain portion of the neck. These include the obliquus
discrepancies exist because the number of verte capitis cranialis, obliquus capitis caudalis, rectus
brae and the number of spinal nerves for each of capitis dorsalis, intermedius, and minor, and the
the several regions are not always the same. cranial ends of the semispinalis capitis and
There are eight pairs of cervical nerves, but only splenius.
seven cervical vertebrae. In most dogs there are The ventral branch o f the first cervical nerve
twenty coccygeal vertebrae, but only the first (ramus ventralis n. cervicalis I) initially lies in
five pairs of coccygeal nerves usually develop. the osseous groove of the atlas which runs trans
Since the three sacral vertebrae are fused to versely to the alar notch from the intervertebral
form the sacrum, there are two dorsal and two foramen. The ventral branch passes through the
ventral pairs of sacral foramina for the passage alar notch and continues in a ventrocaudal di
of the dorsal and ventral branches of the first two rection by passing between the rectus capitis
pairs of sacral nerves. The third pair of sacral lateralis and the rectus capitis ventralis muscles.
nerves pass through the intervertebral foramina It is initially covered by the medial retropharyn
located between the sacrum and the first coc geal lymph node. After appearing at the caudal
cygeal vertebra. border of this node it continues its course down
the neck in close relation to the vagosympathetic
CERVICAL NERVES nerve trunk. It usually anastomoses with the
smaller descending branch of the hypoglossal
There are eight pairs of cervical nerves (nn. nerve to form the cervical loop (ansa cervicalis).
cervicales) (Fig. 11-2), although there are only Variations in the formation of the cervical loop
seven cervical vertebrae. The intervertebral are common. In about 3 per cent of dogs it fails
foramina, through which the first cervical nerves to develop. It may be a long loop measuring 10
pass, are located not between the skull and the cm. Usually the cervical loop extends to a level
atlas, but in the craniodorsal portion of the arch through the third cervical vertebra, but in some
of the atlas. The second pair of cervical nerves dogs it is short, lying on the carotid sheath about
leave the spinal canal through the second pair of 2 cm. caudal to the jugular process. Several
intervertebral foramina, which are located be branches arise from the cervical loop. As the
tween the atlas and the axis. The last or eighth sternothyroid and sternohyoid muscles cross the
pair of cervical nerves pass through the eighth larynx they each receive a branch from the loop.
intervertebral foramina, which are located be Another branch runs caudad on the trachea and
tween the seventh cervical and first thoracic bifurcates near the middle of the neck. The
vertebrae. Therefore the cervical nerves leave shorter of these branches becomes related to the
the vertebral canal cranial to the vertebra of the middle of the lateral border of the sternothy-
same number with the exception of the first and roideus before entering its distal portion. The
last pairs. longer branch follows the lateral border of the
The first cervical nerve (n. cervicalis I) arises sternohyoideus caudally and enters the muscle
from the first segment of the spinal cord, which approximately 4 cm. cranial to the manubrium
is located just caudal to the foramen magnum, of the sternum.
and is surrounded by the cranial portion of the The second cervical nerve (n. cervicalis II)
atlas. Both its dorsal and ventral formative root differs from other typical spinal nerves in three
filaments number from three to five and are respects: (1) Its afferent component is larger
approximately equal in size. In most specimens than that of any of the other cervical or thoracic
a barely distinguishable dorsal root ganglion is nerves. (2) The dorsal and ventral roots fuse
present, while in others it may be 1 mm. in di peripheral to the second intervertebral foramen.
ameter. Upon emerging through the interverte (3) The large dorsal root ganglion lies completely
bral foramen of the atlas the first cervical nerve outside the vertebral canal.
divides into dorsal and ventral branches of equal The dorsal branch o f the second cervical nerve
size, measuring about 2 mm. in diameter. (ramus dorsalis n. cervicalis II) or the greater
The dorsal branch o f the first cervical nerve occipital nerve (n. occipitalis major) is approxi
(ramus dorsalis n. cervicalis I) or the suboccipital mately the same size as the ventral branch (3
nerve (n. suboccipitalis) does not divide into mm. in diameter) in a large dog. It runs caudo-
medial and lateral branches, and no part of it dorsally where it is deeply located under the
supplies the skin. It lies initially under the cranial obliquus capitis caudalis muscle. Emerging be
C e r v ic a l N erves 575
tween this muscle and the spine of the axis, it ramus dorsalis of cervical nerve III supply the
sends muscular branches into the semispinalis middle portion of the complexus muscle. The
capitis and the splenius muscles. It then turns main lateral branches of the rami dorsales of cer
cranially, perforates the overlying muscles, and vical nerves IV and V innervate the middle por
supplies the skin which covers the caudal portion tion of the biventer muscle. The splenius muscle
of the temporal muscle and the base of the pinna. is innervated by branches which come from the
The ventral branch o f the second cervical rami dorsales of cervical nerves III and IV which
nerve (ramus ventralis n. cervicalis II) runs leave near their origins. The branches enter the
caudoventrad on the lateral surface of the cleido- deep surface of the middle and caudal part of
mastoideus muscle for one or two centimeters the muscle. The cranial part of the splenius mus
and divides into the transverse cervical and great cle is supplied by large muscular branches from
auricular nerves. the dorsal branch of cervical nerve II.
The transverse cervical nerve (n. transversus The m edial branches (rami mediales) of the
colli) runs cranioventrad from under the platys rami dorsales of cervical nerves III through VII
ma and crosses the external and internal maxil perforate the intertransversarius dorsalis muscle
lary veins just before they unite to form the and run almost directly dorsad. In their dorsal
external jugular vein. The nerve may branch courses they lie between the multifidus cervicis
before crossing these veins. The branches of this and the spinalis cervicis muscles located medi
nerve arborize in the skin of the mandibular ally, and the complexus and biventer muscles
space. (the two portions of the semispinalis capitis)
The great auricular nerve (n. auricularis mag- located laterally. Finally they cross the lateral
nus) is the larger of the two terminal branches side of the ligamentum nuchae to reach the skin.
of the ventral branch of the second cervical In most specimens the third and fourth medial
nerve. It runs dorsocranially to the base of the dorsal cervical branches further divide into
pinna of the ear and divides into at least two cranial and caudal parts. They thus appear by
branches which run toward the apex of the ear. their spacing in the subcutaneous fascia as if they
Each of these nerves runs about midway be were branches of separate cervical nerves. Bi
tween the intermediate auricular artery and the laterally, these nerves innervate the loose, thick
peripheral arteries—the lateral and the medial skin of the dorsal and adjacent sides of the neck.
auricular arteries as they arborize in their courses The ventral branches (rami ventrales) of
toward the apex of the ear. Variations in both cervical nerves II through V (Fig. 11-4) pass
the numbers and the distribution of the arteries between the muscle bundles of the intertrans
and nerves to the pinna are common. versarius cervicis to reach the medial surface of
The dorsal branches o f cervical nerves III the omotransversarius muscle. They usually do
through VII (rami dorsales nn. cervicales III— not anastomose with each other; therefore only
VII) (Fig. 11-3) vary in both their distributions rarely is a cervical plexus (plexus cervicalis)
and form. Each of these branches sends a small formed. The ventral branches of cervical nerves
branch medially into the multifidus cervicis II, III, and IV regularly anastomose with the
muscle. Only their peripheral portions definitely accessory nerve, however, and a connection be
divide into medial (cutaneous) and lateral (mus tween cervical nerves II and III is not uncom
cular) branches. The dorsal branches of cervical mon. The ventral branch of the large, second
nerves III through VII gradually decrease in cervical nerve has been described. The ventral
size caudally. The seventh dorsal cervical branch branches of the third and fourth cervical nerves
may be reduced to a muscular twig which inner divide into large lateral (cutaneous) branches
vates only the deep muscle fibers which lie (rami laterales) which supply the skin of the
adjacent to it. The dorsal branch of the eighth ventrolateral part of the neck, and smaller m edial
cervical nerve may be absent. The dorsal (muscular) branches (rami mediales) which sup
branches of the middle cervical nerves perforate ply the longus capitis, longus colli, intertrans
the lateral portion of the multifidus cervicis mus versarius cervicis, omotransversarius, and
cle and run dorsally with a slight caudal inclina brachiocephalicus muscles. The medial branches
tion. Upon reaching the ventral portion of the appear as loose clusters of nerves which arise
biventer muscle they usually bifurcate into just peripheral to the intervertebral foramina
medial and lateral branches. The third dorsal and, after short caudoventral courses, enter the
branch divides near its origin, and the fourth several muscles. The size of the ventral branches
dorsal branch may also divide deeply. of the second to the fifth cervical nerves progres
The lateral branches (rami laterales) of the sively decreases. The ventral branch of the
576 Chapter 11. S p in a l N e rv e s
M. m u l t i fid u s —
Cutaneous br. of C5~ -C5, dorsal br
r 'IN iW iO H
F k .. 11-3. Dorsal branches of the cervical nerves, dorsal aspec t. (The muscles on the right side are reflected.)
C e r v ic a l N erves 577
M .s te rn o c e p h o lic u s ( re fle c te d )
R e t r o a u r i c u l o r n. (VII)
A cc e s s a ry
M. o m o t r a n s v e r s a r i u s
M. c l e id o c e r v ic a lis
Pia t y s m a t
M. tr a p e z iu s
Parotid g la nd
- P a r o t i d duct
\ \C e rv ic a l br: of VII
\ xMandibular gland
\ NBr. to mm. cleidomastoideus c
stern oce p ha licu s
\C2, v e n tr a l br.
' G reat a u r ic u la r n.
''Transverse c e r v ic a l n
M. depressor a u r ic u la e
C3, ventral br.
'M. cleidomastoideus
Ext. j u g u l a r v.
second cervical nerve is about three times larger formed by the ventral branches of the sixth,
than that of the fifth. seventh, and eighth cervical and the first and
second thoracic spinal nerves. Occasionally the
NERVES TO THE DIAPHRAGM ventral branch of the fifth cervical nerve also
contributes to its formation; frequently the sec
The phrenic nerve (n. phrenicus) (Fig. 11-2) ond thoracic contribution is lacking. When
reflects the fact that the diaphragm which it either or both the fifth cervical and the second
supplies has a cervical origin. The phrenic nerve thoracic spinal nerves send branches which
regularly arises from the fifth, sixth, and seventh enter into the formation of the brachial plexus,
cervical nerves, and occasionally a small twig they are exceedingly small compared with the
comes from the fourth. other ventral branches which compose the
The branches of origin of the phrenic nerve plexus. In over 250 dissections neither one of
run caudally, medial to the brachial plexus. these nerves was found to be over 1 mm. in di
While running in the fascia adjacent to the ex ameter. Allam et al. (1952) found in fifty-eight
ternal jugular vein, the nerve branches converge dissections that the fifth cervical and the second
and unite to form the phrenic nerve just cranial thoracic nerve contributions to the brachial
to the thoracic inlet. The nerve on each side plexus are more often absent than present.
then passes through the thoracic inlet ventral to Ventral branches of the cervical (C) and tho
the subclavian artery and dorsal to the omo racic (T) spinal nerves which form the brachial
cervical artery. At this site it is joined by a fine plexus are distributed as follows, according to
branch from the caudal cervical ganglion or the Allam et al. (1952):
sympathetic trunk adjacent to the ganglion.
Within the thorax the right phrenic nerve lies in 58.62% formed by C 6, 7, 8, and T 1
20.69% formed by C 5, 6, 7, 8, and T 1
a narrow plica of pleura from the right lamina of 17.24% formed by C 6, 7, 8, and T 1 and 2
the precardial and cardiac mediastinum and the 3.4% formed by C 5, 6, 7, 8, and T 1 and 2
plica venae cavae. The left phrenic nerve lies in
a similar plica from the left pleural sheet of the After the ventral branches of the last three
mediastinum. Each phrenic nerve spreads out cervical and the first and second thoracic spinal
on its respective half of the diaphragm, where it nerves have passed through the intervertebral
supplies this muscle with motor and sensory foramina and the intertransverse musculature,
fibers. Upon reaching the diaphragm each they cross the ventral border of the scalenus
phrenic nerve divides into three main branches: muscle and extend to the thoracic limb by tra
ventral, lateral, and dorsal. This splitting takes versing the axillary space. In this course parts of
place lateral to the middle portion of the tendi these nerves unite with each other to form the
nous center. Each nerve division supplies its various specific nerves which supply the struc
appropriate third of its half of the diaphragm. tures of the thoracic limb and adjacent muscles
Each dorsal branch therefore supplies the crus and skin. The axillary artery and vein lie ventro
of its side. The dog, like man, usually has a con medial to the caudal portion of the brachial
nection between each phrenic nerve and the plexus. The external jugular vein, after it has
sympathetic system at the celiac plexus. In 300 been augmented by the proximal tributary of
dissections we have not seen an accessory the cephalic vein, crosses the ventral surfaces of
phrenic nerve in the dog. The periphery of the the seventh and eighth cervical nerves, from
diaphragm also receives sensory fibers from the which it is separated by the omocervical artery.
last several intercostal nerves (Lemon 1928). It The axillary artery, after having crossed the cra
is generally agreed that the phrenic nerves are nial margin of the first rib, lies closely applied to
the only motor nerves to the diaphragm. In the ventral margin of the scalenus primae costae
vestigators also agree that bilateral phrenicot- muscle and later follows along the craniomedial
omy does not seriously interfere with respiration margin of the radial nerve as both the artery and
in the dog even under moderate exercise. nerve run distad in the brachium. They are
crossed ventrally at the first rib by a muscular
BRACHIAL PLEXUS nerve branch which goes to the deep pectoral
muscle.
The brachial plexus (plexus brachialis) (Figs. Allam et al. (1952) recognize three cords in
11-5, 11-6, 11-7) consists of the large somatic the brachial plexus of the dog to assist the ex
nerve plexus which gives origin to the nerves ploring surgeon by establishing suitable land
which supply the thoracic limb. It is usually marks for electrical stimulation. These lie as in-
(Text continued on page 582.)
B r a c h ia l P lexus 579
S u p r a s c a p u l a r n.N T h o r a c o d o r s a l n.
x
S u b s c a p u l a r n.^ ^ x
To mm. t e r e s maj. v s u b s c a p .
A x i II a r y n.-
R a d i a l n.
To m. s u p r a s p i n a t u s —
To m . p e c t o r a l i s p r o f .
To m. p e c t o r a l i s s u p e r f -------
- L a t t h o r a c i c n.
To m. b r a c h i o c e p h a l i c u s "
To mm. t e n s o r f a s c i a e a n t e b r . v
M u s c u l o c u t a n e o u s n." tric e p s , ca pu t longum
To m. c o r a c o b r a c h i a l i s - To m. t r i c e p s , c a p u t med.
To m. t e r e s m i n o r ' - To m . t r i c e p s , c a p u t ac c e s s .
To m. d e l t o i d e u s / " ^ M e d i a n n.
To m. b i c e p s b r a c h i i 7 U l n a r n.
L a t . c u t a n . b r a c h i a l n. Caud. c u t a n e o u s
a n t e b r a c h i a l n.
To mm. a n c o n e u s v t r i c e p s , 7
c a p u t l at . f
" To m. brae h i a Us
S u p e r f . r a d i a l n . , l a t . br. Med. c u t a n e o u s a n t e b r a c h i a l n.
Tom. p e c t o r a l i s p i o f
To p h r e n i c n.N ,-M s u b s c a p u la r is
Sub s c a p u la r n v -A x i l l a r y n
S u p ra s c a p u la r n „ 'T o m. p e c t o r a l i s p r o f
^ s To mm. s u b s c a p u la ris *■ te re s muj.
To m. b r a c h l o c e p h a l ic u s
- R a d ia l n
To m. p e c t o r a 11s s u p e r f . -
M te re s m a jo r
M. s u p r a s p i n a t u s - ~ _ Thoracodorsal n.
M u s c u lo c u ta n e o u s n . ~ ---- - Median v u ln a r nn
Tom c o r a c o b r a c h i a l i s - - ~ — - Lat. th o r a c i c n
Ax i I l a r y a. -----
~M latissimus dorsi
M, c o r a c o b r a c h i a l is - -
ro m .triceps, long heaJ
C ra n .c irc u m fle x h u m e ra l a.- ~ s' tensor fasc antebr
B r a c h i a l a.
M. tensor fasc antebr
To m. b i c e p s b r a c h i i -
U ln a r n
M. t r i c e p s , acc. h e a d '
M e d ia n n
D e e p b r a c h i a l a.
■ M. tn c e p s , long head
C o lla te ra l u l n a r a.
A n a s t o m o t ic br.
P r o * , c o l la te r o I r a d i a l a.
A r t i c u l a r br
S u p e r f r a d i a l n., m e d i a l br. '
M o m o t r a n s v e r s a r i us C4
/
M. b r a c h i o c e p h a l i c u s . 1 / J u g u l a r v.
* I
Subscapular
/
/ M- l onqus c a p i t i s
S u p r a s c a p u l a r n.\ *
A x i I I a r y n .(
M.supraspinatusv ' M. s c a l e n u s
M u s c u l o c u t a n e o u s n.\ \ ,C6
R a d i a l n., ' \
1 I
M.coracobrachialis ( ' x Omocervi cal a.
C r a n . c i r c u m f l . h u m e r a l a.y
Deep b r a c h i a l a Esophagus
B r a c h i a l a-\ ce
M e d i a n n.s
TI
U ln ar
-Phrenic n
M. biceps v
brachii - F irst rib
' ■ A x i l l a r y a.
M. p e c t o r a l i s superf.
■s L a t th o ra c ic n.va.
M.
coput med' / M. p e c t o r a l i s p r o f .
v caput* longurr
, ' ,I
M. t e n s o r f a s c i a e 1
antebrach ii
M e d i a n r u l n a r nn.1 M subscapularis
/ /
Caud c i r c u m f l e x , h u m e r a l a. . i ' M.teres m a j o r
S u b s c a p u l a r a.I v4th i n t e r c o s t a l n.
M. l a t i s s i m u s d o r s i 3 r d i n t e r c o s t a l n.
T h o r a c o d o r s a l nJ 2 n d i n t e r c o s t a l n.
termediate nerve trunks between the ventral cles of the shoulder as it curves around the cau
branches of the spinal nerves which form the dal border of the subscapularis muscle near its
plexus and the named nerves which innervate distal end. In its intermuscular course proximo-
structures of the limb. Most of these vary. For caudal to the shoulder joint it divides basically
further information on the morphology of the into two portions; one part sends twigs to the
brachial plexus of the dog refer to Russell (1893), subscapularis muscle and completely supplies
Reimers (1925), and Bowne (1959). the teres major muscle. The other portion, ac
The nerves which are branches of the brachial companied by the caudal circumflex humeral
plexus or are direct continuations of the forma vessels, runs laterally to supply the laterally ly
tive ventral branches include the suprascapular, ing teres minor and deltoideus muscles. Before
subscapular, axillary, musculocutaneous, radial, entering the teres minor muscle a branch enters
median, ulnar, dorsal thoracic, lateral thoracic, the caudal part of the shoulder joint capsule
long thoracic, pectoral, and muscular branches. (Fig. 11-8).
The basic plan of the brachial plexus appears The lateral cutaneous brachial nerve (n. cu
as a variable “anastomosis” of the last three cer taneus brachii lateralis) leaves the axillary nerve
vical and first two thoracic spinal nerves whose just prior to the entry of this nerve into the del
fibers run in common for short distances and toideus muscle. It arises, therefore, lateral to the
then segregate in variable combinations to form space between the origins of the lateral and long
the extrinsic and intrinsic named nerves of the heads of the triceps muscle. It runs distally on
thoracic limb. the lateral head of the triceps muscle, where it is
The suprascapular nerve (n. suprascapularis) covered by the deltoideus muscle. It appears
arises primarily and occasionally entirely from subcutaneously caudal to the main portion of
the sixth cervical nerve. It often has a contribu the cephalic vein where it is associated with the
tion from the seventh, but rarely from the fifth cutaneous branches of the caudal circumflex
cervical nerve. It enters the distal end of the in- humeral artery and vein. It supplies the skin of
termuscular space between the supraspinatus the lateral surface of the brachium, lapping in its
and the subscapularis muscles from the medial distribution the area supplied by the intercosto-
side. It is accompanied by the supraspinous ar brachial nerves caudally and the cutaneous
tery and vein. The suprascapular nerve is pri branches of the cervical nerves cranially. It ter
marily a motor nerve to the supraspinatus and minates in the skin of the proximodorsolateral
infraspinatus muscles. Prior to crossing the dis aspect of the forearm. At the elbow joint or just
tal end of the spine of the scapula the nerve sends distal to it, it joins the medial branch of the super
a delicate twig to the lateral part of the shoulder ficial radial nerve, and by means of this nerve its
joint (Fig. 11-8). fibers are carried to the skin of the dorsum of the
The subscapular nerve (n. subscapularis) is antebrachium and possibly the dorsum of the
usually a single, but occasionally double, nerve paw also.
which arises from the union of a branch from the The musculocutaneous nerve (n. musculocu-
sixth and seventh cervical nerves, or if the nerve taneus) gives muscular branches to the coraco-
is double, one part usually arises from the sev brachialis, biceps brachii, and brachialis muscles.
enth cervical nerve directly. It usually divides It continues in the forearm as the medial cuta
into cranial and caudal parts upon entering the neous antebrachial nerve. The musculocutane
medial surface of the distal fifth of the subscapu ous nerve arises mainly from the seventh cervical
laris muscle. The subscapular nerve is about 5 nerve. It is irregular in its formation, occa
cm. long in a medium-sized dog. This permits sionally receiving a branch from the sixth cervi
the extensive sliding movement of the shoulder cal, but more frequently from the eighth cervical
on the thorax during locomotion without nerve and even from the first thoracic nerve in rare
injury. instances. Throughout its course in the brachium
The axillary nerve (n. axillaris), like the sub it lies between or under the cranially lying biceps
scapular nerve, is much longer than the distance brachii muscle and the axillary vessels caudally.
between its origin and its peripheral fixed end. The muscular branches are three in number.
It arises as a branch from the combined seventh Proximally a small twig goes to the coracobrachi-
and eighth cervical nerves. A contribution from alis muscle. This branch is small, and instead of
the sixth cervical nerve may also be present. It arising from the musculocutaneous nerve di
may arise completely or nearly completely from rectly it may exist as a separate twig which
either the seventh or the eighth cervical nerve comes from the eighth cervical or first thoracic
(Allam et al. 1952). It supplies mainly the mus nerve, or both. In reaching the coracobrachialis
B r a c h ia l P l e x u s 583
it follows the cranial circumflex humeral vessels to the accessory and medial heads before it
over a portion of its course. A large branch, the makes contact with the brachial muscle. It fol
proximal muscular branch (ramus muscularis lows this muscle, where it lies related to the nu
proximalis), enters the deep surface of the biceps trient artery of the humerus, in a spiral manner
brachii muscle about 4 cm. from its origin and around the humerus. Upon contacting the lateral
near its caudomedial border. In the distal third head of the triceps it sends a branch to it, and
of the brachium an anastomotic branch (ramus shortly thereafter it bifurcates into deep and
anastomaticus) passes distocaudad usually me superficial branches. The deep branch runs un
dial to the brachial vessels and joins the median der the proximocranial border of the extensor
nerve, which with the ulnar nerve lies caudal to carpi radialis muscle. At the place of bifurcation
the brachial vessels. As the musculocutaneous of the radial nerve a minute twig runs to the deep
nerve winds under the terminal part of the biceps surface of the brachioradialis muscle. The super
brachii muscle from the medial side, it termi ficial branch pursues a more cranial course and
nates in the distal muscular branch (ramus mus becomes superficial between the distocranial
cularis distalis), which enters the distal medial border of the lateral head of the triceps and the
portion of the brachialis muscle, and the small lateral surface of the deeply lying brachial mus
medial cutaneous antebrachial nerve (n. cuta- cle.
neus antebrachii medialis). This cutaneous The deep branch (ramus profundus) of the
branch crosses the lateral side of the tendon of antebrachial part of the radial nerve at first
the biceps brachii muscle and enters the cranial passes under the extensor carpi radialis muscle
surface of the antebrachium from the flexor an near its origin from the lateral supracondyloid
gle of the elbow joint. As the nerve crosses the crest and sends a branch into it. As the deep
cranial surface of the elbow joint it sends a small branch crosses the flexor surface of the elbow
branch to the craniolateral part of it. It freely joint it sends an articular branch to the cranio
branches in its course distad in the forearm as it lateral part of it. The remaining part of the deep
supplies the skin of the craniomedial portion of branch then passes under the supinator muscle
the antebrachium. It ends at the carpus. The which it supplies. Upon emerging from un
cutaneous area which it supplies is overlapped der this muscle it immediately divides into
by the area supplied by the cutaneous branches branches which supply the common and lateral
of the medial branch of the superficial radial digital extensors and a small branch which
nerve cranially and the caudal cutaneous ante closely follows the lateral border of the radius
brachial nerve from the ulnar caudally. and runs distad to innervate the abductor pol
The radial nerve (n. radialis) (Figs. 11-9, 11- licis longus and extensor pollicis longus et indicis
10) arises from the seventh and eighth cervical proprius muscles.
and the first and second thoracic nerves. It is the The superficial branch (ramus superficialis) of
largest nerve of the brachial plexus. It supplies the radial nerve is its more cranial branch. Upon
all the extensor muscles of the elbow, carpal, and emerging from under the cranial part of the dis
digital joints and also the supinator, the brachio- tal border of the lateral head of the triceps mus
radialis, and the abductor pollicis longus muscles. cle it runs obliquely craniodistad on the brachial
The skin on the cranial portion of the antebra muscle, where it is covered by the heavy inter-
chium and paw is also supplied by fibers of radial muscular fascia. After running about 1 cm. in
nerve origin. As the radial nerve approaches the this location it perforates the heavy fascia and
brachium by traversing the axillary space it lies divides unevenly into a larger lateral branch
lateral to the axillary vein and medial to the axil (ramus lateralis) and a smaller m edial branch
lary artery. Upon crossing the medial surface of (ramus medialis). These branches continue to
the conjoined tendons of the teres major and the carpus in relation to the lateral and medial
latissimus dorsi muscles it lies caudal to the branches of the proximal collateral radial arter
brachial vessels which are the continuation of ies respectively. They thus closely flank the
the axillary vessels after these have crossed the medial and lateral sides of the cephalic vein as
conjoined tendon. Upon entering the interval they traverse the antebrachium. From the lateral
between the medial and long heads of the triceps branch of the superficial portion of the radial
muscle the radial nerve divides into a branch nerve the usually double lateral cutaneous ante
which runs proximolaterad and is distributed to brachial nerve (n. cutaneus antebrachii later
the long head of the triceps. The second branch alis) arises. Since the nerves to the skin of the
runs distolaterad and represents the main con lateral side of the antebrachium are appreciably
tinuation of the radial nerve. It supplies a branch larger and longer than those which supply the
584 Chapter 11. S p in a l N e r v e s
^ Scapula
^ A x i l l a r y a v n.
to m. s u b s c a p u l a r i s v
m te re s m a jo r
^ ^ S u b s c a p u la ra .
J o i n t c a p s u le - ^ N .to m. s u b s c a p u l a r i s v
j o i n t c a p s u le
T r a n s , h u m e r a l h q ---- - - T h o r a c o d o r s a l a.
" •* S u b s c a p u l a r a.
Tendon of -
m. b i c e p s b r a c h i i " Br. to j o i n t c a p s u l e
" Caud c i r c u m f l e x h u m e r a l a.
v " >. A x 1 1l a r u a
H um erus
''>.Cran c i r c u m f l e x h u m e r a l a.
C o lla t e r a l r a d i a l a.
P).W£«&V>«
'■ A to m. b i c e p s b r a c h i i
S p ' ne o f s c a p u l a _ ^ -S u p ra s c a p u la r n v a
- N to m s u p r a s p i n a t u s
S u b s c a p u la r a
_ Br. to j o i n t c a p s u l e
N. to m. i n f r a s p i n a t u s —
A
A x i l l a r y a-v n ------------ ♦ - \ - M a jo r tu b e rc le
T h o ra c o d o rs a l a -
S u b s c a p u la r a - -
C ra n c i r c u m f le x h u m e r a l a ~ " N. to m. t e r e s m i n o r v
j/ j o i n t c a p s u le
B r to j o i n t c a p s u l e ' '
C aud-ci r c u m f I e x h u m e r a l a ' ~ — N. to m d e l t o i d e u s
/
N to m d e l to i d e u s ' i " - Lat. c u t a n e o u s b r a c h i a l n,
A x i 11 a r y a ' ‘VjL H um erus
C o lla te ra l r a d ia l a
Fic. 11-8. A. Nerves and arteries of the right shoulder joint, medial aspect.
B. Nerves and arteries of the right shoulder joint, lateral aspect.
B r a c h ia l P lexus 585
d e lto id e u s
M .tr ic e p s , caput laterale
M. tric e p s , caput langum _ _ „ W tricep s, caput accessOrium
In te r c a s ta b r a c h ia l n - _ _ - L a t cutaneous b ra chia l n.
■M. brachiocephalicus
Collateral radial a.-.
M. b ra c h ia l is
W. tric e p s , caput mediate R a d ial n.
Deep ramus
M. b r a c h i o r a d i a li s —
Superf. ramus, med. branch
Superf. ramus, lat. branch
Col l a t e r a l u ln a r a — " M. s u p in a to r
D o rs a l interosseous a ' 'To m.ext. d ig ito ru m communis
Caud. cutan.antebrachial n.- ' ^ Tom. ext. dig itoru m lat. (cut)
M. flex, c a rp i u ln a r is , caput ulnare ' To mm. abd. p o l l ic i s longus v
ext. p o l l ic i s longus et in d ic is proprius
M. ext. c a r p i u l n a r i s
U l n a r n.
'-M. abd. p o llic is longus
M.ext. ca rp i r a d ia lis
M.flex. carpi u ln a r , caput hum erale
M. ext d ig ito ru m communis
M.ext p o l l i c i s longus et in d ic is proprius
P a lm a r b r . '
D a r s a l br.
M.ext. d i g i t o r u m l a t .'
1.HEWSO*4
B r o f p a l m a r in te r o s s e o u s a
skin on the dorsum of the antebrachium, it seems dal to the brachial artery and lateral to the bra
feasible to designate them by distinctive names. chial vein. The median nerve is cranial in position
The more proximal nerve is the larger and is the to the larger ulnar nerve. At the flexor surface
branch which is more constantly present. It of the elbow joint the median nerve dips later
arises just distal to the flexor surface of the elbow ally under the pronator teres muscle and enters
joint and, associated with relatively large cuta the large caudal group of flexor muscles of the
neous branches of the lateral branch of the prox antebrachium. It supplies the pronator teres,
imal collateral radial vessels, it supplies the skin pronator quadratus, flexor carpi radialis, flexor
of the proximal half to two-thirds of the lateral digitorum superficialis, and the radial head of the
surface of the antebrachium. The more distally flexor digitorum profundus muscles. It also sends
located nerve to the skin of the lateral side of the fibers to the ulnar and humeral heads of the
antebrachium, smaller than the more proximally flexor digitorum profundus muscle, and a small
located nerve, also is accompanied by a cutane articular branch to the medial aspect of the el
ous artery and vein which serve the cutaneous bow joint.
area of the region. Occasionally more than two Upon emerging from under the pronator teres,
lateral cutaneous antebrachial nerves are pres to which it sends a small branch, several muscu
ent. Small branches leave both the medial and lar branches (rami musculares) leave the caudal
lateral branches of the superficial radial nerves portion of the nerve. The shortest and most
and innervate the skin on the cranial surface of proximal of these nerves enters the flexor carpi
the antebrachium as the cranial cutaneous ante radialis muscle close to its humeral origin. The
brachial branches (rami cutanei antebrachiales remaining flattened bundle of muscular
craniales). The medial and lateral branches of branches crosses the medial surface of the me
the superficial radial nerves as they innervate dian (brachial) vessels at the place where the
the dorsum of the forepaw are described under common interosseous artery arises and, after run
the description of the nerves to the forepaw. ning under the flexor carpi radialis and through
Because it supplies all the extensor muscles of the humeral head of the flexor digitorum profun
the thoracic limb, except those of the shoulder, dus, most of them end in the superficially lying,
injury to the proximal part of the nerve results flattened flexor digitorum superficialis muscle. In
in a grave syndrome. Uncomplicated radial pa their path to this muscle they lie about 1 cm.
ralysis is rare. Surely what was diagnosed as proximal and parallel to the palmar antebrachial
radial paralysis a generation ago was probably vessels. In this deep location a branch is sent to
an avulsion of the brachial plexus in most cases. the radial head of the flexor digitorum profundus
Clifford et al. (1958) describe such a case. Bowne muscle, which it completely innervates, and
(1959) has demonstrated that the radial nerve smaller twigs enter the humeral and ulnar heads
can be interrupted just distal to the point where of this muscle, which are also supplied by the
the last branch goes to the triceps brachii with ulnar nerve. The small interosseous nerve of the
no permanent impairment of locomotion. He antebrachium (n. interosseus antebrachii ante
observed, however, that his experimentally in rior) first runs on the proximal part of the deli
duced radial paralysis did not produce the same cate interosseous membrane. It then perforates
syndrome as that seen in typical clinical cases. this membrane and runs distally on about the
Worthman (1957) has reported upon many neu proximal half of the pronator quadratus muscle,
rectomies of the nerves of both the fore and hind where it appears as a fine white streak. It enters
limbs in the dog. this muscle in its distal half and innervates it.
The m edian nerve (n. medianus) (Figs. 11-10, The portion of the median nerve which con
11 - 11 ) arises primarily from the eighth cervical tinues distad in the antebrachium, after the mus
and the first and second thoracic spinal nerves. cular branches have arisen, is at first related to
Reimers (1925) does not regard the nerve to be the median artery and vein. When the median
formed until it has received the anastomotic (brachial) artery terminates by dividing into the
branch from the musculocutaneous nerve in the radial and ulnar arteries at about the middle of
distal part of the brachium. Through this anasto the antebrachium, the median nerve continues
mosis the median nerve is augmented by fibers distad in relation to the larger ulnar artery. This
from the sixth and seventh cervical nerves. The portion of the median nerve is small, measuring
median nerve is about twice as large as the ulnar about 0.5 mm. in diameter.
nerve, with which it is loosely bound throughout The ulnar nerve (n. ulnaris) (Figs. 11-10, 11-
the proximal two-thirds of the brachium. The 12 ) arises in close association with the radial and
loosely united median and ulnar nerves lie cau median nerves from the eighth cervical and the
B r a c h ia l Plexus 587
first and second thoracic nerves. After leaving into its lateral surface. Throughout the middle
the caudal part of the brachial plexus the median third of the antebrachium the ulnar nerve lies
and ulnar nerves are flanked by the brachial ar on the caudal border of the deep digital flexor,
tery cranially and the brachial vein caudally. where it is covered by the humeral head of the
They are bound to each other by areolar tissue flexor carpi ulnaris muscle. At about the middle
until they reach the middle of the brachium, of the antebrachium the small, cutaneous dorsal
where they diverge. The ulnar nerve, which branch of the ulnar nerve arises. This branch and
measures about 3 mm. in diameter, runs distad the palmar branch arise as terminal branches of
along the cranial border of the medial head of the ulnar nerve. Both these branches are dis
the triceps brachii muscle and adjacent to the tributed to the structures of the forepaw.
caudal border of the biceps brachii. Upon enter
ing the caudomedial part of the antebrachium
the ulnar nerve runs under the heavy antebra N erves of th e F orepa w (M a n u s)
U In a r n v
C o lla te ra l u ln a r a
M e d i a n n.v >
M u s c u lo c u ta n e o u s n - - Caud c u ta n . a n te b r a c h i a I n ( u ln a r )
B ra c h ia l a - "
Prox c o l I a t e r a l r a d i a l a . - " ' Br. to j o i n t c a p s u l e
N to m b r a c h i a 1 1s - x - N tom. f l e x c a r p i u l n a r i s
(u ln o r h ea d)
Med. c u t a n a n t e b r a c h i a l n.
— J o i n t c a p s u le
D iS t c o lla t e r a l r a d ia l a .-' „
__ R e c u r r e n t u l n a r a
B r to j o i n t c a p s u l e " ,
✓ ■N to m- f l e x o r c a r p i r a d i a l i s
N to m p r o n a t o r t e r e s '
N - to r a d i a l c o l l a t e r a l l i g ■' /V. to mm. f l e x o r d i g i t o r u m p r o f
fle x o r d ig ito r u m su p e rf
M e d ia n a
Comm on t n t e ro s seous Prox d o r s a l i r t e r o s s e o u s a v
N to mm. p r o n a t o r q u a d r a t u s v ' in te ro s s e o u s n
f l e x o r d i q 11 p r o f ( r a d i a l h e a d ) A c c e s s o r y i n t e r o s s e o u s a.
M ed ia n n v P o lm a r i n te r o s s e o u s a
- j - C o l l a t e r a l r a d i a l a.
N u t r i e n t a of h u m e ru s- -
L _/R a d i a l n
■ - - Deep r a d i a l n.
A ------ S u p e r f r a d i a l n.
LljW___N t o m b r a c h i o r a d i a l i s
B M u s c u l o c u t a n e o u s n.
~~ ~ N. to m ext c a r p i r a d i a l i s
D isl c o lla t e ra l r a d ia l a.
- f i r to j o i n t c a p s u l e
U l n aJ rr Lc Uo Ml l Ua ft Ce frUa/l l# i/gcy -— - r — —V *® ?- ■
n.
' v N .to m ext c a r p i r a d i a l i s
B3r to j o i n t c a p s u l e - \ — 3 v ' B r to j o i n t c a p s u l e
P ro x .d a rs a l in te ro s s e o u s a — v N to m s u p i n a t o r
x N to m ext. d i g i t o r u m c o m m u n i s
B r to i n t e r o s s e o u s s p a c e —' a
N.fom ext. d i g i t o r u m lat
N t o m.ext. c a r p i u l n a r i s ' \ ✓ jiv
V II
N- to m .a b d p o l l i c i s l o n q u s ' N. to fn. e x t p e l l i a s l o n g u s et
i n d i c is p r o p r ius
Fn. 11—II). K \erves a.id arteries of the right elbow joint, medial aspect.
B Nerves and arteries of the right elbow joint, lateral aspect.
B r a c h ia l P lexus 589
M e d i a n n. s , U l n a r n.
M u s c u lo c u t a n e o u s n.~ _ _ ,/W. t e n s o r f a s c i a e a n t e b r a c h i i
M. b ic e p s b r a c h i i
..C a u d c u ta n e o u s a n te b ra c h ia l n.
Prox. c o l l a t e r a l r a d i a l a.
To m. b r a c h i a l i s - - C o lla te ra l u l n a r a.
M. b r a c h i a l is - ~
_ - M . p r o n a t o r te res
To m. fle x , d i g i t prof., c a p u t h u m e ra le
N|.NEUV)rt
Fic. 11-11. Nerves of the right antebrachium. media) aspect. Dissection showing median and musculocutaneous nerves.
590 Chapter 11. Sp i n a l N e r v e s
mediales et laterales II, III, et IV) and the m e the deep digital flexor muscle about 1 cm. lateral
dial dorsal digital nerve V (n. digitalis dorsalis to the origin of the ulnar artery. It converges to
medialis V). ward the artery distally in the antebrachium.
The m edian nerve o f the forepaw (n. medi- The vessel and nerve pass through the deep por
anus manus) (Fig. 11-14) near the proximal end tion of the carpal canal together. Lying medial
of the carpus divides into medial and lateral to the accessory carpal bone, the palmar branch
branches. The medial branch, upon reaching the issues a small branch to the carpal pad. A larger
metacarpus, divides again to form the superficial branch runs almost directly medially at the dis
palmar m etacarpal nerves I an d II (nn. meta- tal end of the palmar carpal fibrocartilage and
carpales palmares superficiales I et II). innervates the special muscles of the fifth digit.
The superficial palmar metacarpal nerve I The superficial palmar metacarpal nerve IV
runs to the web of skin of the first interdigital (n. metacarpalis palmaris superficialis IV) arises
space and bifurcates into the proper nerves from the palmar branch of the ulnar nerve as it
which supply part of the skin of the adjacent traverses the carpus. It passes laterally across
palmar sides of the first two digits. These nerves the proximal end of the fifth metacarpal bone
will be described with the portion of the ulnar and sends a branch to the skin on the lateral sides
nerve which innervates the forepaw. The lateral of the fifth metacarpal bone and fifth digit as the
branch into which the median nerve divides be lateral palmar digital nerve V (n. digitalis pal
comes the superficial palm ar metacarpal nerve maris lateralis V).
III (n. metacarpalis palmaris superficialis III). At the proximal end of the metacarpus the
The main superficial palmar metacarpal nerves palmar branch of the ulnar nerve divides into
are formed as follows: nerves II and III from two series of branches. One set is short and is
the median; nerve IV from the ulnar. All three composed of the muscular branches which fre
of these nerves anastomose with the comparable quently arise from the second set or the deep
deep branches of the ulnar to form the common palmar metacarpal nerves.
palmar digital nerves. These nerves and certain The deep palmar metacarpal nerves I, II, III,
irregularities concerning them are described and IV (nn. metacarpales palmares profundi I,
after the description of those portions of the ul II, III, et IV) are usually the terminal branches
nar nerve which are located in the paw. of the palmar branch of the ulnar nerve. The
The ulnar nerve o f the forepaw (n. ulnaris palmar branch passes through the lateral portion
manus) (Fig. 11-14) is represented by the dorsal of the carpal canal where it lies in close associa
and palmar branches, which arise as terminal tion with the relatively large terminal portion of
branches of the ulnar nerve at the junction of the palmar interosseous artery. On the deep
the proximal and middle thirds of the antebra surfaces of the abductors of the second and fifth
chium. digits at their origins the palmar branch of the
The dorsal branch (ramus dorsalis) passes dis ulnar nerve divides into the deep palmar meta
tad toward the lateral aspect of the carpus by carpal nerves I, II, III, and IV and into muscular
passing obliquely laterad between the caudally branches.
lying flexor carpi ulnaris and the cranially lying The muscular branches (rami musculares)
extensor carpi ulnaris muscles. It perforates the arise directly from the palmar branch of the ul
heavy deep antebrachial fascia from 3 to 8 cm. nar or individually from the several deep palmar
proximocaudal to the styloid process of the ulna. metacarpal nerves. They innervate the four in
Closely applied to the skin and in association terosseous muscles, the three lumbricales, the
with a cutaneous branch of the palmar interos special muscles of the first, second, and fifth
seous artery, it obliquely crosses the lateral side digits, and the single, weak flexor digitorum
of the carpus. In its subcutaneous course on the brevis muscle.
dorsolateral surface of the forepaw it is called At first the deep palmar metacarpal nerves
the lateral dorsal digital nerve V (n. digitorum lie on the palmar surfaces of the proximal ends
dorsalis lateralis V), where it supplies the skin of of the interosseous muscles. As they run distad
the dorsolateral surfaces of the metacarpus and toward the digits they lie between the main in
the fifth digit. terosseous muscles in relatively superficial posi
The palmar branch (ramus palmaris) of the tions. Anastomoses occur between the super
forepaw is the main continuation of the ulnar ficial and deep palmar metacarpal nerves near
nerve after the dorsal branch has arisen. As it or at the distal ends of the metacarpal bones.
passes through the distal portion of the ante By these unions the common palmar digital
brachium it lies on the caudomedial surface of nerves 11, 111, and IV (nn. digitales palmares
T h o r a c ic N erves 591
communes II, III, et IV) are formed (Fig. 11-15). vical nerve before it branches to aid in forming
The anastomoses between the members of the the brachial plexus. It runs largely horizontally
two sets—superficial and deep—are irregular on the superficial surface of the thoracic portion
and in some instances multiple. Occasionally of the serratus ventralis muscle which it supplies.
the deep members send slender anastomotic Miller (1934) regards a small branch from the
branches to the proper palmar digital nerves fifth cervical nerve as also belonging to the long
into which the common palmar digital nerves thoracic nerve.
terminally divide. The common nerves are there The dorsal thoracic nerve (n. thoracodorsalis)
fore short as they cross the contact surfaces of (Fig. 11-6) arises primarily from the eighth cer
adjacent metacarpophalangeal joints. From the vical nerve with contributions from the first tho
common nerves II, III, and IV or occasionally racic and/or the seventh cervical nerves. It is
proximal or distal to these nerves the three sen the motor nerve to the latissimus dorsi muscle.
sory nerves arise which innervate the large meta It runs caudodorsad in close relation to the tho
carpal foot pad. Minute twigs from the common racodorsal vessels on the medial surface of the
palmar digital nerves supply the structures of muscle.
the metacarpophalangeal joints of the four main The ventral thoracic nerves (nn. thoracales
digits. ventrales) (Fig. 11-5) represent all the motor
The m edial and lateral palm ar proper digital nerves to the pectoral musculature and the cu-
nerves II, III, and IV (nn. digitales proprii pal taneus trunci muscle (nn. pectorales of N. A.).
mares laterales et mediales) are the terminal Confusion exists in both the veterinary and hu
branches into which each of the three main com man literature on the name of the nerves to the
mon digital nerves divides. They lie on the pectoral musculature. The term “ventral tho
contact sides of the four main digits, dorsal to racic nerves” was chosen since there is a dorsal
the comparable vessels, and like the vessels the thoracic nerve and there is precedence for using
proper nerves which face the axis through the the term (Schaeffer 1953; Goss 1954). The ven
paw (axial branches) are larger and longer than tral thoracic nerves are unusually irregular in
are the abaxial proper nerves. They supply the number and origin. In most specimens a cranial
skin under which they lie and the digital joints and a caudal group can be recognized. Usually
which they cross. At the distal interphalangeal the two nerves which compose the cranial group
joints sensory branches are supplied to the dig derive their fibers from the sixth, seventh, and
ital foot pads. Medial and lateral proper digital eighth cervical nerves. They supply the super
nerves enter the palmar vascular canal of the ficial pectoral muscle. The caudal thoracic
distal phalanges, and thereby nerve fibers reach group is represented by three or four branches
the corium of the horny claw. which innervate the deep pectoral muscle. They
are composed of fibers which come from the
eighth cervical and the first and second thoracic
N e r v e s o f t h e B r a c h ia l P l e x u s W h ic h nerves. They arise from the ventral border of the
S u p p l y E x t r in s ic M u s c l e s o f t h e lateral thoracic nerve (n. thoracolateralis). The
T h o r a c ic L im b long-branched nerve is the sole motor supply to
the cutaneus trunci muscle. According to Lang
The nerves in this group are smaller than the worthy (1924), it runs caudally on the deep sur
nerves which supply the intrinsic structures of face of this cutaneous muscle. At first the lateral
the thoracic limb. They consist of the brachio thoracic nerve accompanies the lateral thoracic
cephalic nerve, long thoracic nerve, dorsal tho artery and vein and lies between the adjacent
racic nerve, and ventral thoracic nerves, includ borders of the latissimus dorsi and deep pectoral
ing the lateral thoracic nerve. muscles after passing medial to the axillary and
The nerve to the m. brachiocephalicus (n. accessory axillary lymph nodes. Langworthy re
brachiocephalicus) (Allam et al. 1952) arises gards this nerve as a branch of the ventral tho
mainly from the sixth cervical nerve, but it may racic (pectoral) nerves rather than vice versa.
be joined by a branch from the fifth cervical
nerve. It passes directly laterad into the brachio THORACIC NERVES
cephalicus muscle cranial to the shoulder joint.
Branches from the cervical plexus also supply The thoracic nerves (nn. thoracici) (Fig. 11-
the brachiocephalicus. 16) number thirteen pairs in the dog, and as a
The long thoracic nerve (n. thoracicus longus) group they retain the simplest segmental form
(Fig. 11-2) usually arises from the seventh cer of all the spinal nerves. Each pair of thoracic
592 Chapter 11. S p in a l N er v e s
„ M. t e n s o r fa s c ia e a n te b r a c h ii
M e d ia n n .^ _
, ^Caud. cutan. a n te b r a c h ia l n.
M u s c u I a c u t o n e o u s n __ \\
. C o l l a t e r a l u l n a r a.
M .bic e ps b r a c h i i __
_ - U ln a r n.
M e d ia n a.—
Prox. c o l l a t e r a l r a d i a l a—
Superf- r a d i a l n.,medial b r .
M.ext. c o r p i r a d i a l i s - _ _ M. fle x o r carpi ulnaris, coput ulnare
Med. cutan. a n t e b r a c h i a l n. —
^Acc. in terosseous o
M .p ro n a to r te re s -
__To m. f(ex. digit, prof., ca pu t ulnare
M . f /ex. c a r p i r a d i a l i s —
__ M.flex. c a rp i u ln a r is , caputhum .
(re fle c te d )
" '- D o r s a l br., u l n a r n.
M. flex, d i g i t o r u m superf.-
fjjjU flex, d ig ito r u m prof., c a p u t hum.
C ran.cutan a n t e b r a c h i a l br. -
P a lm a r a n t e b r a c h i a l a
U l n a r a.- -
M e d ia n n - '
R a d i a l a . - 'I
F k . 1 1 -1 2 . Nerves of the right antebrachium , medial aspect. D issection showing th e ulnar nerve.
T h o r a c ic N erves 593
- S u p e r f . r a d i a l n., m e d br.
P a lm a r in te ro s s e o u s a
- Prox. c o l l a t e r a l r a d i a l a., m e d b r
D ist. d o r s a l i n t e r o s s e o u s a - S u p e r f r a d i a l n., l a t . b r
- P r o x c o l l a t e r a l r a d i a l a., la t. b r
U ln a r n, d o rsa l br -
- - R a d i a l a., d o r s a l b r
B ra n c h e s of p a lm a r in te r o s s e o u s a - - I
-D a rsa l d ig ita l n I
D e e p d o r s a l m e t a c a r p a l ao.~ J r-
- Bn o f u l n a r a-
S u p e rf d o rs a l m e ta c a rp a l a a - t t -
- D o r s a l m e t a c a r p a l n n . I V I I I , II
D o rsa l common d i g i t a l a a
[ - M e d d o r s a l d i g i t a l nn. V, IV ,111,11
P a l m a r common d i q i t a l a a ^ -
L a t d o r s a l d i g i t a l nn ’/ , IV, 111,11
nerves has the same serial number as the verte II) differs from the intercostal nerves which lie
bra which lies in front of their intervertebral caudal to it in two respects. It usually sends a
foramina of exit. All the ventral branches with communicating branch to the first cervical
the exception of the first three or four send twigs nerve, and secondly its distal lateral cutaneous
at their distal ends into the rectus abdominis branch is the largest of all these branches;
muscle, since this muscle traverses both the tho furthermore, this branch runs to the thoracic
rax and the abdomen. Unlike typical spinal limb and supplies a patch of skin in the region of
nerves, the ventral branches do not divide into the elbow joint. It is named the second inter-
medial and lateral branches. costobrachial nerve (n. intercostobrachialis II).
The m edial branches (rami mediales) of the Usually the distal lateral cutaneous branch o f
dorsal branches of the thoracic nerves run es the third intercostal nerve (n. intercostobrachi
sentially parallel and caudal to the first ten cau alis III) also innervates a portion of the thoracic
dally sloping thoracic vertebral spines, with limb proximal to that of the second nerve in most
which they correspond in number. Caudal to the specimens.
eleventh thoracic spine the thoracic spines slope The subcostal nerve (n. subcostalis) is the ven
cranially, and the corresponding nerves cross tral branch of the last or thirteenth thoracic
them obliquely, since they run caudodorsally. nerve. It supplies a band of the abdominal wall
The medial branches supply the multifidus tho which lies adjacent to the caudal border of the
racis, the rotatores, longissimus dorsi, and the last rib and then continues tangentially to the
spinalis et semispinalis thoracis et cervicis mus last rib and costal arch in the abdominal wall. In
cles. It is probable that the vertebrae, ligaments, its area of distribution it lies cranial but parallel
and dura receive branches from these nerves. to the bands of the abdominal wall which are
The medial branches do not end in cutaneous supplied by the ventral branches of the first three
branches. lumbar nerves. It divides into lateral and medial
The lateral branches (rami laterales) of the branches which resemble those of the lumbar
dorsal branches of the thoracic nerves run cau nerves lying caudal to it.
dolaterally at about a 45° angle to a sagittal A typical intercostal nerve (n. intercostalis)
plane. They course between the longissimus (Fig. I I - 17) is disposed as follows: Each begins
dorsi muscle dorsally and the iliocostalis ven where the dorsal branch of the particular tho
trally to reach the medial surfaces of the seg racic nerve arises. For about the first centimeter
ments of the serratus dorsalis muscles where it lies embedded in the dorsal border of the
these are present. They usually perforate these cranioventrally running internal intercostal
segments as well as the iliocostalis dorsi and muscle. It then turns distad on the medial sur
cutaneus trunci muscles to reach the skin, where face of the internal intercostal muscle where it is
they become the proximal lateral cutaneous separated from the caudal border of the cor
branches (rami cutanei laterales). These nerves responding rib by the intercostal vein and the
divide in the superficial fascia into short medial intercostal artery. Variations in this order of
branches and longer ventrolateral branches arrangement occur most frequently in the cranial
which supply the skin of approximately the dor part of the series. This triad of structures is
sal third of the thorax. As these nerves cross the surrounded by a variable quantity of fat. The
medial border of the iliocostalis dorsi muscle nerve lies adjacent to or among the fiber strands
they send branches to it and to the levator costae of the internal intercostal muscle. In the caudal
muscle segments. part of the thorax fleshy sheets of the internal
The ventral branches (rami ventrales) of the intercostal muscle from the intercostal space in
thoracic nerves with the exception of most of the front of it extend over the ribs medially and
first and the thirteenth are more commonly cover the intercostal vessels and nerves which lie
known as the intercostal nerves (nn. inter caudal to the rib. In most intercostal spaces the
costales). intercostal vessels and nerves are covered medi
The major portion of the first thoracic nerve ally only by the pleura.
(ventral branch) passes forward medial to the A typical intercostal nerve has the following
neck of the first rib and contributes appreciably branches:
to the formation of the brachial plexus. The 1. The visceral branch (ramus visceralis), or
intercostal portion of the first thoracic nerve is ramus communicans, which contains efferent
but a delicate twig which enters the muscles of sympathetic fibers with which are intermingled
the first intercostal space. afferent fibers from visceral structures. These
The second intercostal nerve (n. intercostalis branches are connections between the initial
L um bar N erves 595
part of the intercostal nerve and sympathetic alba on the superficial surface of the transversus
trunk, usually at a ganglion. This branch is about abdominis muscle, which they supply before
1 mm. in diameter and 3 mm. long. terminating in the rectus abdominis. These
2. The proximal muscular branch (ramus nerves fail to send ventral cutaneous branches to
muscularis proximalis) leaves the dorsal side of the skin.
the intercostal nerve 1 or 2 cm. from its origin.
Before it perforates the external intercostal mus
cle it sends a long slender branch distally on the LUMBAR NERVES
deep (medial) surface of the external intercostal
muscle, which lies only a few millimeters caudal The lumbar nerves (nn. lumbales) (Figs. 11-
to the corresponding rib. It sends delicate twigs 19, 11-20) are seven in number on each side.
to the external intercostal muscle throughout its Each member of a pair has the same number as
length. Upon contact with the external inter its intervertebral foramen of exit and the verte
costal muscle a branch leaves the nerve which bra which lies cranial to it. The middle of the
runs laterally and supplies the serratus dorsalis first lumbar segment of the spinal cord lies op
muscle. posite the fibrocartilage which connects the first
3. The distal lateral cutaneous branch (ramus two lumbar vertebrae. The nerve roots, there
cutaneus lateralis distalis) (Fig. 11-18) passes fore, of the first pair of lumbar nerves lie essen
through the mid-lateral portion of the thoracic tially in the same transverse area as the foramina
wall and runs distad in the superficial fascia with of exit of these nerves. As traced caudally, the
the like-named artery. The aggregate of these segments of the spinal cord are shorter than the
branches supplies all the skin on the ventro vertebral segments, so that the spinal cord ends
lateral half of the thoracic wall except a narrow, dorsal to the fibrocartilage between the sixth
longitudinal ventral strip. The first two of these and seventh lumbar vertebrae. Because of this
nerves supply skin on the thorax and a portion disproportionate length the last several pairs of
of the thoracic limb. The limb portions of these spinal nerves run increasingly longer distances
nerves (intercostal brachial nerves) supply zones within the spinal canal before leaving their osse
of skin covering the triceps muscle, proximal to ous confines by means of the intervertebral
and over the elbow joint. In the regions of the foramina than do the spinal nerves cranial to
thoracic mammary glands branches from the them. The leash of nerves thus formed is called
distal lateral cutaneous branches (lateral mam the cauda equina. According to Hopkins (1935),
mary branches) ramify under the skin. in a 40- to 50-pound dog the intraspinal extent
4. The distal muscular branch (ramus mus of the lumbar nerves varies from 0 .6 cm. for the
cularis distalis) consists of two branches. One is first pair to 3.5 cm. for the seventh. This author
short and enters the transversus thoracis mus was undoubtedly measuring the nerve roots
cle; the other branch passes to the outside of the rather than the spinal nerves which they form.
rib cage and enters the rectus abdominis muscle. The reason for this spatial disparity between the
5. The ventral cutaneous branch (ramus length of the spinal cord and the vertebral col
cutaneus ventralis) is the terminal part of each umn is a continuation of the growth of the ver
typical intercostal nerve. It runs to the skin by tebral column after the spinal cord has stopped
crossing the lateral surface of the sternum. In growing. The lumbar spinal nerves are formed
the superficial thoracic fascia, which lies on the by the merging of the dorsal and ventral roots
medial portion of the deep pectoral muscle, it is at the intervertebral foramina. As the roots of
closely bound to the only slightly larger ventral the several lumbar spinal nerves run caudally
cutaneous artery. The aggregate of these nerves their proximal halves lie within the dural cover
supplies a zone of skin about 2.5 cm. wide which ing of the spinal cord, and their distal halves lie
lies adjacent to the midventral line. The terminal in dural sheaths in soft epidural fat. Like the
twigs of the ventral cutaneous branches of the typical spinal nerves of the preceding regions,
fifth and seventh intercostal nerves ramify in each lumbar spinal nerve divides upon leaving
the skin covering the medial portions of the two the intervertebral foramen into small dorsal and
thoracic mammary glands when these glands are larger ventral branches. The actual length of
functional. They are named the medial mam each lumbar spinal nerve is only a few milli
mary branches (rami mammarii mediales). meters, and it lies largely in and just lateral to
The ventral extensions of the last three inter the intervertebral foramen through which it
costal nerves leave the intercostal spaces medial passes.
to the costal arch and extend toward the linea The dorsal branches (rami dorsales) of the
(Text continued on page 599.)
396 Chapter 11. S p in a l N erves
R a d ia l a —
U l n a r n., p a l m a r br.
M e d ia n n.—
D o r s a l bn — - P a l m a r in te r a s s e o u s a.
Pal m ar b r — — U ln a r n., d o r s a l br
UI n a r a -
M e d i a l br
L a t e r a l bn
Superf. p a l m a r a r t e r i a l a r c h
■Deep p a l m a r m e t a c a r p a l n n ,ll,llIJ V
- S u p e r f p a l m a r m e ta c a r p a l
nn. II III, IV
P a lm a r c o m m o n
d i g i t a l n n I I I I I , IV —
- » N n to m e ta c a r p a l f o o t pad
Med. p a l m a r p r o p e r d i g i t a l - -
nn ilJU, IV, V ' L a t p a l m a r d i s t a l n.V
L at p a lm a r p ro p e r d ig ita l -
nn.II, III , I V
N n to d i g i t a l f a a t p a d
S u p e rficia l d o rs a l m e ta c a rp a l a IV- -
r dorsal m eta carp al n.IV
- -Superf palmar m e t a c a r p a l a tf n II
S u p e rfic ia l a o rs a ! m e t a c o r p a l a l i i
- Superf palm ar metacarpal a IV
D o rsa l m e ta c a rp a l n. Ill —
C om m on d ig it a l n III
D orsal common d i g i t a l a ll!
Medial d o rs a l p ro p e r —
digi ta l a + n. I V
Medial palm ar p r o p e r d ig ita l a. 4-n IV
- D ig ita l fo o t pad IV
F ig . 1 1 -1 5 . Arlenes anil nerves o f th e fourth digit ami m etacarpus, medial aspect. From Miller. 19581
598 C h apter 1 1 . S p in a l N e r v e s
M. t r a p e z i u s
M. longissim us d o rs i
M m .s p in a l is r s e m i s p i n a / is
th o ra c is r c e rv ic is \ M. le v a to r co s tae
Ao r t o ‘ [- - ~ M .la t is s im u s d orsi
A z y g o s v.
- -Prox. m u s cu la r br.
" M. i n t e r c o s t a l i s int.
M.oblic^uus a b d o m in is ext.
P le u ra
D is ta l m u s c u la r br
x M. r e c t u s a b d o m in is
'M . t r a n s v e r s u s t h o r a c i s
xtnt. t h o r a c i c a. v v.
'/W. p e c t o r a l is p ro f.
‘ V e n t r a l c u tan eo u s br.
S ternu m
F ig . 11-16. Diagram of the sixth thoracic nerve. (Section caudal to the sixth rib.)
L u m bar Nerves 599
lumbar nerves are similar throughout most of dorsal cutaneous branches. The lumbar nerves
the region. Each typically divides into medial caudal to a hyperdeveloped dorsal cutaneous
and lateral branches. The m edial branches (rami branch do have dorsal muscular branches which
mediales) arborize in the longissimus lumborum are dissipated in the epaxial musculature with
muscle which they supply and send terminal out first dividing into medial and lateral parts.
twigs to the multifidus lumborum and the inter- The ventral branches (rami ventrales) of the
spinales lumborum muscles. They run caudo- seven pairs of lumbar nerves are variable, but
dorsad obliquely across the lateral surface of the less so than the dorsal branches. They are usually
cranially inclined spinous processes of the verte described as lumbar nerves without specifically
brae which follow them. They are separated referring to them as ventral branches. Each lum
from the ventral borders of the tendons of the bar nerve is connected to the sympathetic trunk
longissimus lumborum muscle which go to the by a visceral ramus (ramus visceralis), or ramus
accessory processes of the lumbar vertebrae by communicans, with rare exceptions. The con
the large branches of the dorsal branches of the nections are exceedingly variable, as Mehler
lumbar segmental arteries. Only an occasional et al. (1952) have pointed out. In 100 dogs dis
medial branch runs far enough peripherally to sected by these investigators only twenty-three
reach the skin. Pederson et al. (1956) found in specimens had symmetrically located bilateral
the human being that small branches enter the trunk ganglia at every lumbar segment. These
spinal canal and contain sympathetic and sen twenty-three specimens also had at least two
sory fibers which anastomose with each other paired sacral ganglia. The rami communicantes
and supply the dorsal longitudinal ligament, may be double, or the rami from two adjacent
dura mater, periosteum, and blood vessels. The nerves may go to the same ganglion. The first
lateral branches (rami laterales) of the dorsal four or five rami contain preganglionic as well
branches of the first three or four lumbar nerves as postganglionic fibers. The remaining commu
are clearly separated from the medial branches. nicating rami contain only postganglionic and
The dorsal branches of the last three or four afferent fibers. The communicating rami, as they
lumbar nerves do not clearly divide into medial run between the lumbar nerves and the sympa
and lateral portions, but they arborize in the thetic trunk, lie largely under the psoas minor
epaxial muscles of the loin. The lateral branches muscle. They are less than 1 mm. in diameter
of the dorsal branches of the first three or four and about 5 mm. long. Like the brachial plexus
lumbar nerves run caudolaterad through the which gives origin to the nerves which innervate
longissimus and iliocostalis muscles and perfo the thoracic limb, the last five lumbar nerves
rate the iliocostalis mid-laterally in a segmental and all the sacral nerves are joined together to
manner. After continuing in the intramuscular form the lumbosacral plexus (plexus lumbo-
caudolateral direction in the areolar tissue under sacralis), which issues the nerves to the pelvic
the lumbodorsal fascia one or more centimeters, limb. This plexus, therefore, is divided into lum
they perforate the lumbodorsal fascia and ar bar and sacral portions. The first two lumbar
borize in the skin of the dorsolateral parts of the nerves are usually not joined to each other or to
lumbar and sacral regions as the dorsal cutaneous adjoining nerves, but run caudolaterally in the
branches (rami cutanei dorsales) (nn. clunium abdominal wall in series with the last several
superiores of N.A. terminology). caudal thoracic nerves and are therefore not in
The lateral branches of the dorsal branches cluded in the lumbosacral plexus.
also supply the lumbar portion of the iliocostalis The cranial and caudal iliohypogastric nerves
muscle. The cutaneous branches of the dorsal (nn. iliohypogastrici craniales et caudales) (Fig.
branches are unusually variable. Occasionally 11 - 2 0 ) represent the ventral branches of the first
all the lumbar nerves have these branches. In and second lumbar nerves respectively. Each
other specimens a single dorsal branch bifurcates nerve is connected with the sympathetic trunk
deeply within the epaxial musculature, resulting by a single ramus communicans which contains
in two cutaneous nerves which supply the skin both preganglionic and postganglionic fibers.
of each side of the dorsum of the loin. This varia Both nerves send branches to the quadratus lum
tion is similar to those encountered in the borum and the psoas minor muscles.
cervical region. In some specimens the dorsal After leaving the caudal thoracic portion of
cutaneous branch of either the fifth, sixth, or sev the hypaxial musculature by passing between
enth lumbar nerve supplies the skin lying adja the two segments of the quadratus lumborum
cent to it as well as much of that over the rump. muscle the cranial iliohypogastric nerve lies in
In these specimens there is an absence of some the subserous endothoracic fascia at its origin. It
(Text continued on page 606.)
600 C h ap ter 11. S p in a l N e r v e s
M. m u l t i f i d u s
7 t h t h o r a c i c n. . '
M. s p i n a l i s * s e m i s p i n a l i s d o r S i \
D o r s a l br.s v '
\ N '
M lo n g is s im u s d orsi \ N \ \ /M. in t e r s p i n a l i s
\ N v \ /
7th i n t e r c o s t a l n.^ . \ v \ / sM. tr a p e z iu s
M. i l i o c o s t a l i s d o r s i ^ <-D o rsa l b r
M. l e v a t o r c o sto e
M. s e r r a t u s d o r s a l i s . _ - 6 t h th o r a c ic n.
-M. s e r r a t u s vent.
Mm. i n t e r c o s t a l e s int.- " D is t: lat. cutaneous br
M u s c u la r br. ~ ^M. l a t is s i m u s d o rs i
''M . sca le nu s
7th r i b "
"M m . in te r c o s ta le s ext.
P le u ra " 6th r i b
M.abliq_uus e x t -M. re c tu s a b d o m in is
a b d o m in is
^ V e n tr a l cutan eo u s br.
Fic. 11-17. Dissection showing distribution of the thoracic nerves, right lateral aspect.
L um bar N erves 601
13th r i b
D o rs a l b r of T 13 i
V e n t r a l b r of T 13( (
I
C r a n ia l iI io h y p o g a s tr ic n.(LI)^
D o r s a l c u ta n e o u s b r \ '
C audal i l i o h y p o g a s t r i c n .(L 2 ) ^ '
M. i l i o c o s t a l i s lumborum x \ \
\ \ \ i
I l i o i n g u in a l n.(L3) \ \ \
G e n ita l n
M .g lu te u s m edius ■
M. s a r t o r iu s
B r a n c h e s to m re c tu s a b d o m in is 1
PVHguIUM
Med i a l b r of L I 1
Lat. cutaneous b r of LI 1
Mm. tr a n s v e r s u s a b d o m in is , o b liq u u s abd. int., o b liq u u s abd. ext}
F ig . 1 1 -2 0 . Dissection showing the arrangement of the first four lumbar nerves, lateral aspect.
604 Chapter 11. S p in a l Nerves
Wing of sa c ru m
To m . c o c c y g e u s i 7 t h lu m b a r n.
Darsal br. of L5
, V e n tra l br. o f L5
To m. l e v a t o r a n i s
Cutaneous br.''
P u d e n d a l n. -
Caud. c u ta n . f e m o r a l n. - -
P e r i n e a I n. ' Ilio in g u in a l n
L o t c u t a n fe m o r a l n.
Caud. r e c t a l n.' '
>G e n ita l n.
P e l v ie s p l a n c h n i c n " \
>F e m a ra l n.
To mm.quad, f e m o r i s v g e m e l l i
O b t u r a t o r n.
' N. to c a u d a l p o r t i o n o f m. g l u te u s med.
I s c h i a t i c n.
C r a n ia l g l u t e a l n.
To m. o b t u r a t o r int.
C au d al g l u t e a l n.
- - I l i o i n g u i n a l n. ( L3)
P o s tc a v a - -M . q u a d r a t u s lu m b o ru m
-M .p s o a s m in o r
A o rta -
-Lat. cuton. fe m o r a l n (L4)
~M t r a n s v e r s u s abd-
Deep c i r c u m f l. i l i a c o .w .
M.obhg. o b d o m i n i s ext.~ - -M. p s o o s m a j o r
- - G e n i t a l n.
Caud. s u p e r f
e p ig a s tric a . rv. - F e m o r a l n.
~Exr. i l i a c a.
L. u t e r i n e h o r n
~Med.br., i l i o i n g u i n a l n
M. s a r t o r i u s -
Caud. a b d o m i n a l a r v .
Ing u in a l -
m am m ary g lan d F e m o r a l a.rv.
Round h g a m e n l -
~ D e e p f e m o r o l o .rv .
Ext. i n g u m o l r i n g ~
Round l i g of u t e r u s
F e m o ro l a c w—
''Caud. deep
M .p e ctin e u s - e p i g a s t r i c arv.
O b tu ra to r n '
M. a d d u c t o r ' t 1 ( 'M . r e c t u s a b d o m i n is
M .g ro c il i s 1 ' P u d e n d o e p ig a s tr ic t r u n k
G e n ita l n , e x t p u d e n d o l a. v v . 1 P e r i n e a l a.,v. v n.
F ig. J 1-22. Dissection showing course of the genital nerve in the female, ventral aspet t . , The left abdominal wall is reflected.)
606 Chapter 11. Sp in a l N e r v e s
then passes into the subserous transversalis fascia (Fig. 11-21) consists of the intercommunicating
of the abdomen by passing dorsal to the lumbo ventral branches of the last five lumbar and the
costal arch. About 4 cm. ventrolateral to a plane three sacral nerves. It may be divided into lum
through the free end of the lumbar transverse bar and sacral plexuses. Mehler et al. (1952) con
processes the nerve, after having passed through sidered the variations of the lumbosacral sympa
the aponeurosis of origin of the transversus ab thetic trunk in the dog. The communicating rami
dominis muscle, divides into a lateral and a me and sympathetic roots were found to be highly
dial branch. Before dividing it gives branches to variable, particularly in the lumbar segments.
the serosa and to the segments of the quadratus In 65 per cent of the 100 animals they dis
lumborum muscle, against which it lies. It then sected, L4 was the lowest from which communi
runs between two adjacent fleshy bundles of the cating rami originated. The remaining 35 per cent
transversus abdominis muscle as these bundles of animals had communicating rami that arose
arise from a narrow aponeurosis which attaches from spinal nerves L5 and L 6 .
to the ends of the transverse processes of the The lumbar plexus (plexus lumbalis) is a term
lumbar vertebrae. Shortly after entering the ordinarily restricted to the interconnected third,
fascia which separates the transversus abdominis fourth, and fifth lumbar nerves (Havelka 1928).
from the internal abdominal oblique muscle the In some specimens the third lumbar nerve is
iliohypogastric nerve divides into lateral and connected to the second by a fine branch (Brad
medial branches. ley and Grahame 1959), and usually the fifth
The lateral branch (ramus lateralis) passes lumbar nerve is connected to the sixth. The divi
through the internal abdominal oblique to run sion of the lumbosacral plexus into its two com
in the septum between the two abdominal ponents is made primarily because of the location
oblique muscles. In its course ventrocaudally of the plexuses and not its origin. The connection
it sends most of its branches to these muscles, between the third and fourth lumbar nerves
and near the middle of the abdomen it perforates gives origin to the genital branch whereas the
the external abdominal oblique to become connection between the fourth and fifth is usu
subcutaneous as the lateral cutaneous branch ally devoid of branches. The morphology of the
(ramus cutaneus lateralis). It is accompanied lumbar nerves is particularly variable. The lum
by a twig of the cranial abdominal artery and bar plexus may be moved cranially one segment
vein. The lateral cutaneous branch is distrib from the normal (prefixed) or caudally one seg
uted to a ventrolaterally running band of skin ment from the normal (postfixed).
which crosses the junction of the cranial and The ilioinguinal nerve (n. ilioinguinalis) (Fig.
middle thirds of the abdomen caudal to the ribs. 11 - 2 0 ) is the direct ventrolateral continuation of
The m edial branch (ramus medialis) lies the third lumbar nerve. Its ramus communicans
closely applied to the lateral surface of the trans contains both preganglionic and postganglionic
versus abdominis muscle where it appears in fibers. It anastomoses with the fourth lumbar
series with the last five thoracic and the second nerve and divides into medial and lateral
and third lumbar nerves. Like the lateral branch branches which resemble those of the lumbar
it is also accompanied by a small branch of the nerves which precede it. Cutaneous branches
cranial abdominal artery and vein. It supplies a from its lateral portion extend caudoventrally
band of the transversus abdominis muscle and superficial to the lateral cutaneous femoral nerve
peritoneum along its course. It ends in the first and ramify in the skin of the craniolateral sur
lumbar segment of the rectus abdominis muscle face of the thigh. The medial branch of the ilio
as well as the skin and peritoneum covering it. inguinal nerve is small. It runs more caudally
The ventral branch of the second lumbar nerve than ventrolaterally. It is accompanied by a cra
is in all respects similar to the first lumbar nerve nial branch of the ascending branch of the deep
except that it supplies the abdominal wall caudal circumflex iliac vessels. It usually has no grossly
to it and appears at the lateral border of the demonstrable cutaneous branches.
hypaxial musculature after having passed be The lateral cutaneous fem oral nerve (n. cuta
tween the quadratus lumborum and the iliopsoas neus femoris lateralis) is formed primarily by the
muscles at the lumbocostal arch. Occasionally ventral branch of the fourth lumbar nerve, al
one of the iliohypogastric nerves is double with a though there are connections with both the third
reciprocal diminution in size of the nerve caudal and fifth lumbar nerves. It is larger than the lum
to it. Rarely is a single nerve formed by the fusion bar nerves which precede it, but smaller than
of the first and second or the second and third those which follow it. As it runs caudolaterad
lumbar nerves. through the substance of the psoas minor muscle
The lumbosacral plexus (plexus lumbosacralis) it sends branches to it and to the other hypaxial
L um bar N erv es 607
muscles of the region. According to Mehler et al. supplies a zone of skin on the caudomedial sur
(1952), the fourth lumbar nerve is the lowest face of the thigh.
nerve from which a communicating ramus The fem oral nerve (n. femoralis) (Fig. 11-19)
arises. The main portion of the nerve passes arises primarily from the fifth segment of the
through the abdominal wall, lying between the lumbar plexus with a strong root of origin also
deep circumflex iliac artery cranially and the coming from the fourth. A smaller branch of ori
satellite vein caudally. It passes between the gin may come from the third, but rarely does one
lumbar and inguinal portions of the internal ab come from the sixth segment. After being formed
dominal oblique and over the dorsal margin of in the substance of the iliopsoas muscle it con
the external abdominal oblique muscle. Its ter tinues caudally in the substance of this muscle
minal cutaneous branches are variable, but in and leaves the abdomen along with the muscle.
general they follow the accompanying vessels. During its intra-abdominal course it sends mus
A branch ramifies in the skin in the region of the cular branches to the iliopsoas. The prominent
tuber coxae and the adjacent cranial part of the saphenous nerve arises from its cranial side.
rump; other branches supply the skin over Shortly thereafter the femoral nerve enters the
the cranial portion of the thigh, while its longest quadriceps femoris muscle by passing between
branch runs distally, supplying the skin over the the rectus femoris and the vastus medialis mus
thigh and the lateral surface of the stifle joint. cles at the proximal end of the cleft which sepa
Kunzel (1957) illustrates this nerve in his work rates these heads. It supplies all four heads of the
on the topography of the hip joint. quadriceps muscle (rectus femoris, vastus medi
The genital nerve (n. genitalis or ramus geni alis, vastus intermedius, and vastus lateralis) and
talis of the n. genitofemoralis) (Fig. 11-22) is also sends a small twig to the capsularis coxae
called the external spermatic nerve in many muscle. The branches to the various parts of the
veterinary publications. The dog lacks the sep quadriceps largely accompany the cranial fem
arate femoral branch found in man, but does oral artery, which is its chief source of blood
have a cutaneous ramus from the genital branch supply proximally. No cutaneous branches arise
which supplies the skin of the pudendal region. from the femoral nerve, nor could a branch be
The genital nerve arises from the third and fourth found going to the hip joint.
lumbar nerves, the root from the third being The saphenous nerve (n. saphenus) (Figs.
larger than that from the fourth. Bradley and 11-23, 11-30) is the only superficial branch of
Grahame (1959) state that the contribution from the femoral nerve. Arising from the femoral be
the fourth may be absent. Ellenberger and Baum fore this nerve leaves the iliopsoas, the saphe
(1891) state that occasionally the nerve is double, nous becomes related to the medial surface of
and in some specimens the ilioinguinal nerve the tensor fasciae latae muscle and immediately
joins the genital and is distributed with it. Usu divides into muscular and cutaneous branches.
ally the nerve is single, small, and long. It is This division is lacking in many specimens, since
formed in the substance of the medial portion of the muscular branch arises from the femoral
the iliopsoas muscle near the body of the fourth nerve either proximal or distal to the origin of
lumbar vertebra. As it runs caudally it leaves the that part of the saphenous nerve which is cuta
substance of the medial part of the muscle and neous. The muscular branch (ramus muscularis)
continues caudally in the fat which fills the ir bifurcates, one twig going to the cranial belly of
regularities around the postcava and aorta. After the sartorius muscle and the other to the caudal
passing dorsal to the external iliac lymph node to belly. These nerve branches accompany the
which it sends a small branch, it becomes related blood vessels serving the proximal part of the
to the distal portion of the external iliac artery, sartorius muscle. The cutaneous branch (ramus
which it crosses medially. It leaves the abdomen cutaneus) of the saphenous nerve is long and
by passing through the inguinal canal where it slender. It lies in apposition to the cranial sur
lies medial to the spermatic cord in the male or face of the femoral artery as it runs distally across
associated with the round ligament of the uterus the medial surface of the quadriceps muscle. It
in the female. Minute muscular branches are sends branches to the skin of the medial surface
given oif to the external cremaster muscle and of the thigh. Proximal to the stifle a small nerve
the spermatic cord. Upon passing through the accompanies the descending genicular vessels
external inguinal ring it goes to the skin of the to the deep structures of the medial surface of
scrotum and prepuce in the male or to the in the stifle, and a cutaneous branch accompanies
guinal mammary gland and its covering skin in the medial genicular vessels to the skin supplied
the female. In both sexes the terminal branch of by this vessel. Distal to the stifle joint the saphe
the nerve runs caudolaterally and distally and nous nerve continues with the dorsal branch of
N. To c a u d a l p o rtio n of m .g lu te u s med. tP e lv ic s p la n c h n ic n.
Cronial g lu te a l n.{ , N. to m. le va to r am
1st sa cra l n , l s t coccygeal n.
7 th l u m b a r n ( j I / M. coccygeus
' / M. levator ani
'M . c a u d o a n a lis
Cutaneous branches
M. gluteus superf.
■Perineal n.
Pudendal n.
- Caudal g lu te a l n.
■O b t u r a t o r n.
y 11 i u m
, Is c h i a t i c s p i n e
✓S y m p h y s i s p e l v i s
Ext. i l i a c a.
O b t u r a t o r fo r a m e n
Deep fe m a r a l a N. to rn. o b t u r a t o r ext
F e m o r a l a. - _
P u d e n d o e p ig a s tric tru n k - -
J o in t c a p s u le -
C ra n ia l fe m o ra l a -
Nn. to m. a d d u c t o r lon gu s
A a .v n .ta j o i n t ' '
N.’. to m. p e c t i n e u s
O b tu ra to r b r /
Nn. to m. a d d u c t o r magnus et brevis
Med. c i r c u m f l e x f e m o r a l a . '
- Nn- to m. g r a c i l i s
R ig h t fe m u r'
F ic . 1 1 -2 4 . Nerves and arteries o f the right hip join t, medial aspect.
L um bar N erv es 609
Caud. g l u t e a l n .r a t M . g lu t e u s s u p e r f
B r a n c h f r o m S2 ' jCaudat p o r t i o n of m. g lu te u s med.
' >
N.to mm. o b t u r a t o r in te rn u s , ' i Lum bosacral tr u n k
•fe m e lli tr q u a d fe m o r is t ' ' /
/ N. to caud. p o r t i o n of m. g lu te u s med.
M. p i r i f o r m i s x ' ' \
,C ran g lu te a l a .trn
I s c h i a t i c n.\
M u s c u la r b ra nc h es
\ \
M.cjluteus medius
M.abd c r u r i s caud
At biceps feman
gluteus prof.
■M. t e n s e r
fasciae la ta e
Mm gem el 11 -M. s a r t o r iu s
Tendon of
m. o b tu r a to r int.
N.to m capsularis
M.c^uad. fe m o r is
M. a d d u c to r >Cran fe m oral a.
the saphenous vessels and supplies twigs to the The larger ventral branches (rami ventrales),
skin along its course to the paw. A small cuta after leaving the confines of the sacral canal, are
neous branch also follows the plantar branch of connected to the sympathetic trunk by single
the saphenous artery and vein and supplies the rami communicantes. They then continue on the
skin which overlies them. In the paw it supplies medial wall of the pelvis.
the skin of the dorsomedial part of the tarsus and The dorsal branches (rami dorsales) of the
metatarsus and ends in the skin over the second three sacral nerves leave the two dorsal sacral
and first digits when a first digit is present. foramina and the intervertebral foramen be
The obturator nerve (n. obturatorius) (Figs. tween the sacrum and the first coccygeal verte
11-23, 11-24) arises from the fourth, fifth, and bra. The dorsal branches are connected to each
sixth lumbar nerves (Langley and Anderson other by communicating strands forming a small
1896). The sixth root of origin is usually heaviest, dorsal sacral trunk or plexus. This trunk is usually
and the fourth smallest or even absent. The roots joined to the dorsal branches of the last lumbar
of origin do not arise close to the intervertebral and first coccygeal nerves. They decrease in size
foramina, but from the main nerve trunks (sci from first to third and like the dorsal branches of
atic for the sixth and the femoral for the fifth) the lumbar nerves are divided basically into
which are the main continuations of the fifth and medial muscular branches and lateral cutaneous
sixth lumbar nerves. The obturator nerve is branches. The short medial branches supply the
formed within the caudomedial portion of the lateral and medial dorsal sacrococcygeal mus
iliopsoas muscle. It leaves this muscle dorsome- cles. The lateral branches are longer. They run
dially and after crossing the ventrally lying com caudodorsolaterad between the dorsal lateral
mon iliac vein enters the subserosa of the pelvis. sacrococcygeal and dorsal intertransverse coc
After running obliquely caudoventrad across the cygeal muscles to reach the deep gluteal fascia
laterally lying shaft of the ilium it disappears through which they pass, usually accompanied
from view by first passing between the iliopubic by the corresponding dorsal sacral vessels. The
and ischial portions of the levator ani muscle. single lateral branch of the dorsal branch of the
The obturator nerve leaves the pelvis by passing first sacral nerve bifurcates upon passing through
through the cranial part of the obturator fora the gluteal fascia so that two cutaneous nerves
men where it lies in relation to the small obtu run caudoventrad on the superficial gluteal mus
rator ramus of the deep femoral artery which cle and down the thigh. The second and third
ascends through the opening. It sends branches sacral nerves are similarly disposed, supplying
into the external obturator muscle and continues bands of skin caudal to the first. These three
through the cranial part of the obturator fora nerves, the middle clunial nerves (nn. clunii
men. Thereafter it sends branches into the medii) of Ellenberger and Baum (1943), supply
pectineus, gracilis, and adductor muscles, inner a zone of skin which reaches to the middle of the
vating all the muscles which primarily adduct lateral surface of the thigh before ending in a
the pelvic limb. tapering cutaneous zone to anastomose with
each other and with the last two lumbar nerves
to form the sacral plexus.
SACRAL NERVES The sacral plexus (plexus sacralis) is formed by
the large interconnected ventral branches of the
The sacral nerves (nn. sacrales) (Figs. 11-21, last two lumbar nerves and the ventral branches
11-23) leave the three sacral segments of the of the three small sacral nerves which divide and
spinal cord by means of long dorsal and ventral redivide shortly after leaving the spinal canal,
roots, since these segments form that part of the with the resultant branches uniting with adja
conus medullaris which lies opposite the sixth cent nerves to form the sacral plexus. The last
lumbar vertebra. The roots merge to form the two lumbar nerves run caudoventrolaterad, con
sacral nerves within the sacral canal prior to verge, and blend under cover of the lateral ven
their intervertebral foramina of exit. Each of tral sacrococcygeal muscle on the pelvic surface
the first two sacral nerves then divides into dor of the sacrum medial to the sacroiliac joint. Two
sal and ventral branches which leave the sacrum slender branches leave the cranial surface of the
through the first two dorsal and pelvic sacral sixth nerve before it joins the seventh. The first
foramina, respectively. The third pair of sacral branch joins the fifth lumbar nerve and augments
nerves leaves the spinal canal by passing through the size of its continuation, the femoral nerve.
the intervertebral foramina between the sacrum The second branch unites with a like branch of
and first coccygeal vertebra like the other typical the fifth to form the obturator nerve.
spinal nerves. The lumbosacral trunk (truncus lumbosacralis)
Sacral N erves 611
(Fig. 11-25) is the largest and most impor superficial gluteal muscle. It may also send a
tant part of the lumbosacral plexus since it is branch to the middle gluteal muscle. In its course
continued outside the pelvis as the sciatic nerve. of about 2.5 cm. it lies between the sacrotuber-
It arises primarily from the sixth and seventh ous ligament medially, and the large caudal glu
lumbar nerves with a small contribution from teal artery and vein laterally. It is distributed to
the first and occasionally the second sacral the superficial gluteal muscle. It also supplies
nerves (Havelka 1928), and it has a root of origin the piriformis muscle by a delicate branch which
from the fifth lumbar, according to Ellenberger enters the proximal third of the muscle.
and Baum (1891). The lumbosacral trunk has The caudal cutaneous fem oral nerve (n.
two medium-sized branches, the cranial and cutaneus femoris caudalis) (Figs. 11-26, 11-27)
caudal gluteal nerves. The trunk lies in the pel is nearly as large as the pudendal nerve, to which
vic fascia while crossing the shaft of the ilium in it is united for most of its intrapelvic course. It
nearly a frontal plane where it is sandwiched in arises from the first and second sacral nerves and
the space between the origin of the ventral lat seldom from the third sacral (Havelka 1928).
eral sacrococcygeus muscle medially and the Bradley and Grahame (1959) state that it arises
thin levator ani muscle laterally. It is covered from the seventh lumbar and first sacral nerves
by peritoneum and is crossed obliquely on its with a possible addition from the sixth lumbar.
ventral aspect by the parietal branches of the in As it passes out of the pelvis dorsal to the ischial
ternal iliac vessels. It becomes the sciatic nerve arch it divides into perineal branches and the
after the last sacral branch enters it at the greater caudal clunial nerves. The perineal branches
ischiatic foramen. Each of the five constant, (rami perineales) accompany the perineal ves
spinal roots of the sacral plexus is usually united sels to the skin around the anus as several small
to the sympathetic trunk by a single ramus com- twigs. The caudal clunial nerves (nn. clunii cau-
municans. dales) stream out of the ischiorectal fossa in the
The cranial gluteal nerve (n. gluteus cranialis) fat covering the dorsal surface of the internal
arises from the lumbosacral trunk or its roots obturator muscle, usually as three nerves, one
mainly from the sixth and seventh lumbar nerves adjacent to the proximal part of the penis or
and from the first sacral nerves. It leaves the pel labia, one over the semitendinosus and semi
vis by passing immediately through the greater membranosus muscles, and one in the furrow
sciatic foramen and plunges into the muscles of between the biceps femoris and semimembrano
the rump. It is accompanied by the cranial glu sus muscles. These nerves run distally and supply
teal artery and vein. It circles craniad across the the skin of the proximal half of the caudal and
lateral aspect of the shaft of the ilium at the ori the adjacent medial and lateral surfaces of the
gin of the caudal bundle of the deep gluteal mus thigh.
cle. The cranial gluteal nerve continues cranio- The pudendal nerve (n. pudendus) (Fig. 11-
ventral between the middle and deep gluteal 23) usually arises from all three sacral nerves. A
muscles, usually perforates the cranial edge of prefixed origin of the pudendal by one segment
the deep gluteal, and terminates in the tensor is described by Havelka (1928). As the pudendal
fasciae latae muscle. It supplies the deep and nerve runs obliquely caudoventrad to the pelvic
middle gluteal and tensor fasciae latae muscles. outlet it lies lateral to the two coccygeal muscles
It has no cutaneous branches. and appears superficially medial to the superfi
The caudal gluteal nerve (n. gluteus caudalis) cial gluteal muscle. It lies dorsal to the accom
is a small nerve which may be double. It usually panying internal pudendal vessels. At the pelvic
arises from the cranial margin of the lumbosacral outlet the pudendal nerve gives rise to the caudal
trunk or from its seventh lumbar root. Occasion rectal and perineal nerves and the nerves to the
ally the cranial and caudal gluteal nerves arise in external genital organs; the dorsal nerve of the
common; at the other extreme the caudal gluteal penis of the male, and the nerve of the clitoris in
nerve may arise only from the first and second the female. The perineal nerve of both sexes
sacral nerves independently of the lumbosacral continues ventrocranially and ends as the caudal
trunk, as Bradley and Grahame (1959) illustrate. scrotal nerves of the male or the caudal labial
The caudal gluteal nerve runs parallel to the ven- nerves of the female.
trocranial border of the lumbosacral trunk on The caudal rectal nerve (n. rectalis caudalis)
the medial surface of the shaft of the ilium, and is a short nerve which leaves the pudendal at the
after passing through the greater sciatic foramen caudal border of the levator ani muscle and en
it crosses the caudal border of the piriformis mus ters the external anal sphincter muscle a little
cle or passes between the piriformis and middle below the middle. It is accompanied by the more
gluteal muscles to enter the medial surface of the deeply lying caudal rectal artery and vein. It,
612 Chapter 11. S p in a l N erves
S u p e r f lo t. c o c c y g e a l a.
, t C u ta n e o u s b r fr o m v e n t r a l br. o f S3
' /C u ta ne o u s br. f r o m v e n t r a l br. o f SI or2
M. co c cyg e u s / Caud p o r t i o n o f m j l u t e u s m e d
M. l e v a t o r a n i
M s p h in c te r am e x t- M. g l u t e u s s u p e rf .
P e rm e a l a rn ~ _ -C a u d . q lu te a l a m .
Caud. r e c t a l n . r a . -
~ P u d e n d a l n v mt. p u d e n d a l a.
Caud c u ta n e o u s f e m o r a l n
D o r s a l n o f c l i t o r is — -
" T o m o b t u r a t o r tnt. f r o m
is c h ia tic n
S a c r o t u b e r o u s Iiq .
b ic e p s fe m o r is
M. o b t u r a t o r i n t
■M. s e m i t e n d i n o s u s
M .c o n s tric to r v e s t i b u l i /
x 'M . s e m i m e m b r a n o s u s
M. c o n s t r i c t o r v u l v a e 1 M. i s c h i o u r e t h r a l i s
M. i s c h i o c a v e r n o s u s
'A t g r a c i I is
F ic . 11- 26. Nerves, arteries, and muscles of th e fem ale perineum , caudolateral aspect
Sacral N erves 613
M. c o c c y g e u s - Br. o f cran. g l u t e a l a.
M. levator ani -
C u ta n e o u s b r ~ - C a u d a l g l u t e a l a.
" - Cutaneous b r fr o m v e n tr a l
Caud r e c t a l n * a -
b r of SI
'P udendal n. *
D o r s a l a r n. o f p e n is ~ " int. p u d e n d a l a.
'•M. b ic e p s fe m o r is
M. r e t r a c t o r p e n is - ~
' To m. o b tu r a to r int. fr o m
M. b u lb o c a v e m o s u s - i s c h i a t i c n.
NM. o b t u r a t o r int.
C utan eo us b ra n c h e s
i
M. s e m ite n d in o s u s
M. is c h i o c a v e r n o s u s ' 1
M. i s c h i o u r e t h r a l is ' M. s e m im e m b r a n o s u s
and its companion nerve of the opposite side, glans along the dorsal surface of the os penis and
are apparently the sole supply to the external finally ends in the mucosa of the apex of the
sphincter muscle of the anus. Blakely (1957) glans. It is the chief sensory nerve to the penis,
found that unilateral severance of this nerve in mediating afferent impulses which result in or
perineal hernia operations could be performed gasm. The dorsal nerve o f the clitoris (n. dorsalis
without producing incontinence, but that bilat clitoridis) in the female is the homologue of the
eral severance of the caudal rectal nerve resulted dorsal nerve of the penis of the male and medi
in incontinence of feces. ates similar impulses. It is much smaller than the
The perineal nerves (nn. perinei) (Fig. 11-27) comparable nerve of the male and runs to the
are represented by several long twigs which sup ventral commissure of the vulva, where it ends
ply the skin and the mucous membrane in the in the clitoris.
region of the anus. The first branch to leave the The muscular branches of ventral sacral
pudendal nerve goes to the mucosa of the anal nerves (rami musculares) are usually two in num
canal and the skin at the anus. It either crosses ber. One supplies the levator ani and coccygeal
the external anal sphincter superficially or runs muscles, and the second, larger nerve innervates
deeply between the dorsal portion of the anal the rotators of the hip joint. Coming mainly from
sac and the internal sphincter muscle of the anus the second sacral nerve, but also receiving a
to reach the mucosa. The main portion of the small twig from the third, usually, is a small sin
perineal nerve leaves the pudendal near or in gle or double nerve which crosses the lateral
common with the nerve going to the mucosa and surface of the lateral ventral sacrococcygeal
sends several branches to the skin of the peri muscle. Its medial branch arborizes largely on
neum as it runs distally in the furrow located the medial, subperitoneal surface of the levator
between the proximal part of the penis and the ani muscle. Its lateral branch enters the cranial,
buttocks. One of these branches, larger than the medial surface of the much narrower, more cau
others, becomes related to the root of the penis dally located lateral coccygeal muscle. The mus
and as a branched nerve runs distally on the cular branch from the lumbosacral trunk to the
caudolateral surface of the proximal portion of muscles which rotate the hip joint was called the
the penis. The main perineal nerve, however, rotator nerve (n. rotatorius) by Schmaltz (1914).
continues distally and supplies the skin of the It is short. As it leaves the caudal margin of the
escutcheon with its fellow. The terminal portion lumbosacral trunk just prior to the passing of
of this nerve supplies the skin of the scrotum as this trunk through the caudal part of the greater
the caudal scrotal nerve (n. scrotalis caudalis). ischiatic foramen to be continued as the sciatic
In the female the comparable nerve supplies the nerve, a twig runs caudally and arborizes in the
skin of the labia as the caudal labial nerve (n. dorsal surface of the internal obturator muscle
labialis caudalis). The terminal portion of this as it lies on the ischiatic table caudomedial to the
nerve is not confined to innervating the skin of lesser ischiatic foramen. When the rotator nerve
the caudal portions of these external genital is double, the second branch leaves the caudal
parts, but continues cranial to them and is finally border of the lumbosacral trunk or its continua
dissipated in the skin of the inguinal region where tion, the sciatic nerve, about 1 cm. distal to
its area of skin supply is overlapped by the the origin of the branch to the internal obturator,
branches from the genital nerve which passes and curves around the caudal border of the deep
through the inguinal canal. gluteal muscle and sinks into the fascia between
The dorsal nerve o f the penis (n. dorsalis the deep gluteal and the cranial gemellus mus
penis) in the male is the main extrapelvic con cles. Sometimes the two parts of the rotator
tinuation of the pudendal nerve. It curves around nerve arise in common. After crossing the lateral
the pelvic outlet near the symphysis ischii where surface of the shaft of the ischium the second
it is separated from the opposite nerve by the branch supplies both parts of the gemelli and
paired artery and vein of the penis. At this place after passing ventral to the cranial gemellus ter
it issues a thin but long branch which inclines minates in the larger, obliquely running quad
medially where it comes to lie between the dor ratus femoris muscle.
sal vein and the tunica albuginea of the penis. It The sciatic or ischiatic nerve (n. ischiadicus)
usually anastomoses with the larger nerve just (Figs. 11-25, 11-28) is the largest nerve in the
caudal to the bulbus glandis. Upon reaching the body. It is a continuation of the lumbosacral
dorsal surface of the penis the dorsal nerve of the trunk. The division between the two is marked
penis runs cranially on the organ, sending by the second sacral nerve, contributing to this
branches to it, and then enters the caudal part of nerve complex. Since this contribution is lo
the bulbus glandis. It continues through the cated at the greater ischiatic foramen, the ex-
Sacral N erves 615
trapelvic part of the trunk is regarded as the branch then continues distally and divides into
sciatic nerve. It consists of two nerves, the tibial two or three branches. The more caudal branch
and fibular, which are so closely bound together, enters the middle portion of the semitendinosus
proximally, that they appear as one. The two muscle distal to the tendinous intersection which
parts, however, can be forcefully separated back partly divides the muscle. The more cranial part
to their origins from the spinal nerves. The nor of the nerve runs distally and again bifurcates;
mal division of the sciatic nerve is variable. Oc one branch goes to the caudal belly and the other
casionally it is located as far proximally as the to the cranial belly of the semimembranosus. A
hip joint, while at other times it may be as far long, slender, mixed nerve arises from the fibular
distally as the popliteal space. Upon leaving the portion of the sciatic nerve opposite or distal to
pelvis the sciatic nerve first lies on the gemelli the trochanteric fossa. After obliquely crossing
and the tendon of the internal obturator. As it the caudal surface of the tibial part it becomes
passes down the thigh it lies in succession on the associated with the medial border of the ab
quadratus femoris, adductor, and semimem ductor cruris caudalis muscle. In this region it
branosus muscles. It is covered first by the super sends a muscular branch to this small abductor.
ficial gluteal muscle and then by the biceps fem At about the middle of the thigh it leaves the
oris muscle and lies in close association with medial border of the abductor cruris caudalis
the small abductor cruris caudalis muscle, which muscle and obliquely crosses the caudal surface
crosses it obliquely in the proximal third of the of the muscle as it runs distally. It becomes sub
thigh. Its proximal portion is accompanied by cutaneous in the proximal part of the popliteal
the caudal gluteal vessels which lie caudal to it, region where it supplies a small area of skin.
and it is nourished by the small sciatic artery, The pelvic splanchnic nerves (nn. splanchnici
which is partly embedded in its caudolateral pelvini) (Figs. 11-21, 11-23) are the pelvic part
surface. In addition to the rotator nerve, which of the parasympathetic portion of the autonomic
is usually double and arises from the lumbosacral nervous system. They usually arise from the first
trunk, the true sciatic nerve which continues this and second sacral nerves. These two nerves may
trunk outside the pelvis has a single, stout, mus anastomose to form a single pelvic splanchnic
cular branch to the hamstring muscles, and a nerve or may run independently to the pelvic
muscular branch to the abductor cruris caudalis. plexus.
Other main branches of the sciatic nerve include The lateral cutaneous sural nerve (n. cutaneus
the lateral cutaneous sural nerve, caudal cutane surae lateralis) arises from the lateral surface of
ous sural nerve, and the terminal fibular and the fibular portion of the sciatic nerve at about
tibial nerves. the junction of the middle and distal thirds of the
The ramus muscularis of the sciatic nerve is thigh. After running a few centimeters dis
about 1 cm. in length and 3 mm. in diameter. It tally under the biceps femoris it enters the mus
leaves the caudomedial border or fibular portion cle between its smaller, deep head and its larger,
of the sciatic nerve opposite the space between more cranially lying, superficial head. The nerve
the middle gluteal and the cranial gemellus mus passes through the biceps femoris muscle with
cles. It lies cranial and parallel to the sacrotuber- out contributing to its supply and appears sub-
ous ligament and caudal gluteal artery and vein. cutaneously with cutaneous twigs of the caudal
After running about a centimeter in the furrow femoral vessels in the proximal, lateral portion of
(trochanteric fossa) medial to the trochanter the crus. It supplies most of the skin on the
major it sends a branch into the larger super lateral aspect of the crus as it ends near the
ficial portion of the biceps femoris muscle about tarsus.
3 cm. from its main origin on the tuber ischia- The caudal cutaneous sural nerve (n. cutaneus
dicum. The remaining portion of the nerve, surae caudalis) (Fig. 11-29) is known as the n.
which is approximately the same size as the suralis s. communicans tibialis s. cutan. fem. et
branch to the superficial portion of the biceps tibiae post. long, by Ellenberger and Baum
muscle, runs distally and at the distal border of (1891). It is called the n. cutaneus surae medialis
the quadratus femoris muscle branches into by Bradley and Grahame (1959). It resembles
usually four parts which arise variably. The most this nerve of man more closely than any other
caudal portion regularly bifurcates into a smaller, nerve, but the differences in location, area of
distolaterally running branch which supplies the supply, and anastomosis warrant the name given
smaller, deeper portion of the biceps femoris it. It is a long, slender nerve which arises from
muscle and by a shorter, larger branch which the caudal border of the tibial portion of the
innervates the proximal part of the semitendi sciatic nerve or from the tibial nerve directly,
nosus muscle. The main portion of the muscular usually about a centimeter distal to the origin
616 Chapter 11. S p in a l N e r v e s
Nn. L6 r L 7 ^ ^ C ra n ia l g lu te a l a * n .
B r f r o m S I N%v „ L u m b o s a c ra l tru n k
N. s'
to c a u d . p o r t i o n m .g lu t. med-~^\ "1 ' ^ - Bn f r o m SI ? S 2
Br. to ca ud . c u t f e m o r a l n. ' ' „ F e m o ra l a.
P a r i e t a l b r i n t i h o c a. _ - I s c h i a t i c n.
Caud. c / l u t e a l a.~._ _ C a u d a l g l u t e a l n-
S u p e r f . l a t c o c c y g e a l a—~
C r a n i a l f e m o r a l a.
B r to p u d e n d a l n. - •
A to c / l u t e a l mm. v j o i n t Bn to j o i n t c a p s u le
c a p s u le —
"L a t. c ir c u m fle x fe m o ra l
N. to mm. int. o b tu r a to r , -7
g e m e l l i v q uadratus f e m o r i s j o i n t c a p s u le
ir to j o i n t c a p s u l e
^ Rt fe m u r
M u s c u la r bnt ' O b t u r a t o r bn a f deep f e m o r a l a
|/).Ueu)50rJ
F ic . 11 28. \ erv es and arteries of the right hip join t, dorsal aspect.
Sacral N erves 617
_ - P o p lite a l a.
M. s e m ite n d in o s u s -
To m. g a s tro c n e m iu s , med. head - -I •- M- gastrocnem ius, lat. head
Caud. f e m o r a l a —
To m. flex, d ig i t o r u m s u p e rf . —
To mm.popliteus, flex, h a llu c is longus,
Deep f i b u l a r n.
t i b i a l i s post., fle x .d ig ito r u m longus
M. flex, d ig i t o r u m superf. -
- -M .f i b u l o r i s langus
M .g o stra cn e m iu s , lat. head - -
" ~Superf f i b u l a r n.
M. flex, hal lu c is longus- To mm. t i b i a l i s ant. v ext. digit, long.
M. ext. d ig ito r u m lat.- % Cron, t i b i a l a.
M. t i b i a l i s ant.
M . f i b u l a r i s brevis
of the lateral cutaneous sural nerve. It runs a crural fascia to become subcutaneous. At or just
short distance distally, bounded laterally by the proximal to the tarsus it becomes related to the
biceps femoris muscle and medially by the semi small, dorsal division of the saphenous artery,
membranosus muscle. Upon entering the pop which it follows into the dorsum of the paw. Like
liteal region it becomes associated with the the artery, it bifurcates on the dorsal surface of
plantar surface of the gastrocnemius muscle the proximal end of the paw into medial and
running distally on the muscle near the fusion lateral parts, the lateral branch being the larger.
of its two heads and in association with the The lateral branch divides again, and the medial
lateral saphenous vein. Throughout its course it branch may also redivide when a first digit is
sends branches to the skin of the caudal part of present. In this manner the superficial dorsal
the crus. Upon reaching the calcanean tendon it metatarsal nerves are formed.
usually divides into two branches of unequal The superficial dorsal metatarsal nerves II,
size. The smaller branch extends distally. Upon III, and IV (nn. metatarsi dorsales superficiales
reaching the tarsus it usually bifurcates into a II, III, et IV) (Fig. 11-31) run distally in the
small articular branch (ramus articularis) which superficial fascia and send many delicate
runs over the caudal part of the lateral malleolus branches to the skin of the dorsum and sides of
and supplies the lateral portion of the tarsal joint. the metatarsus. They lie over but not in the
The lateral calcaneal branch (ramus calcanei grooves between adjacent metatarsal bones
lateralis) crosses the distolateral surface of the where they are related to the corresponding
tuber calcanei and ends in the skin of this region. arteries and deep to the corresponding veins.
A twig may innervate the tibio-tarsal joint cap Upon approaching the metatarsophalangeal
sule. The larger branch of the caudal cutaneous junctions each nerve sends many rather large
sural nerve runs mediad between the calcanean cutaneous branches to the skin of the distal por
tendon and the tibia and joins the tibial nerve tion of the metatarsus and the dorsal proximal
1 to 3 cm. proximal to the tarsal canal. portions of the digits. Each of the three super
The common peroneal (fibular) nerve (n. ficial dorsal metatarsal nerves receives the much
peroneus [fibularis] communis) (Figs. 11-29, smaller deep dorsal metatarsal nerves at the
11-30) is the smaller of the two terminal distal ends of the intermetatarsal spaces to form
branches of the sciatic. It lies under the thin the several dorsal proper digital nerves.
terminal part of the deep portion of the biceps The deep peroneal (fibular) nerve (n. peroneus
femoris muscle. The nerve runs almost directly [fibularis] profundus) arises as the cranial termi
distad, obliquely crossing the lateral head of the nal branch of the sciatic nerve on the lateral head
gastrocnemius muscle. At the level of the stifle of the gastrocnemius muscle near its cranial
joint it sends an articular branch to the lateral border. It sinks in the proximal part of the crus
collateral ligament. Upon reaching the thin with the distally lying superficial peroneal nerve
lateral border of the flexor hallucis longus mus by passing between the deep digital flexor and
cle about 1.5 cm. distal to the stifle joint it dips lateral digital extensor muscles caudally and the
between this muscle and the lateral digital ex peroneus longus muscle cranially. From its first
tensor on the caudal side, and the peroneus 3 cm. there arise in succession usually four
longus which lies cranial to it and enters the branches. The most proximal branch is a mus
muscles of the cranial part of the crus. The com cular ramus (ramus muscularis) to the deep face,
mon peroneal nerve supplies a small branch to proximal end of the peroneus longus muscle. The
the peroneus longus muscle before dividing into next branch crosses under the long digital ex
superficial and deep peroneal nerves. tensor muscle and enters the medial border of
The superficial peroneal (fibular) nerve (n. the tibialis cranialis muscle. The third branch
peroneus [fibularis] superficialis) leaves the obliquely crosses the cranial surface of the
lateral portion of the parent nerve about 3 cm. delicate extensor hallucis longus muscle which
distal to the stifle joint where it lies in the inter- it supplies and becomes related to the lateral
muscular septum between the flexor hallucis surface of the cranial tibial artery. As it runs to
longus muscle caudally, and the peroneus the tarsus it passes between the artery and the
longus muscle cranially. As it extends distally it tibia, but remains closely united to the artery.
curves under the distal part of the belly of the In the proximal half of the tarsus the deep pero
peroneus longus muscle to enter the septum be neal nerve and cranial tibial artery he in the
tween this muscle and the long digital extensor. groove formed by the tendon of the long digital
At the beginning of the distal third of the crus it extensor, laterally, and the tendon of the cranial
becomes subfascial, and upon approaching the tibial muscle medially. At the tarsus the deep
flexor surface of the tarsus it perforates the peroneal nerve sends delicate branches to the
Sa c r a l N e r v e s 619
three heads of the extensor digitorum brevis portions run to the muscles which lie in the
muscle which lie on the flexor surface of the plantar part of the hindpaw and to the skin and
tarsus. The deep peroneal nerve then divides foot pads of the plantar surface of the hindpaw.
into medial and lateral parts. The muscular branches (rami musculares)
The deep dorsal metatarsal nerve II (n. meta- (Fig. 11-30) are numerous. The first two
tarsei dorsalis profundus II) (Fig. 11-31) is the branches arise in common closely bound to the
metatarsal continuation of the medial branch of cranial border of the caudal cutaneous sural
the deep peroneal nerve. The lateral branch in nerve, arising about 2 cm. before the tibial nerve
the proximal part of the metatarsus bifurcates enters the gastrocnemius muscle. One branch
to form the deep dorsal metatarsal nerves III enters the proximal portion of the caudal border
and IV (nn. metatarsei dorsales profundes III of the lateral head, while the other branch enters
et IV). The three deep dorsal metatarsal nerves the medial head of the gastrocnemius muscle at
anastomose with the corresponding superficial a like place. The muscular branch to the super
ones just proximal to the metatarsophalangeal ficial digital flexor enters the muscle at its origin.
joints to form the dorsal common digital nerves From the cranial border of the tibial nerve, as it
II, III, and IV. lies between the two heads of the gastrocnemius
The dorsal common digital nerves II, III, and muscle, arises the muscular branch which enters
IV (nn. digitales dorsales communes II, III, et the popliteus muscle at about the middle of its
IV) are formed in the distal ends of the three obliquely running distal border. Arising from this
main intermetatarsal spaces by the union of the nerve, or separately from the tibial, or from both
superficial and deep dorsal metatarsal nerves. of these, is the nerve or branches to the deep
They are about 1 cm. long, devoid of gross digital flexor muscle. When single, this nerve
branches, and end opposite or distal to the soon trifurcates into branches which supply its
metatarsophalangeal joints by dividing into two constituent bellies, the flexor hallucis longus
the medial and lateral dorsal proper digital and flexor digitorum longus muscles. A small
nerves II, III, and IV (nn. digitales dorsales twig goes to the tibialis caudalis muscle. After
proprii mediales et laterales II, III, et IV). Since these muscular branches arise, the tibial nerve
the common digital nerves lie between the then runs distad on the flexor hallucis longus
metatarsal bones, their terminal branches are muscle, where it is covered by the lateral head of
not confined to the supply of a single digit, but the gastrocnemius muscle. About 2 cm. proximal
are distributed to the apposed sides of adjacent to the tarsus it receives the caudal cutaneous
digits. For example, the third dorsal common sural nerve from the lateral side cranial to the
digital nerve gives origin to the medial dorsal Achilles tendon. At about the middle of the crus
proper digital nerve IV and the lateral dorsal near the medial surface, the tibial nerve comes
proper digital nerve III, whereas the second into relationship with the plantar branch of the
dorsal common digital nerve provides the proper saphenous artery and vein. About 1 cm. proximal
dorsal digital nerves to digits II and III and the to the tibiotarsal (ankle) joint the tibial nerve
fourth nerve to IV and V. The dorsal proper bifurcates into the medial and lateral plantar
digital nerves, as they run toward the distal ends nerves.
of the digits, lie ventral to the corresponding
arteries and send many branches to the skin of Nerves of th e H in d p a w ( P e s )
the dorsal and adjacent sides of the digit, finally
ending in the corium of the claw. The medial plantar nerve (n. plantaris medi
The tibial nerve (n. tibialis) (Fig. 11-29), alis) (Fig. 11-33) is smaller, more medial and
larger than the peroneal, is the more caudal superficial than the lateral plantar nerve. It
terminal branch of the sciatic nerve. It is about crosses the medial side of the tarsus in the super
5 mm. wide at its origin and is flattened trans ficial fascia outside the tarsal canal. At the proxi
versely as it lies between the caudal portions of mal end of the metatarsus it branches irregularly
the semimembranosis muscle medially and the into the superficial plantar metatarsal nerves II,
biceps femoris muscle laterally. It separates from III, and IV (nn. metatarsei plantares superficiales
the ischiatic nerve gradually in the proximal two- II, III, et IV) and the m edial plantar digital nerve
thirds of the thigh. It enters the crus between II (n. digitalis plantaris medialis II) or superficial
the two heads of the gastrocnemius muscle. The plantar metatarsal nerve I (n. metatarsus plan
tibial nerve supplies all the muscles which lie taris superficialis I) if a first digit is present. The
caudal to the tibia and fibula and sends branches continuation of this nerve is similar to the other
to the stifle, tarsal, and digital joints. Its terminal superficial plantar metatarsal nerves. These
620 Chapter 11. S p in a l N ehves
- Saphenous n.
- F e m o r a l a-
Fe m u r ~
- D e s c e n d in g g e n i c u l a r a.
— Branch to j o i n t
P a t e 11 a
— S a phenous a.
P a t e l l a r l i g ---- - C audal g e n ic u l a r a.
- T i b i a l col l a t e r a l l i g
-M e d ia l g e n ic u la r a
F ib u la
- D orsal b r, saphenous a.
T ib ia -
- P l a n t a r br, saphenous a
F em u r - _ Coudal fe m o r a l a.
F e m o r a l a. _ _ P o p l i t e a l a-
I s c h i a t i c n.~ \ „ to fe m u r
F i b u l a r n. — \ P a te lla
T ' b i o l n.- - Tendon of m. p o p lite u s
B r a n c h e s to j o i n t - -
C a u d c u t a n e o u s s u r a l n .-
N.to m . g a s t r o c n e m i u s med.- - J o i n t c a p s u le
N to m g a stro cn e m iu s lot— I \ _ -N .to m . f i b u l a r i s longus
N. to m. f l e x o r d ig s u p e r f - l j
__C au d a l t i b i a l a.
Caud. g e n i c u l a r a- _Superf. v deep f i b u l a r nn.
N. to mm. t i b i a l i s caud., popl iteus, '— C r a n ia l t i b i a l a.
v fle x, d i g i t o r u m longus
N to f i b u l a r c o l l a t e r a l l i N. to mm. t i b i a l i s cran. vext. digit- long,
Nn to m .fle x . h a l l u c i s Ion to m. ext. d i g i t o r u m lat.
N. to m. flex, d i g i t lo r m-ext. d ig i t o r u m longus
F i b u la
to m. ext. h a llu c is longus
N. to m. f i b u l a r i s b r e v is '
to m. ext. d ig i t o r u m longus
Fic. 11-30. A. Nerves and arteries of the right stifle joint, medial aspect.
B. Nerves and arteries of the right stifle joint, lateral aspect.
S acral N frves 621
Superf fib u la r n —
— Saphenous n.
Deep f i b u l a r n.— /'•
-A- Cran. ti b i a l a
Cran. t i b i a I a , superf. br.~
- ~v" “ Saphenous a, d o r s a l bn
B r of caud. c u t a n e o u s —
S u r a l n. Superf. bn of t i b i a l n.
(- - -Dorsal pedal a.
- - T r a n s v e r s e m e ta ta rs a l a.
Dorsal common
digital nn. IV, III, II
Med d o r s a l p ro p e r
d i g i t a l nn. V, IV. Ill Med. d o rs a l
~ d i g i t a l n .II
Lat. d o r s a l -
d i g i t a l n.V "Lat. d o rsa l p ro p e r
d i g i t a l nn. IV,I I I , I I
F ig . 1 1-31. Nerves and arteries of the right hindpaw, dorsal aspect. Inset of nerve supply to a double dewelaw
622 Chapter 11. S p in a l N erves
nerves extend obliquely distolaterad in the fascia muscle. When the special muscles of the first
covering the four branches of the superficial digit are developed, they receive their nerve
flexor tendon and send minute twigs to the supply from this source also.
quadratus plantae, lumbricales, and interflexorii The deep plantar metatarsal nerves I, II, III,
muscles. At or near the distal end of the meta and IV (nn. metatarsei plantares profundi I, II,
tarsus branches are given off their plantar sides III, et IV) are similar in location and termina
to the metatarsal foot pad. Their small continua tion to the comparable nerves of the forepaw.
tions anastomose with the deep plantar meta After running distad under the special muscles
tarsal nerves II, III, and IV to form the common of the second and fifth digits and on the interossei
plantar digital nerves. they anastomose with the superficial plantar
The lateral plantar nerve (n. plantaris lateralis) metatarsal nerves near the metatarsophalangeal
(Fig. 11-33) is larger, more lateral and deeper joints to form the common plantar digital nerves.
than the medial plantar nerve. Caudal to the Adjacent deep plantar metatarsal nerves anasto
distal part of the tarsus it enters the interval be mose, and cutaneous branches arise from their
tween the superficial and deep digital flexor distal ends and innervate the metatarsal pad.
tendons and obliquely runs distad between them The common plantar digital nerves II, III, and
a short distance before terminating as the deep IV (nn. digiti plantares communes II, III, et IV)
plantar metatarsal nerves II, III, and IV and are formed by the union of the superficial and
muscular branches. The lateral plantar nerve deep plantar metatarsal nerves in the distal ends
near its origin gives rise to the lateral plantar of the three main intermetatarsal spaces. After
digital nerve V (n. digitalis plantaris lateralis V), running 1 or 2 cm. each nerve bifurcates into two
which runs distad on the lateral surface of the plantar proper digital nerves (nn. digiti plantares
fifth metatarsal bone and digit and terminates proprii) which supply the skin of the apposed
by dorsal and plantar branches in the terminal and plantar surfaces of contiguous digits. Each
part of the fifth digit. The plantar branch goes nerve terminates as a dorsal branch which inner
to the fifth digital foot pad, and the dorsal branch vates the corium of the claw and the distal
goes to the corium of the claw. This nerve sends interphalangeal joint, and a plantar branch
twigs all along its course to the skin of the lateral which innervates a portion of the digital pad of
side of the paw. It issues a branch proximally its side.
which supplies the small quadratus plantae
muscle. COCCYGEAL NERVES
Usually upon entering the proximal part of
the metatarsus the lateral plantar nerve divides The paired coccygeal nerves (nn. coccygei)
into many branches. Three of these are the deep (Fig. 11-32) vary in number from four (Havelka
plantar metatarsal nerves II, III, and IV, and the 1928) to seven (Ellenberger and Baum 1891).
remaining branches go to the interosseous mus Hopkins (1935) found five pairs in each of nine
cles and the special muscles of the (first), second carefully dissected mongrel specimens. Like the
and fifth digits, and the quadratus plantae mus other spinal nerves, the coccygeal nerves branch
cle. The first muscular branch (ramus muscu immediately upon leaving their intervertebral
laris) arises from the lateral plantar nerve medial foramina into dorsal and ventral branches. Each
to the distal portion of the fibular tarsal bone; it pair of coccygeal nerves is numbered according
crosses the plantar surface of this bone and enters to the vertebra which precedes the interverte
the small spindle-shaped belly of the abductor bral foramen through which it runs. The dorsal
digiti V muscle. Another branch closely associ branch of the first coccygeal nerve is joined by
ated with it goes to the small transversely run the dorsal branch of the third or last sacral nerve,
ning quadratus plantae muscle. The remaining and the dorsal branches of the five coccygeal
muscular branches arise rather closely together nerves anastomose with each other. In this
at the proximal end of the metatarsus. The first manner the dorsal coccygeal trunk (tr. coccygei
branch leaves the lateral border of the lateral dorsalis) is formed. In a similar manner the
plantar nerve and enters the proximal end of ventral coccygeal trunk is formed. Baum and
the fifth interosseous muscle. A few millimeters Zietzschmann (1936) name these trunks the n.
distal to this origin the several nerves, one to collector caudae dorsalis and the n. collector
each of the interosseii (IV, III, and II), arise from caudae ventralis, respectively. Some precedence
the proximal portions of the deep metatarsal for the names, dorsal and ventral coccygeal
nerves. From the second or most medial of the trunks, is found in Sisson and Grossman’s text
deep plantar metatarsal nerves arises the branch (1952). Other authors have named the coccygeal
which supplies the abductor digiti secundi nerve trunks and their branches the coccygeal
C o ccygeal Nerves 623
M u s c u la r b ra nch es
Cutaneous branches
7 7 -7 / ,
Ventral coccygeal tr u n k
plexuses. They appear to be too highly organized nerves joining it. The trunk reaches its greatest
to warrant being called plexuses. The dorsal width of about 2 mm. as it is flattened against the
coccygeal trunk lies directly dorsal to the trans fibrocartilage located between the fifth and sixth
verse processes and the intertransverse muscles coccygeal vertebrae. The ventral sacrococcygeal
which extend only between several of the most muscles cover it ventrally. It lies ventral to the
proximal transverse processes. The dorsal sacro transverse processes and the intertransverse
coccygeal muscles lie directly dorsal to the nerve muscles. It is accompanied by the ventrolateral
trunk which is accompanied by the dorsolateral coccygeal artery; the two structures lie closely
coccygeal artery. The delicate muscular lateral to the hemal arches and more distally
branches (rami musculares) which leave it ex lateral to the hemal processes. The dorsal and
tend dorsocaudally into the dorsal lateral and ventral coccygeal trunks extend to the tip of the
medial sacrococcygeal muscles. In some seg tail. Muscular branches (rami musculares) leave
ments there are two muscular branches leaving both the dorsal and ventral surfaces of the trunk.
the trunk between the dorsal coccygeal nerves The delicate dorsal branches supply both parts
which contribute to its formation. In several dis of the intertransverse caudae muscle and the
sections not more than two of these branches vertebrae. The ventrally running branches inner
could be traced to terminations in the skin in any vate the lateral and medial ventral sacrococ
single specimen, and these were found near the cygeal muscles. The first seven to nine ventral
root of the tail. branches terminate in dissectable cutaneous
The ventral coccygeal trunk (tr. coccygeus branches (rami cutanei) which innervate the skin
ventralis) is larger than the dorsal coccygeal of the tail. Some of these fine nerves can be
trunk. It is united with the ventral branch of the traced several centimeters distally in the super
last sacral nerve at its beginning, and thereafter ficial fascia. Their main trunks lie in close associ
the trunk is augmented by the total volume of ation with the large, superficial lateral coccygeal
each of the ventral branches of the coccygeal vein and its small accompanying artery.
624 Chapter 11. S p in a l N erves
- T i b i a I n.
S u p e r f i c i a l br- o f t i b i a l n —
L a t p l a n t a r a. •
■Med. p l a n t a r n
Med. p l a n t a r a. - Lat p la n ta r n
T i b i a l n—
Med- p l a n t a r n.- Superf. p l a n t a r
L a t p l a n t a r n~ m e ta ta rs a l aa. -
Deep p la n ta r
m e ta ta rs a l aa.— Deep p l a n t a r m e t a t a r s a l
n n II. I l l I V
Superf. p l a n t a r m etatarsal
"nn.JJ.JIJ. IV
P l a n t a r common d i g i t a l
- nn. II, III. IV
•N n to m e t a t a r s a l footpad
Med. p l a n t a r Med. p la n ta r proper d ig ital
d ig ita l n il - -n n .III.IV , V
L o t p la n ta r p ro p e r- ~~ Lat. p la n t a r d ig i t a l n.V
d i g i t a l n n II, HI. IV
- N n t o d i g i t a l fo otpa d
F ig . 1 1 -3 3 . Nerves and arteries o f the right hindpaw plantar aspect. Inset of nerve supply to a double dewclaw
C o ccygeal N erves 625
If one remains aware of the artificiality of segre innervated. One neuron (preganglionic) of each
gating any portion of the nervous system from pair is located in the central nervous system and
the whole, a most useful concept can still be sends its myelinated axon out as part of the
gained by studying the autonomic nervous sys peripheral nervous system. These preganglionic
tem (systema nervosum autonomicum) as a more axons synapse with the second neuron (post
or less separate unit. Consideration must also be ganglionic) of the chain, and the nonmyelinated
given to interrelations between the autonomic axons of these postganglionic neurons end
and the somatic nervous systems. among or on glandular, cardiac muscle, or
The autonomic (general visceral efferent) sys smooth muscle cells. The postganglionic neu
tem may be defined as that portion of the rons ordinarily occur in clusters referred to as
nervous system concerned with the motor inner ganglia. The larger ganglia are found in specific
vation of smooth muscle, cardiac muscle, and locations and are named accordingly. Many
glands. Its counterpart with respect to somatic smaller, unnamed ganglia may be found scat
(derived from somites of embryo) innervation is tered along the paths of the autonomic nerves.
somatic efferent—the portion of the nervous It must further be emphasized that variability
system concerned with motor innervation of of details of autonomic nerve distribution is com
striated muscle. mon.
The foregoing definition of autonomic pur Autonomic ganglia are classified as vertebral,
posely excludes sensory innervation of the vis collateral (prevertebral), and terminal (periph
cera, since this is best classified separately as eral). Vertebral ganglia are more or less
visceral afferent. It should be noted here that segmentally distributed along the paired sym
this is not a universally accepted definition of pathetic chains which in turn lie along the ven
the word. Many authors prefer to include vis tral sides of the heads of the ribs. Collateral
ceral afferent structures under the term “auto ganglia are found more peripherally and in the
nomic.” For example, the Nomina Anatomica abdominal cavity are related to some of the
lists under autonomic nervous system the various larger arteries such as the celiac. Terminal
gross structures which contain both visceral ganglia are usually small and located in various
motor and visceral sensory nerves. Nevertheless body organs.
as a basis for rational consideration of function Both sympathetic and parasympathic divi
it is felt best to equate autonomic with general sions of the autonomic system have been shown
visceral efferent. References will be made to to be under direct control of the hypothalamus.
certain details of visceral afferent innervation For example, Beattie et al. (1930) showed that
when pertinent to the general picture. extrasystolic arrhythmia produced by chloro
On anatomical, pharmacological, and func form anesthesia in the cat could be prevented
tional bases the autonomic nervous system is either by the combination of sectioning sympa
further subdivided into sympathetic (pars sym- thetic nerves to the heart and removal of the
pathica) and parasympathetic (pars parasym- adrenal glands or by destruction of a portion of
pathica) portions. Both are so-called two-neuron the hypothalamus. These authors were also able
systems in the sense that two neuron chains link to trace hypothalamic efferent fibers as far as
the central nervous system to the structure the second lumbar segment of the spinal cord
626
P a r a s y m p a t h e t ic D iv is io n 627
(see also Katsuki et al. 1955). The hypothalamus glion ciliare) and leave the oculomotor nerve as
is regulated by certain areas of the cerebral the short root of the ciliary ganglion. Synapse
cortex (Uvnas 1954) and in turn reciprocally occurs in the ciliary ganglion, and the post
influences the cortex. Certain groups of neurons ganglionic fibers leave as the short ciliary nerves.
in the rostral hypothalamus have been shown to They supply the ciliary muscle which regulates
exert control over the parasympathetic system, lens curvature and the sphincter of the iris
whereas neurons in the caudal hypothalamus which, when activated, reduces pupillary di
influence sympathetic activity. Overlap of these ameter.
higher autonomic centers exists, and other parts
of the brain (Landau 1953; Lofving 1961) are Facial (VII) (Fig. 12-2)
also involved in control of autonomic activity.
Therefore it may be useful to think of impulses Small cells located in and about the motor
from several parts of the brain converging upon nucleus of the seventh cranial nerve send their
preganglionic neurons of the autonomic system preganglionic axons out with the pars intermedia
in a manner similar to the convergence of neural of the facial. These fibers then run in the major
influences upon lower motor neurons of the petrosal nerve (n. petrosus major) and nerve of
somatic nervous system. the pterygoid canal (n. canalis pterygoidei) to the
A good deal of autonomic activity is reflexly site of synapse in the pterygopalatine ganglion
regulated and is influenced by a wide variety of (ganglion pterygopalatinum). The nerve of the
sensory input. Schofield (1961) describes enteric pterygoid canal emerges through a small fora
neurons in the wall of the gut which may be men rostral to the foramen rotundum. The
afferent in function and therefore possibly play ganglion is described as lying medial to the
a part in the peristaltic reflex. Such an arrange sphenopalatine nerve on the pterygoid muscles.
ment could allow for reflex activity without Postganglionic distribution to the lacrimal gland
'mediation by the central nervous system. See is not well defined. Apparently some fibers may
also Bakeyeva (1957). Visceral afferent impulses go directly as small filaments, and some may run
play a large part in these reflexes, and most such with the lacrimal nerve. Other postganglionic
impulses do not give rise to sensations in human fibers go to glands and smooth muscle of the
beings. We should remain aware that no means nasal and oral cavities via branches of the fifth
exist for determining subjective sensations in cranial nerve (Jung et al. 1926).
animals, and so we sometimes resort to personal The rostral salivatory nucleus is located in the
experience and feelings for interpretation. dorsal part of the reticular formation dorsal to
the motor facial nucleus and medial to the
PARASYMPATHETIC DIVISION nucleus of the descending (spinal) root of the
trigeminal. These preganglionic axons leave the
Parasympathetic preganglionic axons leave brain stem with the pars intermedia of the facial.
the brain stem as part of cranial nerves III, VII, They join the main trunk of the facial nerve (Van
IX, and X (Figs. 12-1,12-2). The axons in nerves Buskirk 1945) and then leave as part of the
III, VII, and IX are distributed to the head re chorda tympani nerve (Foley 1945), which later
gion, whereas the vagus nerve distributes auto joins the lingual branch of the mandibular.
nomic fibers to the cervical, thoracic, and ab Synapse occurs in the submandibular ganglion
dominal viscera as far caudally as the left colic (ganglion submandibulare), and the postgangli
flexure. Parasympathetic preganglionic fibers onic axons go to the sublingual and submaxillary
also leave the spinal cord as part of the ventral salivary glands.
roots of the sacral nerves and become part of the
pelvic plexus. It is this brain stem and sacral Glossopharyngeal (IX) (Fig. 12-2)
spinal cord origin of parasympathetic fibers
which is described by the term “craniosacral” On a line caudal to the rostral salivatory nu
as a synonym for parasympathetic. cleus and also in the dorsal reticular formation is
found the caudal salivatory nucleus. The pre
Oculomotor (III) (Fig. 12-2) ganglionic axons leave with rootlets of the ninth
cranial nerve and run as part of its tympanic
Preganglionic neurons lie in the nucleus of branch (n. tympanicus), tympanic plexus (plexus
Edinger-Westphal, which is unpaired in the dog. tympanicus), and lesser petrosal nerve (n. petro
The axons run as part of the third cranial nerve sus minor). Synapse occurs in the otic ganglion
as far as the level of the ciliary ganglion (gan (ganglion oticum) near the origin of the mandib-
628 Chapter 12. T he A u t o n o m ic N ervous System
IPARASYMPATHETIC
.................................... • - - - I SY M PATH E T I C
I
G e n e r a liz e d
CRANIAL CERVICAL GANGLION
Lacrimal, Mandibular& Sublingual G.
-E y e
Parotid and M ucosal Glonds Salivary Glands
-S w e a t and Mucosal Glands
Sym pathetic
Blood Vessels
In n erva tio n
R E C U R R E N T LARYNGEAL N.
S T E L L A T E GANG
H ea rt and Lu ngs------
of
Sweat Glands, Blood
-SYMPATHETIC TRUNK
THORACIC (GREATER)
’SP LA N C H N IC N E R V E
CELIAC GANG,
r-rCRANIAL MESENT.G.
A bdom inal Viscera —
CELIAC MESENT.
PLE XU S Vessels
AORT/CORENAL
8 Erector
AND
GONADAL GANG.
CAUDAL MESENTERIC
GANG, and PLEXUS
PiU M u s c le s
P E L V IC G A N G L IA
F ig . 1 2 -1 . G eneral distribution of peripheral elem ents of sym pathetic and parasym pathetic divisions.
K
P a ro tid G landJ
Auriculotemporal Nerve
Sublinguol Gland —
— S p n m c te r of In c
Nerve of P terygoid
Canal
Fic. 12-2. Routes of distribution of preganglionic and postganglionic parasympathetic fibers in the bead region.
C O N T R IB U T IO N TO
NODOSE GANGLION\ PHARYNGOESOPHAGEAL N. CRANIAL LARYNGEAL N.
PHARYNGEAL BR VAGUS*
AURICULAR BR VAGUS
JUGULAR GANGLION
6LOSS OPHARYNGEAL N - M
with branch to carotid
YNGOESOPH. N
CRAN/AL CERVICAL GANGLII
HYPOGLOSSAL N —
SYMPATHETIC
OCCIPITAL A (C u t/ TRUNK
BR OF I I TO PHARYNGEAL PLEXi
RNAL BR. CRANIAL
RYNGEAL N
EXTERNAL MAXILLARY
UNGUAL A.
ular branch of the fifth nerve, and the post sible to peel the epineurial sheath away from the
ganglionic axons run with the auriculotemporal vagus and its ganglia, thereby clearly demon
branch of the trigeminal to their destination on strating the branches which leave the vagus. In
the gland cells of the parotid salivary gland. Ap some specimens a small, unnamed bundle of
parently the orbital salivary gland also receives vagal fibers may be seen to bypass the nodose
parasympathetic postganglionic fibers from the ganglion.
otic ganglion. Distal to the nodose ganglion the vagus joins
the sympathetic trunk, and the two are bound
in a common sheath along the entire cervical
Vagus (X ) (Figs. 12-1, 12-3, 12-4) region. The sympathetic trunk may retain a
rounded contour (in cross section) or may as
The vagus nerve contains a number of func sume a crescent shape where it is closely applied
tional components, one of which is the general to the vagus. In some specimens it is nevertheless
visceral efferent (parasympathetic) division. possible to separate the two easily by dissection.
Cells of origin of the preganglionic axons are At the level of the caudal cervical ganglion
found in the dorsal (motor) vagal nucleus. The (Fig. 12-4) the right recurrent laryngeal nerve (n.
paired nucleus is located in the dorsal part of the laryngeus recurrens) leaves the right vagus and
caudal medulla oblongata beneath and caudal passes dorsally around the caudal side of the sub
to the small eminences referred to as the alae clavian artery. As it runs anteriorly on the trachea
cineriae. Various branches of the vagus will be it lies deep to the carotid artery in the angle be
mentioned in this discussion, but distribution of tween the longus colli muscle and the trachea.
the branches other than autonomic is described Near its origin the right recurrent laryngeal
in the section on cranial nerves. The vagal root nerve gives off a fairly large branch, the recurrent
lets leave the brain along the dorsolateral sulcus cardiac nerve (right middle cardiosympathetic
of the medulla oblongata in series with the root nerve), which may receive contributions from
lets of cranial nerves IX and XI. Close associa the caudal cervical ganglion, the vagal trunk
tion of some of the vagal fibers with those of the (right cardiovagal nerve of Nonidez), and
accessory (XI) nerve for a very short distance the left recurrent laryngeal nerve. It is distrib
has led to the description of these vagal fibers as uted mainly to plexuses along the right and
the bulbar or internal root of the accessory nerve left coronary arteries. The nerve contains
in some species. Chase and Ranson (1914) con postganglionic sympathetic, preganglionic para
cluded that a true bulbar root of the eleventh sympathetic, and visceral afferent fibers for
cranial nerve does not exist in the dog, a fact the heart.
that can be demonstrated by careful dissection. Nonidez (1939) has described parasympa
A few millimeters distal to the origin of the thetic neurons of the terminal ganglia of the
vagus is located the small superior (jugular) gan heart as differing markedly in size. As a conse
glion (ganglion superius). It contains unipolar quence the postganglionic axons also vary in di
sensory type neurons whose dendrites are dis ameter. These axons lie in the walls of the atria
tributed with the auricular branch (r. auricularis) and the interatrial septum and form extensive
of the vagus. This branch leaves approximately plexuses. Smaller branches leave the plexuses
at the level of the superior ganglion and has been and end as ring-shaped, club-shaped, or reticu
shown to be distributed in part via branches of lated enlargements, which contact the surface
the seventh cranial nerve. of the cardiac muscle fibers. Similar enlarge
The pharyngeal 'branch (r. pharyngeus) of the ments may also occur along the course of the
vagus is given off between the superior and the finer branchings. Terminations are especially
more distally lying inferior (nodose) ganglion abundant among the specialized muscle fibers of
(ganglion inferius). The latter ganglion is also the sino-atrial and atrioventricular node. Post
composed mainly of unipolar sensory type neu ganglionic parasympathetic axons of the cardiac
rons which have their peripheral distribution in ganglia are also distributed via nerve plexuses
the viscera. along branches of the coronary arteries. These
The cranial laryngeal nerve (n. laryngeus su axons form plexuses in the adventitia and finally
perior) leaves the vagus at the level of the in end in relation to the smooth muscle cells of the
ferior ganglion. The cranial cervical ganglion outer media. Primarily these fibers innervate the
(sympathetic) is located medial to the nodose arteries of the atrium and proximal part of the
ganglion. The epineurial sheaths of the two ventricles. Most of the parasympathetic distri
ganglia may be separate or may show variable bution is to structures above the coronary sulcus,
degrees of fusion. By careful dissection it is pos whereas sympathetic fibers innervate the ven-
P a r a s y m p a t h e t ic D iv is io n 631
\ '
<SPINAL GANGLION
VENTRAL ROOT
COMBINED VERTEBRAL N. and
N To LONGUS C O L L ! MUSCLE
VERTEBRAL ARTERY
N To OMOCERVICAL A.
VAGOSYMPATHETIC TRUNK
CAUDAL CERVICAL GANGLION
L T RECURRENT LARYNGEAL N
R T RECURRENT LARYNGEAL N
'ESOPHAGUS
Fic. 12-4. Dissection of thoracic region, right side. The right brachiocephalic (innominate) nerve(s), the sympathetic contribu
tion to the phrenic nerve, and several of the arterial plexuses have been omitted.
632 Chapter 12. T he A u t o n o m ic N ervous System
tricles. See Holmes (1956, 1957) and Tcheng tral branches. The ventral branch of the right
(1950, 1951) for further information on intrinsic fuses with its left counterpart to form the ventral
innervation of the heart. vagal (ventral esophageal) trunk. A similar anas
The right vagus nerve, having usually sepa tomosis occurs between right and left dorsal
rated from the sympathetic trunk a short dis vagal branches more distally to form the dorsal
tance cranial to the level of the caudal cervical vagal (dorsal esophageal) trunk.
ganglion, runs ventral to the subclavian artery Both dorsal and ventral vagal trunks supply
and continues caudally along the lateral surface branches to the esophagus before passing
of the trachea. At the level of the subclavian ar through the diaphragm at the esophageal hiatus.
tery the ansa subclavia may be intimately at Upon reaching the abdominal cavity the ventral
tached to the vagus for a short distance. Also in vagus becomes plexiform for a short distance.
this region the vagus and recurrent laryngeal Branches from this plexus supply mainly the
nerves may receive branches from the caudal liver and stomach. The hepatic branches (usu
cervical ganglion or ansa subclavia. Distal to the ally two or three) run in the lesser omentum to
origin of the recurrent nerve the vagus gives off the liver. Chiu (1943) describes three superficial
two or more fine branches called the cranial and hepatic branches, two of which come from the
caudal cardiovagal nerves. The cranial cardio- ventral vagus and one from the dorsal. These
vagal nerves go mainly to the pretracheal plexus. run in the lesser omentum and pass between the
The caudal cardiovagal nerves distribute to the caudate and left lateral lobe of the liver to form
dorsal wall of the right atrium. Electrical stimula a simple plexus just rostral to the porta hepatis.
tion of the right vagus nerve at various levels indi Chiu also describes two more plexuses in the
cates that most of the inhibitory fibers going to course of distribution of these fibers. One of the
the right atrium run in the cardiovagal nerves. plexuses receives contributions from the celiac
Most of the fibers in these nerves are nonmyeli plexus. Autonomic nerve branches are distrib
nated, but nevertheless are considered to be uted from the plexuses to the cystic duct, gall
preganglionic parasympathetic (numerous ex bladder, left lateral lobe of the liver and com
ceptions exist to the “rule” of preganglionic axons mon bile duct. Fiber groups are also described
being myelinated and postganglionic axons un as passing along the right gastric and cranial
myelinated). Daly and Hebb (1952) have pro pancreaticoduodenal arteries to the pancreas
vided evidence that the cervical sympathetic and along the right gastric artery to the pylorus.
trunk of the dog may contain cardio-inhibitor, A small filament may go directly to the duode
bronchoconstrictor and pulmonary vasomotor num.
(vasopressor) fibers. Such fibers could possibly be The second group of fibers (three or four gas
collaterals of ascending preganglionic sympa tric branches) from the ventral vagus supplies
thetic fibers or postganglionic fibers which origi the ventral surface of the stomach. Mizeres
nate in the cranial cervical ganglion. Further (1955a) mentions the presence of a number of
more, the apparent presence of cardio-acceler- other filaments which join the sympathetic
ator fibers in the vagus has been demonstrated plexus on the branches of the left gastric artery.
by Shizume (1952) and others. Pannier (1946) The dorsal vagal trunk supplies the cardiac
placed the origin of these fibers in the medulla region of the stomach and then forms a plexus
oblongata. on its dorsal surface. Distribution is mainly to
The main trunk of the right vagus continues the lesser curvature and pyloric regions of the
caudad dorsal to the root of the lung. At this stomach. Stimulation experiments in unanesthe
level it gives off several prominent branches tized decerebrate dogs (Okabe 1958) indicate
along the bronchi. Parasympathetic ganglia are that the vagus nerves carry motor fibers to the
found in the lung, and postganglionic parasym cardiac sphincter and that sympathetic fibers in
pathetic fibers have been traced to smooth mus the greater splanchnic nerve mediate inhibition
cle and glandular structures. The lung is also of cardiac sphincter motility (Okabe 1959).
richly innervated with a wide variety of receptor Branches of the dorsal vagal trunk may also join
structures as far distally as the alveoli. Receptors the celiac, left gastric, hepatic, and cranial mes
have been described in smooth muscle, tracheal enteric nerve plexuses associated with the cor
and bronchial epithelium, respiratory bronchi responding arteries. Mizeres (1955a) has also
oles, alveolar ducts, and alveolar walls (Elftmann described filaments to the hepatic plexus.
1943). The vagus supplies branches directly to Information on vagal parasympathetic distri
the trachea and esophagus and also via the re bution caudal to the stomach is sparse, but there
current laryngeal nerve. seems to be general agreement that these fibers
Immediately caudal to the root of the lung reach as far caudally as the left colic flexure.
both right and left vagi split into dorsal and ven
S y m p a t h e t ic D iv is io n 633
Branching and distribution of the left vagus these cardio-inhibitory nerves are nonmyeli
are similar to the right as far distally as the level nated.
of the caudal cervical ganglion. Here a branch Cranial to the root of the lung the left vagus
leaves the vagus to join a branch from the caudal may contribute a variable number of filaments
cervical ganglion, resulting in formation of the to the cardiosympathetic nerves as previously
left brachiocephalic (innominate) nerve (appar noted. Several branches leave the left vagus for
ently left cardiovagal nerve of Nonidez). This is distribution along the bronchi much as occurs
distributed to the base of the brachiocephalic on the right side. Distal to the root of the lung
artery and ventral surface of the aortic arch the left vagus divides into dorsal and ventral
(Mizeres 1955a). A significant component of this branches each of which joins its counterpart of
nerve is afferent (depressor). the right side to result in dorsal and ventral vagal
It must be emphasized that details of auto trunks. Their distribution has been discussed.
nomic nerve distribution to the heart are ex Both right and left thoracic vagus have been
tremely variable, particularly with respect to shown to contain a bundle of sympathetic fibers
origin of fibers. Many of the nerves to the heart near the periphery of the main trunk.
are mixed in the sense that they carry sympa Parasympathetic postganglionic neurons may
thetic, parasympathetic, and visceral afferent be found scattered in the walls of structures
fibers. Terminology is varied and often confus which they supply or else as part of well-defined
ing. In general it can be said that impulses prop plexuses in similar regions. Most prominent of
agated in sympathetic fibers tend to accelerate these are the myenteric and submucous plexuses
heart rate and force of contraction, whereas of the digestive tract (Lawrentjew 1931; Filo-
parasympathetic stimulation reduces heart rate. gamo 1950; Richardson 1960; Schofield 1961).
Filaments leave the left recurrent laryngeal The myenteric (Auerbach’s) plexus (El’bert
nerve near its origin and go to the left atrium or 1956; Learning and Cauna 1961) (plexus myen-
plexuses which supply it. Other filaments to the tericus) is located between the longitudinal and
left atrium leave the vagus distal to the origin of circular muscle layers, and the submucous
the recurrent nerve. The vagus also contributes (Meissner’s) plexus (plexus submucosus) lies in
fibers to certain other cardiac nerves which will the submucosa of the digestive tube. Prominent
be discussed with the sympathetic system. ganglionated plexuses are also found in the atria
Fibers which make up the left recurrent laryn of the heart. Much of their regulating effect is
geal nerve leave the vagus at the level of the aor thought to be mediated by way of the sino-atrial
tic arch and loop around the arch distal to the and atrio-ventricular nodes.
ligamentum arteriosum. The recurrent laryngeal Cell bodies for the sacral parasympathetics
nerve then proceeds anteriorly along the left are found in the sacral spinal cord, and the pre
ventrolateral surface of the trachea adjacent to ganglionic axons leave as part of the ventral roots
the ventromedial side of the esophagus. of the sacral segments. These fibers join more
Branches are supplied to the trachea and esoph distally as the pelvic nerve (nervus erigens), lo
agus as the nerve passes to its termination in the cated on the lateral wall of the distal portion of
larynx (see Hwang et al. 1948 for more detail on the rectum. On both sides the pelvic nerve ex
innervation of the esophagus). Gross dissection pands into a plexus which also receives the hypo
indicates that in the cervical region the left re gastric (sympathetic) nerves. One or more gan
current laryngeal supplies more branches to the glia (presumably parasympathetic) are located
esophagus than it does to the trachea. The oppo within the plexus (Jayle 1935). Distribution of
site is true of the right recurrent laryngeal nerve. postganglionic autonomic fibers is via branches
In dogs whose esophagus lies to the right of the and plexuses to the pelvic viscera and reproduc
midline in the cervical region there is a more tive organs. Parasympathetic fibers from the
even distribution of branches of right and left sacral region have been traced through the
recurrent laryngeal nerves to both trachea and length of the large intestine (Schmidt 1933).
esophagus. Iljina and Lawrentjew (1932a, 1932b) have de
Electrical stimulation experiments conducted scribed parasympathetic ganglia in the wall of
on the left vagus of the dog indicate that cardio- the rectum and urinary bladder.
inhibitory fibers leave the left recurrent laryn
geal nerve as it passes around the arch of the SYMPATHETIC DIVISION
aorta and go to the left atrial region. Other in
hibitory fibers leave the left vagus distal to the Sympathetic preganglionic axons take origin
origin of the recurrent nerve and are also dis from small cells in the intermediolateral cell col
tributed to the left atrium. Most of the fibers in umn of spinal cord segments T1 through L4 or
634 Chapter 12. T he A u t o n o m ic N ervous System
L5. In some instances preganglionic fibers may 3. Fibers which leave vertebral ganglia such
originate as far caudally as the sixth lumbar seg as the stellate to go directly to a visceral struc
ment (Mehler et al. 1952). The myelinated small- ture such as the heart via a cardiac nerve.
diameter axons leave in the ventral roots of the 4. Fibers which leave vertebral ganglia such
above-named segments and for a short distance as the cranial cervical for distribution as plex
are part of the corresponding spinal nerves. uses along arteries of the head region.
These fibers then leave each spinal nerve as one 5. Fibers which leave collateral ganglia such
or more communicating rami which attach to as the celiac for distribution as plexuses along
the sympathetic trunk. The trunk is a paired arteries to the abdominal viscera.
strand of preganglionic and postganglionic sym Other modifications of distribution route may
pathetic nerve fibers located ventrolateral to the occur.
bodies of the vertebrae in the thoracic and lum Reference is sometimes made to gray versus
bar regions. Continuations of the two trunks white rami communicantes, the gray being
caudally into the sacral and cranial coccygeal made up mainly of postganglionic fibers and the
region may be partially fused and lie ventral to white mainly of preganglionic. At most segmen
the bodies of the vertebrae. Both sympathetic tal levels of the thoracic and upper lumbar re
trunks are continued anteriorly into the cervical gions of the dog the rami are mixed in the sense
region where each runs in a common sheath that they contain both preganglionic and post
with the vagus nerve. Each trunk terminates in ganglionic fibers. Rostral and caudal to the tho
the cranial cervical ganglion. With certain ex racic and upper lumbar levels the communicat
ceptions sympathetic chain ganglia are located ing rami are made up almost entirely of non
at segmental intervals along the sympathetic myelinated fibers, since no preganglionic
trunk. contributions to the sympathetic trunk arise
The ganglia are numbered according to the from the cervical, lower lumbar, and sacral re
spinal nerve to which the postganglionic fibers gions. The comparatively few myelinated fibers
attach. The part of the sympathetic trunk which which may be present in these “gray” rami are
joins adjacent ganglia is the interganglionic (in- probably visceral afferent.
ternodal) segment. These ganglia contain the The small size of the sympathetic filaments
multipolar neurons which give rise to the which are grossly dissectable belies their exten
predominantly nonmyelinated postganglionic sive distribution. In general it appears that sym
axons. Synapses may occur, for example, be pathetic postganglionic fibers are as widely dis
tween preganglionic fibers of T 12 ramus com tributed as the arterial system.
municans and cells in T12 sympathetic ganglion, Both microscopic and macroscopic ganglia
or the preganglionic fibers may proceed rostrad may be found more or less randomly scattered
or caudad in the sympathetic trunk to synapse in along the peripheral portions of the sympathetic
ganglia at other segmental levels. In general, system. These are for the most part unnamed
preganglionic fibers from T I to T5 are directed ganglia either proximal or distal to ganglia such
forward in the sympathetic trunk, whereas cau as celiac, cranial mesenteric, or those of the ver
dal to T5 the fibers go caudally. Thus pregan tebral chain. For this reason any given segment
glionic fibers from a given segmental level of the of sympathetic nerve cannot be considered to
spinal cord may be distributed to vertebral carry all preganglionic or all postganglionic fi
ganglia at many different levels. bers. Microscopic examination of a sympathetic
Preganglionic fibers may also leave the sym nerve could not be expected to identify com
pathetic trunk (as for example in the splanchnic pletely the nature of the fibers, since not all pre
nerves) and synapse in collateral ganglia such as ganglionic fibers are universally myelinated
the celiac. throughout their length. Postganglionic fibers
Postganglionic fibers may be distributed in a usually have no myelin sheath, but this also is
number of different ways: probably not a constant characteristic. Appar
1. Fibers which leave the vertebral ganglia ently, sympathetic fibers acquire myelin sheaths
via the communicating rami to return to the after birth, whereas myelinization of vagus fibers
spinal nerve at the same level. These are distrib begins before birth (Diamare and deMennato
uted to smooth muscle (vasomotor and pilomo 1930).
tor fibers) and glands by way of the spinal nerve. S y m p a t h e t ic D is t r ib u t io n to H ead and
2. Fibers which run either rostrally or cau N eck
dally in the sympathetic trunk and join the adja
cent spinal nerves via their respective rami. Preganglionic fibers from the upper thoracic
S y m p a t h e t ic D iv is io n 635
region run as part of the vagosympathetic trunk and plexuses along arteries. The vascular nerve
to their site of synapse in the cranial cervical plexuses lie in the tunica media and tunica ad
ganglion (ganglion cervicale superius). The cer ventitia. Cunliffe (1961) has found no definite
vical sympathetic trunk of the dog has been histological evidence of a secretomotor nerve
shown to contain approximately 5000 to 13,000 supply to the thyroid and suggests that rate of
fibers, of which about half appear to be myeli secretion and removal of the secretory product
nated (Foley and DuBois 1940). The cranial cer are indirectly controlled by rate of blood flow.
vical ganglion lies deep to the tympanic bulla Essentially the same conclusions were reached
and the proximal portions of cranial nerves IX, by Nonidez (1935), who also demonstrated an
X, XI, and XII. A large portion of the postgan afferent nerve supply of the thyroid.
glionic fibers leaving the ganglion continue as
plexuses along the arteries of the head region. Sym p a t h e t ic D is t r ib u t io n in T h o r a c ic
For example, rather prominent bundles of fibers R e g io n
can be seen going to both external and internal
carotid arteries. Sympathetic postganglionic Fusion of vertebral ganglia in the anterior
fibers also follow the common carotid artery. In thoracic and caudal cervical regions has resulted
general the arterially distributed sympathetic in formation of the stellate and caudal cervical
fibers supply sweat and salivary glands and ganglia. The structure referred to as the caudal
smooth muscle such as that of the arterial walls, cervical ganglion in the human is a part of the
nictitating membrane, dilator pupillae, and the stellate ganglion in the dog. Therefore the caudal
erector pili. Those sympathetic nerves which cervical ganglion of the dog is comparable to the
follow the arteries may be fairly discrete bundles middle cervical ganglion of the human. The
or plexiform (see Billingsley and Ranson 1918a stellate (cervicothoracic) is the largest autonomic
and 1918b for distribution in the cat). ganglion in the dog and is located on the lat
Filaments from the cranial cervical ganglion eral surface of the longus colli muscle at the level
may in some specimens be seen to join the ninth, of the first intercostal space. It receives mixed
tenth, eleventh, and twelfth cranial nerves, the rami from T l, T2, and T3 spinal nerves and
connection to the tenth being most common. A sometimes from T4. Preganglionic fibers ending
fairly constant branch to the first cervical nerve in the stellate ganglion may originate as far back
occurs (Fig. 12-3), and this may supply filaments as T5 spinal cord segment. Gray rami connect to
to the second and third cervical nerves also. C 8 and C7 spinal nerves. Some of the rami at
Other branches may join the pharyngeal and tached to the stellate ganglion may be double.
cranial laryngeal branches of the vagus. In some The ramus to C7 may be fused with the vertebral
dogs one may demonstrate a pharyngeal plexus nerve (n. vertebralis), and both may be fused
formed by contributions from cranial nerves IX with a third nerve, the branch of the seventh
and X and the cranial cervical ganglion, whereas cervical nerve to the longus colli muscle. For this
in other specimens there is little intermingling reason the vertebral nerve may appear decep
of these fibers. Filaments also attach the cranial tively large. Actually, the vertebral nerve is com
cervical ganglion to the region of the carotid paratively small. Supposedly the nerve plexus
sinus and carotid body (glomus). A larger branch which runs along the vertebral artery supplies
(Fig. 12-3) leaves the caudal end of the cranial gray rami to spinal nerves anterior to C7 as far as
cervical ganglion, contributes to the pharyngeal C3. These connections are difficult to demon
plexus, and sends fibers (thyroid nerve of Noni strate. In some instances the vertebral nerve
dez) along the thyroid artery to the gland. The (Fukuyama and Yabuki 1958) leaves the stellate
rest of the nerve continues caudally in the con ganglion as a separate entity. The nerve enters
nective tissue membrane which joins the vago the transverse foramen of the sixth cervical ver
sympathetic trunk to the common carotid artery. tebra along with the vertebral artery and usually
(Intermingling of several functional types of continues as a plexus along the artery.
fibers and variations in branching preclude the Cranioventrad to the stellate ganglion the
positive identification of some nerves in this area sympathetic trunk splits to pass around the sub
by other than physiological means.) It is joined clavian artery as the ansa subclavia. One or both
by small branches of the cranial laryngeal and sides of this loop may be double.
glossopharyngeal nerves and continues caudally At the junction of the vagosympathetic trunk
as the pharyngoesophageal nerve. The thyroid with the ansa subclavia medial to the subclavian
gland receives sympathetic postganglionic fibers artery lies the caudal cervical ganglion of the
which are distributed as interfollicular nerves sympathetic division (Fig. 12-4). It lies in the
636 Chapter 12. T he A u t o n o m ic N ervous Sy st e m
receive contributions from the ansa subclavia, Plexuses on the smaller arteries have already
stellate ganglion and/or vagal trunk. been discussed.
L eft B rachiocephalic (I nnominate
N erves (L eft C ardiovagal ). These nerves orig Pressoreceptor and Chemoreceptor
inate from the caudal cervical ganglion and Afferents
vagus. They go to the base of the brachiocephalic
A rise in blood pressure activates pressorecep
artery and ventral surface of the aortic arch.
tors located at a number of different sites in the
Pressoreceptor afferents are carried in these
body which are primarily related to the larger
nerves in addition to autonomic fibers.
vessels. As a sequel to increased pressoreceptor
activity there follows a reflex fall in blood pres
Other Sympathetic Branches in the sure. (Lofving [1961] has delineated a cortical
Thoracic Region depressor area [sympatho-inhibitory] in the ros
Prominent branches can be seen to follow tral part of the cingulate gyrus and a pressor
each of the arteries just rostral to the heart, such area [sympatho-excitatory] in the subcallosal
as axillary, omocervical, costocervical. These region. These experiments were done on the
may be referred to as plexuses which follow the cat.) Pressoreceptor cells are known to be located
respective arteries and are named accordingly. in the wall of the carotid sinus (innervated by
Contributions to these branches are mainly from the carotid sinus branch of cranial nerve IX) at
the ansa subclavia and caudal cervical ganglia the bifurcation of the internal and external ca
with occasional filaments from the stellate gan rotid arteries and also in the walls of the large
glion. This ganglion may also supply a branch to vessels as they enter and leave the heart. Afferent
the phrenic nerve. fibers (aortic depressor fibers) which carry im
Filaments from the rostral portions of the tho pulses from the latter group of receptors run
racic sympathetic trunk going directly to the rostrally with the vagus nerve and enter the
heart have been reported, but apparently are medulla as part of the rootlets of the vagus. Cell
not constant. Small ganglia may be located along bodies of these afferent fibers are in the nodose
the paths of these nerves. Mizeres (1955a) has ganglion. Nonidez (1937) has described in con
described the autonomic plexuses of the thoracic siderable detail the location of the pressorecep
region, and the following is based on his work. tors and also of the group of chemoreceptor cells
The pretracheal plexus is found ventral to the (aortic glomi or aortic bodies) in this area. The
tracheal bifurcation and dorsal to the right pul epithelioid bodies themselves are further de
monary artery. Contributions to this plexus are scribed as being composed of one type of cell
from the left recurrent laryngeal nerve, ventro with granular cytoplasm and a round or oval
medial cervical cardiac nerve, recurrent cardiac nucleus. Some are closely packed, while others
nerve, dorsal cervical cardiac nerve, and cranial form irregular strands separated by capillaries.
cardiovagal nerves. This plexus supplies the dor These receptor regions are abundantly supplied
sal atrial walls and continues as the right and left by nerves composed of mostly medium and small
coronary plexuses along the coronary arteries. diameter fibers. Some fibers form “baskets”
The latter plexuses receive the main part of the around the epithelioid cells. Finer branches end
recurrent cardiac nerve. Distribution is to the as rings or club-shaped enlargements on the cells.
ventricular walls and the ascending aorta. The Garner and Duncan (1958) have reported on
dorsal ventricular plexus lies on the dorsal wall electron microscopic studies of the structure of
of the left atrium and left ventricle. The fibers, the carotid body (glomus) in the cat.
which apparently supply the ventricular wall,
come mainly from the ventrolateral cervical Routes of Sympathetic Fibers
cardiac nerve and also from the left recurrent to the Heart
laryngeal and the left stellate cardiac nerves.
The right pulmonary plexus receives branches Effects of stimulating sympathetic fibers to
from the vagus and stellate cardiac nerves. The the heart are generally those of increased rate
left pulmonary plexus is supplied by the vagus (accelerator effect) and increased force of con
and stellate cardiac nerves. It also receives fila traction (augmentor effect). In stimulation ex
ments from the dorsal ventricular plexus. The periments described by Mizeres (1958) the
pulmonary plexuses carry autonomic and sen greatest acceleratory effect was usually ob
sory fibers from the parenchyma and vasculature tained from T2 and T3 levels of the sympathetic
of the lung. outflow on the right side. Stimulation at the same
638 Chapter 12. T he A u t o n o m ic N ervous S ystem
level on the left side was usually without effect Lumbar splanchnic nerves may arise from
on heart rate. An augmentor effect, however, each of the lumbar sympathetic segmental levels,
could always be obtained by stimulating the those from the first five being most constant.
appropriate sympathetic rami on the left side. They are named according to the level from
Thus most of the accelerator fibers come from which they arise. Most originate as grossly dis-
the right communicating rami (T1 to T4), pass sectable single filaments, but some may be
through both limbs of the ansa subclavia, and double, and anastomotic branches may join
proceed via the right stellate cardiac nerves to adjacent nerves. Origin may be from a lumbar
the right atrial wall. Alternate routes appear to ganglion or from interganglionic segments of the
be via the cranial and caudal cardiovagal nerves. sympathetic trunk. As noted earlier, the first
Augmentor fibers, which take origin mainly lumbar splanchnic nerve may have connections
via the left upper thoracic rami communicantes, with the thoracic splanchnic nerve. In general
probably run mainly in the ventrolateral cervical the first four lumbar splanchnic nerves distribute
cardiac nerve. to one or more of the following: aorticorenal,
cranial mesenteric, and gonadal ganglia; inter-
mesenteric, renal, and gonadal plexuses. The
S y m p a th e tic D i s t r i b u t io n in t h e
fifth through seventh lumbar splanchnic nerves
A b d o m in a l R e g i o n (Fig. 12-5) go to the caudal mesenteric ganglion. Filaments
also go to the postcaval vein and iliac arteries.
Branches of the thoracic and abdominal sym Correspondingly named plexuses of variable
pathetic trunk supply preganglionic fibers to the complexity are found along the arteries of the
abdominal and pelvic regions. In addition, these abdominal region. Those at the root of the celiac
branches also carry a few postganglionic fibers and cranial mesenteric arteries form a dense mat
and a considerable number of visceral afferent or sheath called the celiacomesenteric plexus.
fibers. As is true of other areas of autonomic This is continuous with the plexuses which are
nerve distribution in the body, the specific distributed with the branches of these two
nerve branchings in this region may vary con arteries. Interwoven in the celiacomesenteric
siderably from one animal to the next. plexus are the paired celiac and unpaired cranial
The thoracic (greater) splanchnic nerve leaves mesenteric ganglia.
the thoracic sympathetic trunk approximately Small sympathetic ganglia containing approx
at the level of the thirteenth thoracic ganglion. imately twenty-five to 100 neurons have been
The nerve is larger in diameter than the continu demonstrated along the course of the larger
ation of the sympathetic trunk. Both pass under arteries of the abdominal cavity. Since these lie
the lumbocostal arch lateral to the crus of the distal to the large ganglia such as celiac and
diaphragm and enter the abdominal cavity. cranial mesenteric, it follows that the nerve
Other branches designated as lesser splanchnic plexuses along the arteries can be expected to
nerves may leave the sympathetic trunk just contain some sympathetic preganglionic fibers
caudal to the greater splanchnic nerve. The in addition to the preponderance of sympathetic
thoracic splanchnic nerve supplies filaments to postganglionics. These plexuses also include
the thoracic aorta and the adrenal gland. It parasympathetic preganglionic and visceral
terminates mainly as several branches to the afferent fibers.
celiacomesenteric plexus. The adrenal ganglia The cellular makeup of the adrenal medulla
may also be supplied by this nerve. The lesser and the chemical mediators involved indicates
splanchnic nerves, if present, distribute to the that this portion of the adrenal glands is much
same general area. Filaments may sometimes be like a sympathetic ganglion. The medulla is
found joining the greater splanchnic to the first richly innervated by preganglionic sympathetic
lumbar splanchnic nerve. fibers. This is in contrast to the sparsely inner
Immediately after the thoracic splanchnic vated adrenal cortex (Teitlebaum 1942; Arikh-
nerve branches off, the lumbar sympathetic bayev 1957; Wilkinson 1961).
trunk is small, but becomes larger as it proceeds Filaments which follow the external and in
caudally. Ganglia may be present for each of the ternal iliac arteries apparently may arise from
seven lumbar segments, or variable fusion of the more caudally located lumbar splanchnic
adjacent ganglia may occur. Preganglionic con nerves, the hypogastric nerves, and as continua
tributions to the lumbar sympathetic trunk ap tions of the aortic plexus.
parently do not occur caudal to L 6 . In most The right and left hypogastric nerves repre
instances the caudal limits of the preganglionic sent largely postganglionic connections between
contributions are L4 or L5. the caudal mesenteric ganglion and the pelvic
S y m p a t h e t ic D iv is io n 639
Branches to A o rta
Dorsal Gastric N.
To Phrenic N.
Thoracic S p la n ch n ic N.
Lt. G a s t r i c A.
Common H e p a t ic A. Splenic A.
Celiac A. A drenal 6 . & P lexus
Rt. C e liac G. Lt. C e lia c G.
~ L u m bar Splanchnic N. I.
Cranial M esenteric A.
Lt. P h re n ic o-A b d o m in al A.
Splanch nic G.
Cranial M e s e n t e r i c G. A o rtic o re n al G.
L um bar S p l a n c h n i c N. n .
Renal G.
Gonadal G.
G o n adal P lexus
Rt. Gonodal A.-
^ T ~ L . 3Z.
-------—----------Lum bar Splanchnic N. 3L
Caudal M esenteric G.
Caudal M esenteric A.
Deep C i r c u m f le x Iliac A.
E xternal Ilia c A.
Internal Ilia c A.
M edian S a c r a l A.
Celiacomesenteric Celiac and cranial mesenteric Dorsal vagus, thoracic See structures supplied
splanchnic nerves by branches of celiac
and cranial mesenteric
arteries
Splenic .................................. Left gastroepiploic Left celiac ganglion Spleen, pancreas, greater
Continuation of splenic artery curvature of stomach
to spleen and pancreas
Phrenicoabdominal ............. Phrenic (may arise directly Celiac ganglion Diaphragm, part of ab
from aorta) Adrenal ganglia dominal wall
Branches to adrenal gland and Adrenal gland
abdominal wall
Daly, M. deB., and C. Hebb. 1952. Pulmonary vasomotor li Ingersoll, E. H., and L. L. Jones. 1958. Effect upon the urinary
bers in the cervical vagosympathetic nerve of the dog. bladder of unilateral stimulation of hypogastric nerves in
Quart. J. exp. Physiol. 37: 19-44. the dog. Anat. Rec. 130: 605-616.
Derom, E. 1945. Experimental researches on the vasomotor Jayle, G. E. 1935. Le centre hypogastrique du chien. Arch.
innervation of the dog’s front paw. Bull. Acad. roy. Med. Anat. (Strasbourg) 19: 357-367.
Belg. 10: 427-459. Jung, L., R. Tagand, and F. Chavanne. 1926. Sur l’innervation
Diamare, V., and Mario de Mennato. 1930. Contributo all’ana- excito-secretoire de la muqueuse nasale. C. R. Soc. Biol.
tomia ed alio sviluppo del sistema nervoso simpatico. Atti (Paris) 95: 835-837.
R. Accad. Sci. Fis. e. Mat. (Naples) 18: 1-115. Katsuki, S., T. Okajima, and T. Matsumoto. 1955. Experimen
El’bert, M. E. 1956. On the problem of the cyto-architecture tal studies on the autonomic function of the preoptic area;
of Auerbach’s plexus of the small intestines in the cat and influences of electrical stimulation of the preoptic area
dog. Tr. Mosk. Vet. Akad. 1956(18): 35-38. upon blood pressure, and movement of pelvic organs
Elftman, A. G. 1943. The afferent and parasympathetic inner (urinary bladder, rectum, and uterus). Kumamoto med. J.
vation of the lungs and trachea of the dog. Amer. J. Anat. 8: 180-186.
72: 1-23. Kuntz, A., and G. Saccomanno. 1944. Sympathetic innerva
Erez, B. M. 1955. The problem of the extracardiac nerve sys tion of the detrusor muscle. J. Urol. (Baltimore) 51: 535-
tem in dogs. Uch. Zap. Tadzh. Inst. 1955(6): 145-153. 542.
Filogamo, G. 1950. Ricerche sul plesso mienterico. Arch. ital. Kuru, M., T. Kurati, and Y. Koyama. 1959. The bulbar vesico-
Anat. Embriol. 54: 401-412. constrictor center and the bulbo-sacral connections aris
Foley, J. O. 1945. The sensory and motor axons of the chorda ing from it; a stucly of the function of the lateral reticulo
tyrnpani. Proc. Soc. exp. Biol. (N.Y.) 60: 262-267. spinal tract. J. comp. Neurol. 113: 365-388.
Foley, J. O., and F. S. DuBois. 1940. A quantitative and experi Landau, W. 1953. Autonomic responses mediated via the cor
mental study of the cervical sympathetic trunk. J. comp. ticospinal tract. J. Neurophysiol. 16: 299-311.
Neurol. 72: 587-601. Lawrentjew, B. J. 1931. Zur Lehre von der Cytoarchitektonik
Folkow, B. 1955. Nervous control of blood vessels. Physiol. des peripherischen autonomen Nervensystems; die Cyto
Rev. 35: 629-663. architektonik der Ganglien des Verdauungskanals beim
Fukuyama, U., and M. Yabuki. 1958. On the vertebral nerve Hunde. Z. mikr.-anat. Forsch. 23: 527-551.
and the communicating rami connecting with the inferior Learning, D. B., and N. Cauna. 1961. A qualitative and quan
cervical ganglion in the dog. Fukushima J. med. Sci. 5: titative study of the myenteric plexus of the small intes
63-88. tine of the cat. J. Anat. (Lond.) 95:160-169.
Garner, C. M., and D. Duncan. 1958. Observations on the fine Lindgren, P., A. Rosen, P. Strandberg, and B. Uvnas. 1956.
structure of the carotid body. Anat. Rec. 130: 691-710. The sympathetic vasodilator outflow; a new cortico-spinal
Greenberg, S. R. 1954. Sympathetic components of the vagus autonomic pathway. J. comp. Neurol. 105: 95-109.
cardiac nerves of the dog. Anat. Rec. 118: 304-305. Lofving, B. 1961. Cardiovascular adjustments induced from
--------------- 1956. A fiber analysis of the vagus cardiac rami the rostral cingulate gyrus. Acta physiol, scand. 53 (Suppl.
and the cervical sympathetic nerves in the dog. J. comp. 184): 1-82.
Neurol. 104: 33-48. Mehler, W. R., J. C. Fischer, and W. F. Alexander. 1952. The
Hassin, G. B. 1929. The nerve supply of the cerebral blood ves anatomy and variations of the lumbosacral sympathetic
sels; a histologic study. Arch. Neurol. Psychiat. (Chic.) 22: trunk in the dog. Anat. Rec. 113: 421-435.
375-391. Miyake, T. 1958. Effects of intestinal distention upon the
Hayasi, T. 1959. On the effects of distension of the urinary movements of the colon. J. Physiol. Soc. Japan 20: 744-
bladder upon the colon. J. Physiol. Soc. Japan 21: 380- 751.
385. Mizeres, N. J. 1955a. The anatomy of the autonomic nervous
Hillarp, N. A. 1959. The construction and functional organ system in the dog. Amer. J. Anat. 96; 285-318.
ization of the autonomic innervation apparatus. Acta --------------- 1955b. Isolation of the cardioinhibitory branches
physiol, scand. 46 (Suppl. 157): 1-38. of the right vagus nerve in the dog. Anat. Rec. 123: 437-
Holmes, R. L. 1956. Further observations on the nerve endings 446.
in the adult dog heart. J. Anat. (Lond.) 90; 600. _----- -------- 1957. The course of the left cardioinhibitory fibers
--------------- 1957. Structures in the atrial endocardium of the in the dog. Anat. Rec. 127: 109-115.
dog which stain with methylene blue, and the effects of --------------- 1958. The origin and course of the cardioacceler-
unilateral vagotomy. J. Anat. (Lond.) 91: 259-266. ator fibers in the dog. Anat. Rec. 132: 261-279.
Hwang, K., M. I. Grossman, and A. C. Joy. 1948. Nervous con Nonidez, J. F. 1935. Innervation of the thyroid gland; Distri
trol of the cervical portion of the esophagus. Amer. J. bution and termination of the nerve fibers in the dog.
Physiol. 154: 343-357. Amer. J. Anat. 57: 135-170.
Iljina, W. J., and B. J. Lawrentjew. 1932a. Zur Lehre von der --------------- 1937. Distribution of the aortic nerve fibers and
Cytoarchitektonik des peripherischen autonomen Ner- the epithelioid bodies (supracardial “paraganglia”) in the
vensystems; Ganglien des Rektums und ihre Beziehungen dog. Anat. Rec. 69: 299-313.
zu dem sakralen Parasympathikus. Z. mikr.-anat. Forsch. --------------- 1939. Studies on the innervation of the heart; dis
30: 530-542. tribution of the cardiac nerves with special reference to
--------------- 1932b. Experimentell-morphologische Studien the identification of the sympathetic and parasympa
iiber den feineren Bau des autonomen Nervensystems; thetic postganglionics. Amer. J. Anat. 65: 361-407.
Uber die Innervation der Harnblase. Z. mikr.-anat. Okabe, Y. 1958. The vagus innervation of the cardiac sphinc
Forsch. 30: 543-550. ter. J. Physiol. Soc. Japan 20: 752-763.
6 44 Chapter 12. T he A u t o n o m ic N ervous System
--------------- 1959. The splanchnic innervation of the cardiac Shizume, K. 1952. The cardio-accelerator fibers in the vagus
sphincter. J. Physiol. Soc. Japan 21: 43-49. nerve. Nippon J. Angio-cardiol. 16: 8-14.
Pannier, R. 1946. Contribution a 1’innervation sympathique Tcheng, K. T. 1950. Etude histologique de Tinnervation car-
du coeur; les nerfs cardioaccelerateurs. Arch. int. diaque chez le chien. C. R. Soc. Biol. (Paris) 144: 882-883.
Pharmacodyn. 73: 193-259. --------------- 1951. Innervation of the dog’s heart. Amer. Heart
Polley, E. H. 1955. The innervation of blood vessels in striated J. 41: 512-524.
muscle and skin. J. comp. Neurol. 103: 253-268. Teitlebaum, H. 1942. The innervation of the adrenal gland.
Richardson, K. C. 1960. Studies on the structure of autonomic Quart. Rev. Biol. 17: 135.
nerves in the small intestine, correlating the silver im Uvnas, B. 1954. Sympathetic vasodilator outflow. Physiol. Rev.
pregnated image in light microscopy with the perman 34: 608-618.
ganate fixed ultrastructure in electron microscopy. J. Van Buskirk, C. 1945. The seventh nerve complex. J. comp.
Anat. (Lond.) 94: 457-472. Neurol. 82: 303-330.
Schmidt, C. A. 1933. Distribution of vagus and sacral nerves Wilkinson, I. M. S. 1961. The intrinsic innervation of the supra
to the large intestine. Proc. Soc. exp. Biol. (N.Y.) 30: 739- renal gland. Acta Anat. (Basel) 46: 127-134.
740.
Schofield, G. C. 1961. Experimental studies on the innervation
of the mucous membrane of the gut. Brain 83: 490-514.
CHAPTER 13
TH E DIGESTIVE SYSTEM
A N D ABDOM EN
The digestive system (apparatus digestorius), also establishes free communication between
or systema digestorium) consists of the diges the two parts of the oral cavity. When the
tive tube and accessory organs. The primary mouth is open the vestibule is largely effaced
parts of the digestive tube are the mouth, phar and the oral cavity proper is greatly enlarged.
ynx, esophagus, stomach, small intestine, large The parotid and zygomatic salivary ducts
intestine, and anus. That portion caudal to the open into the dorsoposterior part of the vesti
pharynx, including all structures from the esoph bule. The parotid duct opens through the cheek
agus to the anus, is termed the alimentary canal on the small parotid papilla (papilla parotidea),
(canalis alimentarius). The accessory organs in located opposite the posterior part of the upper
clude the teeth, tongue, salivary glands, liver, fourth premolar tooth, approximately 5 mm.
gall bladder, and pancreas. from the fornix of the vestibule, which is formed
The wall of the digestive tube is richly sup by a reflection of the mucosa from the cheek
plied with secretory epithelium and intrinsic to the gum. The main duct of the zygomatic
glands. It is lined throughout by a mucous gland opens lateral to the posterior part of the
membrane (tunica mucosa) which is continuous upper first molar tooth on a small papilla near
with the surface integument at the mouth and the vestibular fornix. A small ridge connects the
anus. main zygomatic and parotid duct openings.
Usually one to four small accessory ducts from
THE MOUTH AND ASSOCIATED the zygomatic gland open posterior to the main
STRUCTURES duct. The submucous glands are few and are
confined to the lower lip and the adjacent part
Mouth of the cheek. The secretion of these glands is
discharged through about 10 openings located
The mouth (os), in a restricted sense, includes opposite the four lower premolar teeth near the
only the opening between the lips. In a broad fornix of the vestibule.
sense it designates the oral cavity (cavum oris), The oral cavity proper (cavum oris pro-
in which are located the teeth and tongue. The prium) is bounded dorsally by the hard palate
oral cavity is divided into the vestibule and the and a small part of the adjacent soft palate;
oral cavity proper. laterally and anteriorly the dental arches and
The vestibule of the mouth (vestibulum oris) teeth form its boundary; the tongue and the
is the space external to the teeth and gums and reflected mucosa under it form the floor of this
internal to the lips and cheeks. It opens to the cavity. When the mouth is closed the tongue
outside anteriorly by means of the U-shaped nearly fills the oral cavity proper. The sublin
slit, the oral fissure (rima oris), between the lips. gual and mandibular ducts open under the body
This opening is in essentially a frontal plane of the tongue, on the inconspicuous sublingual
through the anterior margins of the orbits. caruncle (caruncula sublingualis).
When the mouth is closed the vestibule com Extending posteriorly from the caruncle to
municates with the oral cavity proper by means about a transverse plane through the lower
of the interdental spaces, which vary greatly in shearing teeth is a low ridge of mucosa about 2
size. A space on either side, posterior to the last mm. wide and 1 mm. high. This is the sub
cheek tooth, and nearly 1 cm. long in large dogs, lingual fo ld (plica sublingualis). It lies close to
645
646 Chapter 13. T he D ig e s t iv e S ystem and A bdo m en
the body of the mandible and is formed by the The wide orbicularis oris muscle and the inser
underlying mandibular and sublingual ducts tions of several other facial muscles form the
and the anterior part of the polystomatic por media of the lips.
tion of the sublingual gland.
Just posterior to the upper central incisor Cheeks
teeth is the incisive papilla (papilla incisiva), a
rounded eminence which extends posteriorly The cheeks (buccae) form the posterior por
to blend with the first transverse ridge formed tion of the lateral walls of the vestibular cavity.
of the mucosa covering the hard palate. On each The cheeks are small in the dog because of the
side of this papilla the incisive canal, or naso unusually long posterior extension of the com
palatine duct (ductus nasopalatinus), opens. missures of the lips. The distance between the
This duct leaves the oral cavity by a slit-like commissures and the masseter muscles, forming
opening and extends posterodorsally for 1 or 2 the posterior limits of the laterally expandable
cm. through the palatine fissure to open into cheeks, averages only 3 cm. The cavity of the
the floor of the nasal fossa. The oral cavity is cheeks, however, runs medial to the masseter
continuous posteriorly with the isthmus of the muscles and extends as far posteriorly as the at
fauces and with the oral pharynx. tachment of the buccinator muscles on the
mandible and maxilla at their alveolar borders
Lips opposite the last two cheek teeth of each jaw
and the intervening anterior border of the basal
The lips (labia oris) form the anterior and half of the coronoid process. In rodents and
most of the lateral boundaries of the vestibule. most herbivores the cheeks are large and serve
Upper and lower lips (labia maxillaria et mandib- as storage space, especially during mastication
ularia) are recognized, which meet at acute and transportation. This function is of minor
angles posteriorly, forming the commissures o f importance in the dog.
the lip (commissurae labiorum). The lips bound The morphology of the cheeks is closely re
the oral fissure, the external opening of the oral lated to that of the lips, with which they are
cavity. Their margins are narrow and devoid continuous. The lips and the cheeks consist of
of hair except the anterior two-thirds of the three basic layers. The external layer is the
upper lip on either side. Toward the commis hairy integument; the middle layer consists of
sure, on either side, the posterior third progres muscle and fibroelastic tissue; and the inner
sively increases in width, to form a rounded layer consists of the mucosa (tunica mucosa).
border measuring as much as 1 cm. The margin Projecting posterolaterally from the posterior
of the lower lip as far posteriorly as a level part of the skin of the cheek are usually two
through the canine teeth is devoid of hair over coarse vibrissae, 3 to 5 cm. long. These are in
a zone about 5 mm. wide. Posterior to the addition to the vibrissae of the upper lip and
canine teeth this smooth zone increases to 1 cm. muzzle. The middle layer of the cheek consists
in width, and the narrow border becomes ser primarily of the buccinator muscle, although
rated by the formation of about 15 conical lateral to this are the m. zygomaticus and certain
papillae several millimeters high. fasciculi of the cutaneous fascia. The dorsal buc
No definite frenulum, or median mucosal cal glands in carnivores are consolidated to form
fold, attaches the lower lip to the gum, and the the zygomatic gland, a large mixed salivary
median mucosal fold of the upper lip is poorly gland located in the ventral part of the orbit.
developed, being thick but narrow. The mucosa The ventral buccal glands consist of a few small,
of the lower lip is firmly attached to the gum on solitary glands located in the submucosa, an
either side in the space between the canine and terior to the masseter muscle and medial to the
the first premolar tooth (interdental space). A fibers of the ventral part of the buccinator. The
deep, straight, narrow cleft, the philtrum, marks mucosa of the lips and cheeks is thinly corni-
the union of the two halves of the upper lip, fied stratified squamous epithelium which, in
anteriorly. The hair of both lips slopes postero- some breeds, is partly or wholly pigmented.
ventrally. It is thinner and shorter in front, Although the lips are not used as prehensile
longer and thicker farther back. A few of these organs, the various muscles in them provide
are tactile hairs, or vibrissae. On the upper lip movement. Dogs express affection or rage
and adjacent dorsal part of the muzzle the largely by the movements of their lips and
vibrissae are imperfectly arranged in four rows. cheeks.
T he M o u th and A s s o c ia t e d Stru ctures 647
I n c i s i v e p d p i Ila~
O r i f i c e of i n c i s i v e c a n a l
Hard p a la te
/ B u c c i n a t o r m.
/
Orifice o f p a r o t i d / / S t y l o g l o s s u s m.
/ / / M y l o h y o i d e u s m.
O r i f i c e s of
ZLjCjomai i c d u c t s | \ E p i h i j o i d bone
S oft p a la te
R i d y e f o r me d by m a s s e t e r m - ^
I'XVI' ' M\
/
/
^ M a n d i b u l a r a l v e o l a r ruj.tv.
.Masseter m
G i o s s o p a l a t i ne a r c h - 4 -
^ Hypoglossal
Pa la tin e tonsil
^ ^ P h a r y n y e a l mm.
F a c i a l v. ^Diyastricus m
Pharynyopalatine a r c h ___ ^ ^ S t y l o h y o i d e u s m.
^ l^ h a r y n y e a l isthmus — _ - M a n d i bulor/ln.
l \ - ~ T h y roic^/^artilage
Me d . t f e t r o p h a r y n c j e a l In
Dorsal^walI o f
laryncjeal pharunx
- - - E x t . m a x i l l a r y v.
A n n u l a r fold
^ ___ P a r a t h y r o i d j l a n d
Ext . j u g u l a r v.
H
- T h y r o i d cjland
Esophacj < .--------- jr--------S t e r n o c e p h a l i c u s rn.
------ Common c a r o t i d a.
O — \— lnf j u3 ular v-
N, V
F ig . 1 3 -1 . Frontal section o f head and neck through the digestive tube, ventral aspect.
T he M o u th and A s s o c ia t e d Stru ctures 649
extrinsic muscles, the levator and tensor veli Lying directly dorsal to the palatine aponeu
palatini, blend with the palatine aponeurosis. rosis are the small mixed dorsal palatine glands
The right and left pterygopharyngeal muscles (Ellenberger 1911). The epithelium which
pass lateral to the posterior part of the soft pal covers them, and on which their numerous
ate from origins on the pterygoid bones. Right ducts open, is of the pseudostratified ciliated
and left palatopharyngeal muscles arise near the columnar type. It is continued forward on the
median plane from the palatine aponeurosis. dorsum of the hard palate to line the nasal
They sweep laterally and upward in the phar pharynx and the nasopharyngeal duct. Poste
ynx, forming the bases for the palatopharyngeal riorly, before reaching the posterior border of
arches. the soft palate, it is continued by stratified squa
Vessels of the palate. The right and left mous epithelium which is the epithelium of the
major palatine arteries (aa. palatinae majores) whole ventral surface of the palate.
are the main arteries to the hard palate. The
main arteries to the soft palate are the minor Teeth
palatine arteries (aa. palatinae minores), aided
by the ascending pharyngeal (a. pharyngea The teeth (dentes) (Fig. 13-2) are highly spe
ascendens) and the major palatine. They are cialized structures which serve as weapons of
dwarfed by the palatine plexus of veins of the offense and defense, as well as for the procur
soft palate, which lies mainly lateral to the two ing, cutting, and crushing of food. Each tooth
slender palatine muscles. The main part of the is divided into three parts. The crown (corona
palatine plexus is located in the deep part of the dentis) is the exposed part, the part which pro
glandular mantle of the soft palate. It arises trudes above the gums and is covered by the
from the smaller, less developed venous plexus shiny white enamel. All the crowns of the teeth
of the hard palate, which is located in the sub of the dog except the canines end in tubercles
mucosa covering the bony palate. The lym (tubercula dentis). The neck (collum dentis) is a
phatics go to the medial retropharyngeal lymph slight constriction of the tooth located at the
nodes. gum line, where the enamel ends. The root
Nerves of the palate. The major palatine (radix dentis) is the portion below the gum, and
branch of the maxillary division of the trigem for the most part it is embedded in the alveolus.
inal nerve (n. trigeminus) courses through the Its pointed end is the apex o f the root (apex
palatine canal and supplies sensory fibers to the radicis dentis). Many teeth have more than one
oral side of the hard palate. The nasal side is root. Once teeth are fully erupted in the dog
supplied by the posterior nasal or sphenopala they cease growing. Because of the increased
tine branch of the same nerve. The main sen use and greater dependence on the teeth which
sory supply to the soft palate is the minor come about through growth, two sets of teeth
palatine branch of the maxillary nerve (n. maxil- are necessary. The first set is fully erupted and
laris), which follows the minor palatine artery functional early in the second month after
to enter the anterior end of the soft palate. birth. These teeth, known as deciduous teeth
Branches from the glossopharyngeal (n. glosso (dentes decidui), serve the animal during its
pharyngeus) and the vagus nerve (n. vagus) also most active puppyhood. Upon approaching ma
enter the soft palate and supply the pterygo turity, when the jaws have become longer and
pharyngeal and palatopharyngeal muscles. The larger and the small deciduous teeth are no
vagi contribute most to the supply of these longer adequate, they are shed and immediately
muscles; the glossopharyngeal nerves supply replaced by the permanent teeth (dentes per-
the lateral walls of the oral pharynx and, to a manentes), which last throughout adult life.
lesser extent, the soft palate. They are sensory These are larger than the deciduous teeth. The
also to the posterior part of the tongue. jaws continue to grow, so that additional per
Aponeurosis of the palate. The palatine apo manent teeth are added in back of the perma
neurosis (raphe palati) consists essentially of nent premolar teeth. These added teeth are the
the fanned-out terminal tendons of the right and true molars. There are two on each side of the
left tensor veli palatini muscles. It is located be upper jaw, and three on each side of the lower
tween the ventral margins of the right and left jaw.
perpendicular parts of the palatine bones and Observations on the development of the de
the pterygoid bones, and attaches anteriorly to ciduous teeth in the fetal dog have been
the posterior margin of the hard palate. It is thin reported by Hormandinger (1958), Satrapa-
but strong. Binder (1959) and Williams (1961). Postnatal
650 C h ap ter 13. T he D i g e s t i v e Sy s t e m and A bd o m en
calcification and eruption of the deciduous and rated from the comer incisors by an interdental
permanent teeth have been studied by Meyer space of about 3 mm. on the upper jaw and by
(1942), Hoppner (1956), and Amall (1961). less than this distance on the lower jaw in mes
The teeth are arranged as upper and b w er aticephalic heads. They are by far the longest
dental arches (arcus dentalis maxillaris et man teeth of the dog, having large roots which are
dibularis). The lower arch is narrower and nearly two times as long as their crowns. The
usually shorter than the upper (Seiferle and canine teeth of the upper jaw are about the
Meyer 1942). The teeth which compose the same length as the lower ones, but are more
dental arches are anchored in sockets, or alveoli, massive. They are transversely compressed.
of the upper and lower jaws. They are so placed When the mouth is closed the crown of the
that many of the teeth fail to meet or else over lower canine tooth occupies the interval be
lap or protrude between adjacent teeth lying tween the upper corner incisor and the canine
opposite to them. The function of these non- tooth. The roots produce arciform alveolar juga
occlusal or imperfectly occluding teeth is to which lie peripheral to the roots of the first pre
grasp, puncture, or shear. Stockard (1941) refers molar teeth. The canine teeth are pointed at
to the area between the canine and camassial each end and have a middle portion which may
teeth as the premolar carrying space. In dogs be slightly wider within the alveolus than at the
the food is bolted rather than masticated. gum line. This necessitates a flap operation of
Tooth surfaces. The surface of the tooth the maxilla over the root of the tooth to facili
which faces the lip or cheek is the labial or buc tate its proper removal.
cal surface (facies labialis or f. buccalis), and the All the teeth posterior to the canines are
surface which faces the tongue is the lingual often referred to as the cheek teeth (St. Clair
surface (facies lingualis). The surface adjacent and Jones 1957). They are divided into pre
to the next tooth in the dental arch is the con molars and molars. In the permanent dentition
tact surface (facies contactus). For the incisors there are four premolar teeth (dentes premo-
the contact surfaces are medial and lateral; for lares) on each side in each jaw. The first premo
the canine and cheek teeth they are anterior lar tooth in the dog is not replaced, and in this
and posterior. The surface which faces the op respect it is similar to the molars. It erupts be
posite dental arch is the occlusal or masticatory tween the fourth and fifth postnatal month and
surface (facies masticatoria). usually remains throughout life. Some authors
Tooth groupings. The upper teeth are at consider the first premolar to belong to the de
tached in the alveoli of the incisive (premaxil ciduous dentition, whereas others prefer to re
lary) and maxillary bones. Those with roots gard it as a member of the permanent dentition.
embedded in the incisive bones are the incisor Tims (1902) mentions other animals in which
teeth (dentes incisivi). The upper incisors premolar I erupts late in the growth period and
usually are placed slightly in front of the lower is not replaced.
incisors. The incisor tooth nearest the mid plane The placement of the premolar teeth may be
on each side in each jaw is incisor I, or central altered by changing the shape of the head
incisor. The second is incisor II, or intermediate through selective breeding, but regardless of
incisor, and the third is incisor III, or comer in head shape the teeth remain relatively constant
cisor. They are long, slender teeth, arched in form and size (Stockard 1941). The first pre
slightly forward and laterally compressed. They molar is the smallest, and has a single peglike
increase in size from the central to the corner root. The last, or fourth, premolar is the largest;
incisor. The central tubercles of the upper cor in the upper jaw it has three roots, in the lower,
ner incisors are largest and slightly hooked pos only two. The middle two premolar teeth are
teriorly, whereas those of incisors I and II are similar in both jaws. Each has two roots. The
irregularly trident transversely. Of the three crowns of the first three premolars are similar
projections or tubercles, the central one is larg to those of the corresponding teeth on the other
est and extends farthest distally; the small me side of the same jaw and of those in the oppo
dial projections are farther from the gums than site jaw. The first premolar tooth has a single
the lateral ones. A V-shaped ridge of enamel, tubercle which usually hooks slightly posteri
with its apex near the gum, connects the side orly. The ledge of enamel at the gum line is
tubercles on the lingual surface. Tubercles wear relatively straight medially and convex laterally.
off early in adult life or remain to old age, de The second and third premolars have similar
pending on the chewing habits of the dog. crowns. On each tooth the anterior border
The canine teeth (dentes canini) are sepa slopes diagonally backward to end in a strong,
652 C h ap ter 1 3 . The D ig e s tiv e S y s te m and A bdom en
pyramid-shaped point which inclines slightly shaped that the greatest compression force is
backward. The posterior border of this tubercle transmitted through it to the compact bone
is steep, ending in a relatively level, transversely which lies adjacent to the neck of the tooth.
rounded border. Adjacent to the base of the The lower molar teeth are peculiar in that
main tubercle on this border is a second smaller the first is about twice as large as the other two
tubercle 1 or 2 mm. high. The posterior mar together; it is the cam assial tooth of the lower
gins of the crowns of the two middle premolars jaw. The posterior third of each lower camas
are prominent. They may form small tubercles. sial tooth is adapted for crushing and grinding;
The upper fourth premolar teeth are the the anterior portion is sharp and pointed, and
largest cutting teeth of the upper jaw. They are suited for shearing action. The shearing portion
the cam assial (dentes camassei) or sectorial of each lower camassial tooth possesses the
(shearing) teeth. Each has three stout diverg strongest tubercles of any teeth of the lower
ing, conical roots. The two roots on the labial jaw and is roughly quadrilateral in form. This
side form prominent alveolar juga that reach as part of the crown is longer than the inner sur
far dorsally as a frontal plane through the infra face of the posterior part of the crown of the
orbital canals. The posterior root of each upper upper camassial tooth with which it forms a
camassial tooth is triangular and somewhat scissors. A small fossa is formed in the hard pal
transversely flattened, becoming wide at the ate, opposite its most salient tubercle, to receive
neck. The lingual root, a fourth shorter than the the tubercle. Seiferle and Meyer (1942) have
labial, is flattened in an oblique plane and described and illustrated the normal dentition
slightly twisted. A small tubercle is located on of the German shepherd.
the short crown which lies opposite its most Dental formulae. Since the teeth are grouped
medial part. According to Khuen (1952), the according to position and form, it is possible to
upper fourth premolar or camassial tooth is the express their arrangement as a dental formula.
tooth most commonly involved with root ab The abbreviation representing the particular
scesses. Such an abscess of one of the lateral teeth (I, incisor; C, canine; PM, premolar; M,
roots manifests itself usually by the formation of molar) is followed by the number of such teeth
a persistent fistula which discharges on the face on one side of the upper and the lower jaw.
anteroventral to the eye. A permanent cure in The formula for the deciduous dentition of
volves extraction of the affected tooth. To ac the dog is:
complish this, the tooth may be split so that the
roots can be removed separately. The fourth
premolar tooth in the lower jaw is similar in 1 1 C I PM | = 28
form to the two teeth in front of it but is slightly
larger. It is not the camassial tooth of the lower
jaw. The permanent dentition is represented as:
The molar teeth (dentes molares) have no de
ciduous predecessors. There are two on each
side of the upper jaw and three on each side of l | c ± P M | M | = 42
the lower jaw. In each jaw the first are the larg
est and the last are the smallest. The mastica
tory surfaces of the upper molars are multitu- Eruption of teeth. The deciduous second,
berculate and are at two levels. The anterior third and fourth premolar teeth are replaced by
parts are at a higher level than the posterior. the permanent premolars. The permanent teeth
The anterior masticatory surfaces of the upper develop beneath or to one side of the deciduous
molar teeth are irregularly flattened and make teeth which they are destined to dislodge. The
normal contact with the last two molars and permanent incisors erupt slightly posterior to
about the posterior third of the first molar of the the deciduous incisors. The central and inter
lower jaw. The lower molars, although multi- mediate deciduous incisors and the canine teeth
tuberculate, contact only the anterior parts of of both jaws have usually erupted by the end of
the upper molar teeth. the first month. The corner incisors erupt dur
Each upper molar tooth has three slightly ing the fifth or sixth week; the deciduous pre
diverging roots. The lingual root of each of molars, between the fourth and eighth weeks.
these teeth is more massive than either of the The time of eruption of the permanent teeth is
two buccal roots, although it is shorter. It is rather closely correlated with the life span of
slightly compressed anteroposteriorly and is so the breed. Variations are numerous, but the life
T h e M outh and A s s o c ia t e d St r u c t u r e s 653
span of a Great Dane or St. Bernard is about 8 The pulp (pulpa dentis), the only soft tissue
years; that of a setter, pointer, or hound, 13 contained in a tooth, is composed of sensory
years; and that of the toy breeds, 17 years. The nerves, arteries, veins, lymphatic capillaries, and
larger the breed, the shorter the life span. The a rather primitive connective tissue which holds
teeth erupt earliest in the large breeds. Table those structures together. The pulp is contained
13-1 gives the normal range of time for the in the pulp cavity (cavum dentis). A small ap
eruption of each of the permanent teeth, which ical foram en (foramen apicis dentis), at the end
erupt at about the same time, in each jaw. of each root, allows free passage of the vessels
Eruption of the permanent canine teeth usually and nerves in and out of the tooth through the
precedes the shedding of the corresponding root canal (canalis radicis dentis).
deciduous ones; for two or three weeks, or even Dental anomalies. Deviations from the nor
longer, the permanent canine is visible antero- mal number or placement of the teeth are the
medial to the corresponding deciduous tooth. most common dental anomalies reported in
Table 13-1. Eruption of Permanent Teeth dogs. Such anomalies seldom occur in the doli
GROUP TOOTH E R U P T IO N P E R IO D
chocephalic breeds, but are encountered very
Incisors Central 2 to 5 months commonly in the brachycephalic dogs. As the
Intermediate 2 to 5 months muzzle is shortened, deviation in placement of
Corner Most breeds 4 to 5 months the teeth results. The size of the teeth does not
Canine 5 to 6 months decrease proportionately with a reduction in
Premolars First 4 to 5 months the length of jaw (Stockard 1941). In one Bos
Second 6 months ton terrier only three lower cheek teeth were
Third 6 months present on one side of the lower jaw, and four
Fourth 4 to 5 months on the other. There was no evidence of filled-in
Molars First 5 to 6 months alveoli, and the dental arch was full. A few rela
Second 6 to 7 months tively large teeth, properly placed, are probably
Third 6 to 7 months more efficient than many teeth placed at dis
torted angles.
Structure of teeth. The dense, pearly-white Supernumerary teeth are occasionally found
outer layer of the crown is enamel (enamelum). in all breeds. Extra incisors and premolars are
It is the hardest substance in the body, and most common. Usually, when an increase in the
gives sparks when struck with steel. Only about number of premolars occurs, two single-rooted
5 per cent of enamel is organic matter. It is premolars are located behind the canine. An
thickest on the wearing surfaces of the teeth, extra incisor is commonly in series with the
and its hardness gradually increases as the dog other incisors, although an extra incisor located
gets older. Glock et al. (1942) found that it is on the palate has been recorded (Colyer 1936).
possible to remove enamel from the teeth of Supernumerary teeth are usually unilateral, and
puppies up to 17 weeks of age by scraping with occur in the upper jaw more frequently than in
a scalpel. the lower. The number of teeth is seldom re
Dentin (dentinum), similar to bone in chem duced, except in brachycephalic breeds. The
ical composition, forms the bulk of the tooth, greatest reduction involves the cheek teeth of
enclosing the pulp cavity internally and under the lower jaw. In old specimens lost teeth with
lying the enamel externally. It is known as a resultant obliteration of the alveoli must not
“ivory,” and is yellowish white in color. It con be mistaken for a congenital decrease in num
tains few nerves, but through the processes of ber.
the cells (odontoblasts) that form it a conductive The third upper premolar is the first tooth to
system is established which gives sensitivity to rotate as the muzzle is shortened by selective
touch, cold, and other stimuli. Gradual erosion breeding. Later, all upper premolars may be
of the dentin, as occurs from wear in the aged, rotated. The second and third premolars may
is not accompanied by pain, as the conducting rotate either clockwise or counterclockwise, al
fibers recede or calcify in advance of the wear though rotation which brings their anterior roots
ing surface. Injured dentin is repaired only un nearer the median plane is the more common.
der favorable conditions. Little additional room is gained by rotation of
The cementum in the dog is a thin covering the first premolar, since its width nearly equals
of bonelike tissue found only on the roots. its length. The long fourth upper premolar,
Grossly, the cementum cannot be differentiated however, usually rotates so that its anterior roots
from the dentin which it covers. lie nearer the median plane than in the normal
654 C h ap ter 13. T h e D ig e s tiv e S y s te m a n d A bdom en
specimen. Sometimes the second and third pre nally. On the root or posterior part it becomes
molars he parallel to each other transversely or slightly rounded, particularly from side to side.
sagittally. The molar teeth seem to be little A deep, narrow, straight median groove (sulcus
affected by rotation. medianus linguae) divides the dorsum of the
All anomalous placements of teeth are more organ into symmetrical halves throughout its
common in the upper than in the lower dental anterior three-fourths. No terminal sulcus, fora
arch. Wood and Wood (1933) describe a dog men cecum, or thyroglossal duct persists in the
which had three permanent molars on one side dog. The ventral surface o f the tongue (facies
of the upper jaw, and cite nine other accounts ventralis linguae) is convex transversely and
of this rare anomaly. These authors give four lies on the mucosa covering the mylohyoid mus
explanations for extra teeth in general: mechani cles. The margin o f the tongue (margo linguae),
cal splitting of the tooth germ, retention of a marking the junction of its dorsal and ventral
deciduous tooth, mutation producing a new surfaces, is thin at the apex (free end) but grad
tooth, and reversion to an ancestral condition. ually thickens over the body. The root o f the
The findings in the specimens mentioned by tongue (radix linguae) is the caudal third of
Wood and Wood are probably explained by the the organ, being the portion through which the
reversion principle, since these authors state three paired extrinsic muscles enter. The body
that the third upper molar was probably lost by o f the tongue (corpus linguae) is the long slender
the ancestors of the modern dog before the part anterior to the root. The body ends ante
middle of the Oligocene period. riorly by becoming the tip or apex o f the tongue
Deviation from the normal number of roots (apex linguae), which is its highly mobile, free
or fusion of the roots or of whole teeth is rarely extremity.
found. Colyer (1936) and Meyer (1942) describe The mucosa of the tongue (tunica mucosa
several different kinds of dental anomalies in linguae) is comified, stratified squamous epithe
the German shepherd. Graves (1948) cites the lium which is closely attached to the dorsum
case of an English sheep dog which grew a third and sides of the organ and helps to form the
set of teeth after extraction of 11 of the perma various types of papillae. On the ventral surface
nent teeth when the dog was nine years old. it is thin, less heavily cornified, and loosely at
Dechambre (1912) gives a brief description of tached to the tongue musculature. An unpaired,
an English toy terrier which had a total absence ventral median fold of mucosa, the lingual
of all teeth. frenulum (frenulum linguae), runs from the
floor of the mouth mainly to the body of the
Gums tongue. Posteriorly, it contains the right and
left closely apposed genioglossal muscles. An
The gums (gingivae) are composed of dense
teriorly, the two layers of mucosa are united by
fibrous tissue covered by smooth, heavily vas
delicate areolar tissue. On each side of the
cularized mucosae. The lamina propria is thick
frenulum on the ventral surface of the tongue
and strong. It extends around the necks of the
is the fim briated plica (plica fimbriata). It is a
teeth and down into the alveoli to be continuous
slightly protruding, rounded fold of mucosa, the
with the alveolar periosteum. The gums are con
lateral border of which essentially parallels the
tinuous externally with the mucosa of the vesti
lateral border of the tongue. This plica begins
bule, and internally with that of the floor of the
near the anterior end of the frenulum and fades
oral cavity proper or of the hard palate. In those
away after running posteriorly a distance of
breeds with pigmented oral mucosa the gums are
about 2 cm. Its free margin is only slightly fim
likewise pigmented.
briated in the dog; the plicae are only partly
effaced by raising the tongue. The lingual vein
Tongue
lies closely under the mucosa between the
The tongue (lingua) (Fig. 13-3) is a muscular frenulum and the lateral border of the fimbri
organ which forms most of the floor of both the ated fold; it may be used for intravenous in
mouth and the oral pharynx. It consists basically jections.
of a mass of interwoven bundles of muscle fibers The papillae of the tongue (papillae linguales)
with a distinct dorsal longitudinal stratum and are projections of the corium and lamina propria
a surface mucosa. covered by stratified squamous epithelium.
The dorsum o f the tongue (dorsum linguae) They are located on the dorsal surface and mar
is nearly flat, both transversely and longitudi gins of the tongue. Some of the papillae house
T he M o uth and A s s o c ia t e d Stru c tu r es 655
A r e a of f o l i a t e p a p i l l a e
Fungiform p a p illa e
F i l i f o r m papi l lae
Lingual frenulum
taste buds; others constitute the rough surface Function of the tongue. The tongue with its
which aids in grooming and feeding. Based on papillae and taste buds serves primarily as a
their shape, five types of papillae are recog ladle for fluids, a receptor for gustatory sensa
nized: filiform, fungiform, vallate, foliate, and tions, and a swab for the cleansing of wounds
conical. These and the taste buds are discussed or the grooming of the hair. A brood bitch uses
in Chapter 17. her tongue to dry, clean, and stimulate her
Muscles of the tongue. The muscles of the young. By means of licking, she provides the
tongue (musculi linguae) are divided into ex stimulus necessary for the initiation of defeca
trinsic and intrinsic groups, both of which are tion and urination in the newborn. The tongue
described in Chapter 3. The extrinsic muscles aids in reducing body temperature through lol
consist of the genioglossus, styloglossus, and ling, and assists in swallowing, prehension, and
hyoglossus, all of which are paired. The intrinsic mastication. Bennett and Hutchinson (1946)
muscles of the tongue have never been exten investigated the movements of the tongue in the
sively studied (Bennett and Ramsay 1941). They dog. They concluded that protrusion of the
consist of dorsal, ventral, longitudinal, trans tongue is due to the action of the genioglossus
verse, and vertical muscle fiber systems. muscles which pull the base of the tongue for
The lyssa is a filiform body about 4 cm. long ward, and the intrinsic muscles which undergo
and 2 mm. wide at its greatest diameter. It lies elongation. Experimentally, if the tongue is pro
in the median plane between right and left truded by the action of the genioglossus and
genioglossal muscles. It extends to within 1 or 2 intrinsic muscles of one side, the non-contracting
mm. of the tip of the tongue. Its anterior part half affects the direction of the protrusion in
lies directly adjacent to the mucosa. Muscle such a manner that the tip is directed contra-
fibers appear to insert on its cranial part. Traut- laterally.
mann and Fiebiger (1957) state that the lyssa
contains adipose tissue and may contain islands S a l iv a r y G l a n d s
of cartilage in its posterior part. They mention
the presence of a small amount of striated mus The salivary glands, broadly speaking, are all
cle in the middle part of the lyssa. A branch of of those glands which pour their secretions into
the hypoglossal nerve supplies it. Bennett (1944) the oral cavity. These are the parotid, mandib
found that stimulation of this nerve caused a ular, sublingual, and zygomatic glands, which
visible elongation of the lyssa. He suggested that are the salivary glands in the usual or restricted
the lyssa may act as a stretch receptor. sense.
Vessels and nerves of the tongue. The lin
gual artery is the main blood supply to the Parotid Gland
tongue; the sublingual artery is distributed
mainly to the mylohyoid and the anterior por The parotid gland (glandula parotis) (Fig.
tion of the digastric muscle. The veins of the 13-4) lies at the junction of the head and neck
tongue are satellites of the arteries. The vessels and closely embraces the basal portion of the
of the tongue are described in Chapters 4 and 5. auricular cartilage. Its outline is V-shaped as
Brown (1937) describes arteriovenous anasto viewed from the surface. The apex is directed
moses in the tongue of the dog which have an ventrally, and the two diverging dorsal limbs
extremely rich nerve supply. It is suggested that lie on each side of the ventral part of the auricu
these anastomoses function in the elimination lar cartilage. The parotid gland is bounded
of body heat. The lymph vessels drain into the posteriorly by the sternomastoideus and cleido-
medial retropharyngeal lymph nodes. The cervicalis muscles, and anteriorly by the mas
tongue is supplied by three different nerves. seter muscle and the temporomandibular joint.
The hypoglossal nerve is motor to the muscles The gland weighs about 7 gm. and has an over
of the tongue. The lingual branch of the man all length of approximately 6 cm. It is thickest
dibular portion of the trigeminal nerve furnishes ventrally, measuring about 1.5 cm. The gland
ordinary sensory fibers to the anterior two-thirds is a dark flesh color, with coarse lobulations visi
of the organ, and the fibers carried by the ble through its thin capsule. It is divided into
chorda tympani, a branch of the facial, join superficial and deep portions, and possesses
those of the lingual to be distributed to the taste three angles, and three borders, and two sur
buds of the tongue. The posterior third, or root faces.
of the tongue, receives its entire sensory inner The superficial portion (pars superficialis) of
vation through the glossopharyngeal nerve. the parotid gland consists of the two limbs of the
T he M o u th and A s s o c ia t e d Stru ctures 657
L o c a t i o n of p a l a t i n e t o n s i l
P a ro tid duct,
i T e m p o r a l m.
{Z y g o m a t i c a r c h
P t e r y g o i d e u s me d . m.
Palatine glands
prbitat ligament
L a c rim a l gland
Z y g o m a t i c gl and
/ O r i f i c e of m a j o r z y g o ma t i c du c t
,O r i f i c e o f p a r o t i d d u c t
O r i f i c e o f m a n d i b u l a r d uct
St erno-
Ai G e n i o g i a s s u s m.
t h y r o i d e u s m! S u b l i n g u a l gland (polijstom atic p a rt)
Cricothyroideus m Mylohyoideus rn.
Sternohyoideus m ' D i g a s t ri c u s m
V and the dorsally concave, thin-edged portion fascia as it runs straight forward to the cheek
which connects the two limbs. parallel, or nearly parallel, to the fibers of the
The deep portion (pars profunda) of the pa masseter. It opens into the buccal cavity at the
rotid gland is wedge-shaped and lies ventral to anterior end of a blunt ridge of mucosa by a
the cartilaginous external ear canal. It is dorsal small papilla. This is located opposite the poste
to the anterior pole of the mandibular gland rior margin of the fourth upper premolar, or
and extends medially toward the tympanic bulla camassial (shearing) tooth. The ridge of mucosa,
and wall of the nasal pharynx. on the anterior end of which the duct opens, runs
Of the three angles of the parotid gland, one posteriorly to the end of the dental arch lying
points dorsoanteriorly, one dorsoposteriorly, and in or near the fornix formed by the attachment
the third ventrally. The borders are dorsal, an of the cheek to the upper jaw.
terior, and posterior. The superficial surface is Accessory parotid glands (glandulae parotis
nearly flat transversely and only slightly convex accessoriae) are usually present on one or both
longitudinally. It is crossed vertically by the sides. They range in size from single lobules to
straplike m. depressor auriculae (m. parotideo- small oval glandular masses over 1 cm. long.
auricularis) which, in turn, is covered ventrally They usually lie above the parotid duct and may
by fascicles of the platysma. Near the ventral be placed at any level along it. Their small ducts
angle the gland is usually tunneled by the inter empty into the main parotid duct.
nal maxillary vein. From under its anterior bor The fascia covering the parotid gland is thin
der emerge the palpebral, auriculotemporal, and and of the areolar type. At the borders of the
the dorsal and ventral buccal nerves. The pa gland it blends with the superficial fascia of
rotid lymph node usually lies for the most part the head, ear, and neck. Similar fascia lines the
under the anterior border near the ventral space in which the gland lies. The fascia sepa
angle. The anterior auricular artery and vein rating the lobules is abundant and loose.
and the transverse facial artery run under or The parotid artery, a branch of the external
along the anterior border. The posterior border carotid, is wholly distributed to the parotid
is circled by branches of the intermediate auric gland and usually is its main blood supply. The
ular blood vessels. Some of these structures may great auricular, masseter, transverse facial, and
run through the gland. anterior auricular arteries all send twigs to the
The dorsal border is not related to any large parotid gland. The veins which drain the pa
nerves or vessels, but its position is important rotid are radicles of the superficial temporal and
because of its close proximity to the external ear great auricular veins. The lymphatics from the
canal. Blakeley (1957) gives data on the success parotid gland drain into the parotid and medial
of the operation to extend the intertragic inci retropharyngeal lymph nodes. The parotid
sure to a level of the horizontal part of the ex gland receives parasympathetic nerve fibers
ternal ear canal. This operation is highly through the auriculotemporal branch of the
successful in correcting chronic otitis externa. trigeminal nerve. These fibers originally come
In such an operation the dorsal border of the from the glossopharyngeal nerve and run to the
parotid gland, unless it was retracted, would be auriculotemporal in the short tympanic nerve.
partly incised. The deep part of the parotid The sympathetic fibers reach the gland from the
gland is related to the facial nerve and its termi external carotid plexus of nerves by branches
nal branches as they emerge posterior to the which follow mainly the parotid artery to the
osseous external acoustic meatus. The deep por gland. Histologically, the parotid salivary gland
tion of the gland is also related to the maxillary is a serous, compound tubulo-alveolar gland.
and superficial temporal arteries. The internal
maxillary vein is related to the parotid gland at Mandibular Gland
a more ventral and superficial level than are the
nerves and arteries. The mandibular gland (glandula mandibularis)
The parotid duct (ductus parotideus) is about (Fig. 13-4) is an ovoid body lying largely be
1.5 mm. in diameter and 6 cm. long. It is formed tween the external and internal maxillary veins
by two or three converging radicles which leave just posterior to the angle of the jaw. It is about
the ventral third of the anterior border of the 3 cm. long, 2.5 cm. wide, and 1.5 cm. thick. It
gland and unite with each other on the mas weighs approximately 8 gm., being a little heavier
seter muscle several millimeters from the gland. than the parotid gland. Its lobules, which are of
The duct is rather closely united to the lateral about the same dimensions as the parotid lobules,
surface of the masseter muscle by superficial are fitted together much more compactly, with
T he M o u th and A s s o c ia t e d Str u c t u r e s 659
less connective tissue separating them. The out the remaining part of its course the man
whole organ is a light buff color and is not sharply dibular duct is closely related to the sublingual
separated from the smaller monostomatic por duct, and is described with it following the
tion of the sublingual gland, which is slightly description of the sublingual gland.
darker in color. This portion of the sublingual The largest artery supplying the mandibular
gland butts against the ventral part of the cranial gland, the glandular branch of the facial,
pole of the mandibular gland on which it leaves enters the gland where the mandibular duct
a nearly flat, oblique impression. leaves it. Entering the dorsal part of the deep
From a superficial gross appearance this por surface of the gland are one or two small
tion of the sublingual gland constitutes the branches from the great auricular artery. The
pointed cranial pole of the salivary mass in this chief vein draining the gland leaves its deep sur
location since both of the salivary masses are face and terminates usually in the lingual vein
contained within the same heavy fibrous cap as this vessel enters the external maxillary. A
sule. The common capsule is a specialization of second vein leaves the posterior part of the
the buccopharyngeal fascia and does not send gland and terminates either in the facial, maxil
trabeculae into the glands. It is derived primar lary, or lingual vein. Its parasympathetic fibers
ily from the deep cervical fascia. The man come from the facial nerve. These fibers first
dibular gland has cranial and caudal poles, and run in the chorda tympani to the mandibular
superficial and deep surfaces. Its cranial pole is branch of the trigeminal nerve and continue in
truncate and is related to the major portion of the lingual branch of the latter to the mandib
the sublingual gland; the caudal pole forms an ular ganglion, where they synapse with post
even arc vertically as it unites the superficial ganglionic neurons. Secretory fibers from these
and deep surfaces at an acute angle. The super cells run with the mandibular duct to the
ficial surface is slightly rounded in all planes gland. Sympathetic fibers reach the gland by
and is grooved dorsally for the internal maxil means of the perivascular plexus around the
lary vein. Its dorsocranial part is variably over glandular artery. The lymphatics drain into the
lapped by the parotid gland; ventrally it is medial retropharyngeal lymph node.
related to the mandibular lymph node, or nodes,
lying dorsal to the external maxillary vein. The
deep surface is further divided into subsurfaces
by the following structures on which it lies: the Sublingual Gland
muscle and terminal tendon of the stemomas-
toideus, dorsocaudally; the medial retropharyn The sublingual gland (glandula sublingualis)
geal lymph node and larynx, medially; and the is the smallest of the four pairs of major salivary
digastric and ribbon-like stylohyoid muscles, glands. It weighs about 1 gm. and consists of an
cranially. aggregation of two or more lobulated masses.
The mandibular gland is a mixed gland. The nearly flat, truncated base, or posterior
Stormont (1928) gives a brief review of the liter surface, of the largest division of the gland is
ature concerning the histology of the salivary closely related to the blunt anterior end of the
glands of carnivores. mandibular gland. Both glands are enclosed
The mandibular duct (ductus mandibularis) in the same heavy, fibrous capsule, the sublin
leaves the medial surface of the gland near gual gland being distinguishable by its slightly
the ventromedial part of the impression formed darker color. The tapered extremity of the sub
by the sublingual gland. As the initial part lingual gland extends anteromedially between
of the duct runs anteromedially it lies in the posteromedial border of the masseter mus
relation to the medial surface of the sublingual cle, anteriorly and the anterolateral surface of
gland. In association with this gland the duct the digastric muscle, posteriorly. It curves ante
lies between the masseter muscle and mandible riorly upon reaching the lateral surface of the
laterally, and the digastric muscle medially. In styloglossus, so that it lies medial to the body of
this location the duct runs obliquely antero the mandible.
medially. Upon reaching the lateral part of the Usually, separated from the anterior end of
pharyngeal mucosa it arches forward and, with this portion of the gland, is an ovoid cluster of
lobules of the anterior part of the sublingual lobules which may reach a length of 3 cm. and
gland, runs in the intermuscular septum be a width of 1 cm. These lobules lie directly under
tween the medially located m. styloglossus and the mucosa which reflects from the jaw to the
the laterally located m. mylohyoideus. Through tongue. Their secretion is poured into the main
660 C h ap ter 13. T he D ig e s t iv e Sy stem and A bd o m en
sublingual duct through four to six short excre zygomatic arch. Found only in the dog and cat,
tory ducts. The sublingual duct lies ventral to among the domestic mammals, it represents a
the gland. Since these portions of the gland posterior condensation of the largely unilobu-
empty into the main sublingual duct, they are lated dorsal buccal glands of other mammals. It
known as the pars monostomatica of the sub is globular to pyramidal in shape, with its base
lingual gland. The pars polystomatica consists directed upward and backward and lying
of that portion of the sublingual gland which against the ventral part of the periorbita. It is
discharges its secretion directly into the oral surrounded by soft fat outside of a feebly devel
cavity without its passing through the main sub oped capsule. Because of the soft tissue adjacent
lingual duct. The gland consists of 6 to 12 small, to it, its lobules are much more distinct than are
usually isolated lobules of salivary tissue which those of the mandibular gland, which it re
lie under the mucosa on each side of the body sembles in color. From its blunt apex, which
of the tongue anterior to the lingual branch of lies lateral to that part of the maxilla containing
the trigeminal nerve. They are so small that no the roots of the last upper molar tooth, it sends
definite sublingual fold is formed by them. They one major duct and two to four minor ducts to
open into the ventral fornix of the vestibule by the posterior part of the buccal cavity. The
several ducts. major duct opens about 1 cm. posterior to the
The sublingual duct (ductus sublingualis) is parotid papilla on the ridge of mucosa that ex
closely related to the mandibular duct through tends to a plane through the posterior surface
out its course in the intermandibular space, of the last upper cheek tooth. The smaller,
being located dorsal to it. On reaching the minor ducts open on this ridge, posterior to the
posterior margin of the mylohyoid muscle opening of the major duct.
which is located in a transverse plane less than Usually, the first branch of the infraorbital
1 cm. anterior to the angle of the jaw, the two artery, as it enters the infraorbital canal, sup
ducts pass medial and dorsal to this muscle. plies the zygomatic gland. The main vein which
Anterior to the lingual branch of the trigeminal leaves it enters the reflex vein, which grooves
nerve, which crosses the lateral surfaces of the its lateral surface.
ducts at a transverse level just posterior to
the orbital openings, the ducts lie between the
PHARYNX
genioglossal and mylohyoid muscles. The ducts
open on a small sublingual papilla (caruncula The pharynx (cavum pharyngis) (Figs. 13-1,
sublingualis), which is located lateral to the an 13-6) is a passage which is, in part, common to
terior end of the frenulum at its attachment pos both the respiratory and the digestive system.
terior to the symphysis of the mandibles. It extends approximately from a transverse
According to Michel’s (1956) observations (30 plane through the head at the level of the orbital
dogs), in two-thirds of the specimens the ducts openings to a similar plane through the second
open individually; in one-third they have a com cervical vertebra. The anterior part of the phar
mon opening. When the openings are separate ynx is divided by the soft palate into an exclu
the mandibular opening lies anterior to the sively respiratory, or dorsal, portion, and an
sublingual. Michel describes his technique for alimentary and respiratory, or ventral, portion.
cannulating these ducts. At the posterior end of the soft palate the two
The glandular branch of the facial artery sup portions cross or become coextensive, forming
plies the monostomatic portion, and the sub the pharyngeal isthmus (isthmus pharyngeus).
lingual artery, a branch of the lingual, supplies The isthmus is bounded posteriorly by the ven
the small polystomatic part of the sublingual tral surface of the epiglottis when the opening
gland. These arteries are accompanied by satel to the larynx is closed. When the laryngeal
lite veins which drain the gland. Lymphatic entrance is open the cavities of the pharynx and
drainage is into the medial retropharyngeal larynx are directly continuous.
lymph node. Time-honored names for the anterior parts
of the pharynx are: nasal part, for the respiratory
Zygomatic Gland portion, and oral part, for the digestive portion.
Posterior to the pharyngeal isthmus the diges
The zygomatic gland (glandula zygomatica) tive tube is continued dorsal to the larynx as the
(Fig. 13-5), also known as the orbital gland, laryngeal part of the pharynx, and the respira
weighs about 3 gm. and is located ventral to the tory tube is continued ventrally as the larynx,
P harynx 661
------- — A Deep f a c i a l v.
P t e r y go p h a r y n ge u s m, Levator v e li palatini m.
Hypophysis( 1 f i Palatopharyngeus m.
Soft palates \ ' / /Rt tonsillar a
N a s a l p harynx y Hyopharyngeus m-
P t er ygo id e us med m , K e r a t o h y oi de us m.
Cut edge a f mucosa - ' C u t edcje of mucosa
M y l o h y o i d e u s m- ~Br of c r an i a l l aryngeal a.
Sublingual salivary g l a n d - - L or yng ea l p h a r y n x
( monostomati c p a r t ) _ - Inte r a r yt en oi d c a r t i lage
Oral pharynx - -Cricoid cartilage
Styloglossus m - ~ - Thyr ohyoi d bone
L m g u a l b r of IX n - - Vocal f o l d
Genioglossus m " " " C r i c o i d c a r t i l ag e
' ,
Cemohyoideus m-' , 1 \ \ ' St er na hyo i de uS m.
/
Mylohyoideus m/ 1 v ' T h y r o i d c a r t i lacje
Epi hyoi d bone' \ ' Ventricle
i
Hyogl ossus my t ' Epiglottis ( r e t r a c t e d )
Rt h y o e p i g l o t t i c u s m- B a s i h y o i d bone
Fic. 13-6. Mid-sagittal section through the pharynx. The location of the right palatine tonsil is indicated by a dotted circle.
The biiccopharyngeal fascia deep to the tonsil was removed and the soft palate elevated.
662 C h ap ter 13. T he D ig e s t iv e Sy st e m and A bd o m en
which in turn is continuous with the trachea. The palatine tonsil has no afferent lym
The respiratory portion of the pharynx is dis phatics. Its efferent vessels drain into the medial
cussed in Chapter 14. retropharyngeal lymph node. It receives its
The isthmus of the fauces (isthmus faucium) nutrition mainly from the tonsillar artery, which
is the short tubal connection between the oral is derived from the lingual. The tonsillar artery
cavity and the oral part of the pharynx. It is enters the middle or widest portion of the tonsil
bounded on each side by the palatoglossal by about three branches. The posterior pole of
arches, ventrally by the tongue, and dorsally by the tonsil may receive twigs from the hyoid
the soft palate. branches of the lingual. The small veins from
The oral portion (pars oralis) of the pharynx the tonsil enter the palatine plexus of veins. Be
extends from the isthmus of the fauces to the sides the palatine tonsils other lymphoid tissue
pharyngeal isthmus dorsally and to the larynx lies in the base of the tongue and in the nasal
ventrally. It is bounded dorsally by the soft part of the pharynx. These are referred to as the
palate, ventrally by the root of the tongue, and lingual and pharyngeal tonsils. The lingual ton
laterally by the tonsillar sinus, with its contained sil (tonsilla lingualis) is so diffuse, it cannot be
palatine tonsil. The lateral wall of the oral part seen in gross. The pharyngeal tonsil (tonsilla
of the pharynx, which provides the site for the pharyngea), or adenoid, is described with the
palatine tonsil, is called the fauces. nasal part of the pharynx in Chapter 14.
The palatine tonsil (tonsilla palatina) is a The laryngeal part (pars laryngea) of the
long, relatively thin lymph node, located in the pharynx (Fig. 13-7) is that portion which lies
lateral wall of the oral part of the pharynx dorsal to the larynx. It extends from the pha
(fauces), just caudal to the palatoglossal arch. ryngeal isthmus and the nasal part of the phar
It is hidden from casual observation because of ynx in front to the beginning of the esophagus
its position in the tonsillar sinus (sinus tonsil behind. Its caudal limit, therefore, reaches to a
laris). The medial wall of the sinus is formed by transverse plane which passes through the
the thin, falciform tonsillar fo ld (plica semi caudal border of the cricoid cartilage and the
lunaris), which is a fold from the ventral surface middle of the axis. The pharyngo-esophageal
of the lateral portion of the soft palate. junction in the dog is distinctly marked by an
The palatine tonsil is divided into a protrud annular ridge of tissue known as the annular
ing, fusiform portion which composes the major fo ld (plica annularis).
portion of the organ, and a usually smaller, Although the function of the laryngeal part
deeper, minor portion which lies under the of the pharynx is both respiratory and alimen
mucosa forming the anterior part of the lateral tary, its chief importance is in deglutition. The
wall of the tonsillar sinus. The deep portion of bolus of food ingested is conveyed to it by
the gland may be formed as a result of tonsillitis, the plunger-like movement of the base of the
since it is usually absent in young healthy speci tongue. Six pairs of extrinsic muscles control
mens. It develops in the submucosa directly on the shape and size of the nasal and laryngeal
the thick buccopharyngeal fascia. Lateral to this parts of the pharynx. Three pairs of these mus
fascia lies the lingual branch of the glossopha cles are constrictors, two pairs are shorteners,
ryngeal nerve, and the styloglossal and medial and the muscles of one pair act as a dilator.
pterygoid muscles. Dyce (1957) uses the name, laryngopharyngeus,
The palatine tonsil has a long, narrow hilus to designate the combined cricopharyngeal and
which may be thickened in its middle so that thyropharyngeal muscles, which are the two
anterior and posterior fossae are formed which caudal pairs of constrictors. As Dyce states,
separate the main portion of the tonsil from the these two muscles are largely fused to each
lateral wall of the tonsillar sinus. When the other and they are also blended caudally with
major portion of the organ is forcibly pulled out the spiral muscular coat of the esophagus. The
of the tonsillar sinus the deeper, minor portion muscles of the pharynx are described in Chap
is also drawn downward in a fold of mucosa ter 3.
formed by the traction, so that complete extir The laryngeal part of the pharynx receives
pation of the organ is made possible. The aver its nutrition through the paired pharyngeal
age dimensions of the major portion are: length branches of the cranial thyroid and the ascend
2.5 cm., width 0.5 cm., and thickness 0.4 cm. ing pharyngeal arteries. The soft palate, the
The pointed ends of the fusiform organ are not dorsal wall of the oral part of the pharynx,
free but are firmly attached to the dorsolateral receives most of its nutrition from the paired
parts of the wall of the sinus. minor palatine arteries.
P h arynx 663
Nasal pharynx
C u t edge of p h a r y n c / e a l mm
„ Soft p a l ate
LaryngeaI o r i f i c e -
_- E p ig lo tt i s
A rye p ig lo ttic fo ld - .
L o c a t i o n of d o r s a l
b o r d e r of t h y r o i d c a r t i l a g e —C uneiform tu bercle
-Laryngeal pharynx
L o c o t i o n of c r i c o i d c a r t i l a g e '
Muc ou s g l a n d s
A n n u l a r fold
Esophagus
Trachea
by serosa, its adventitia blends with that proper fibers of the inner coat, instead of becoming
fascia of the organs with which it comes in more transverse, become nearly longitudinal
contact. and pass to the outside. They continue on the
The muscular coat (tunica muscularis) (Fig. visceral wall of the stomach as its outer longi
13-8) consists essentially of two oblique layers tudinal layer. The longitudinal fibers of the
of striated muscle fibers. The external muscular dorsal surface of the esophagus continue on the
coat arises on the ventral side of the esophagus dorsal wall of the stomach. The nearly trans
from the medial dorsal crest of the cricoid and verse, inner muscular fibers of the esophagus
the comiculate portions of the arytenoid carti partly blend with the circular and oblique fibers
lages by means of the cricoesophageal tendon of the stomach. The division between the
(tendo cricoesophageus). This tendon is a dis striated musculature of the esophagus and the
tinct thin band of collagenous tissue about 0.5 smooth musculature of the stomach cannot be
cm. wide and 1 cm. long. It tapers to a point determined by gross examination.
caudally as muscle fibers leave each side of it. The cervical portion of the esophagus pos
It, more than any other structure, serves as the sesses, in addition to the two oblique coats,
fixed point (punctum fixum) of cranial attach several poorly developed groups of longitudinal
ment of the esophagus (Sauer 1951). fibers. The right and left lateral longitudinal
The first few muscle fibers to arise arch bands are best developed. They arise under
sharply outward and upward on each side and neath the cricopharyngeal muscles from the
meet dorsally as transverse fibers. They are not fascia adjacent to the lateral borders of the dor
distinct from the caudal fibers of the cranially sal cricoarytenoid muscles. Some fibers contrib
lying cricopharyngeal muscles. Helm (1907) uting to these bands come from the cricoesoph
appropriately called this initial portion of the ageal tendon. The muscles are 1 to 2 mm. wide.
external esophageal muscle coat, the cricoesoph They usually fade away on the caudal portion
ageal muscle (m. cricoesophageus). The fibers of the cervical part of the esophagus, but they
from each side do not decussate mid-dorsally may extend to the mid-thoracic part. Inner and
but blend with each other without demarcation. outer ventral longitudinal esophageal muscle
Although the first few fibers are nearly trans fibers can usually also be recognized. The inner
verse in direction, the subsequent fibers become ones arise from the cricoesophageal tendon and,
increasingly more oblique so that in fusing with after running about 2 cm., become dispersed
their fellows they form progressively narrower on the main inner muscular coat. They lie on
loops or ellipses caudally. the dorsal surface of the ventral decussations
The main musculature of the esophagus which they partly cover. The outer longitudinal
caudal to the cricoesophageal fibers is in the muscle fibers lie ventral to the ventral decussa
form of spiral fibers. These start about 5 cm. tions. They are so feeble that they cannot always
from the beginning of the esophagus and con be dissected in gross with certainty. In some
tinue to within 5 to 10 cm. of the cardia of the specimens extremely delicate longitudinal mus
stomach. The superficial fibers on one side be cle bundles have been seen lying between the
come the deep ones on the other side. These inner and outer muscular coats in the cranial
apparently continuous oblique bundles spiral part of the cervical portion of the esophagus.
around the esophagus in such a way that they The submucous coat (tela submucosa) loosely
cross each other at nearly right angles in mak connects the mucous and the muscular coats. It
ing up the two main muscular coats of the allows the relatively inelastic mucous coat to be
organ. The lines of decussations are dorsal and thrown into heavy longitudinal folds when the
ventral in position. The line of ventral decussa esophagus is contracted. It contains blood ves
tion seems to stop about 10 cm. from the cardia, sels, nerves, and mucous glands. Trautmann and
whereas the dorsal decussations end about 5 cm. Fiebiger (1957) state that the esophageal glands
from it. The decussations do not end abruptly (gl. esophageae) form a continuous stratum that
as there is a gradual shifting of the direction of extends to the vicinity of the stomach. The
the fibers of the inner and outer muscle coats. muscular layer o f the mucosa (lamina muscu
The fibers of the inner coat become more laris mucosae), according to these authors, is
transverse in direction, while those of the outer present only in the caudal half of the esophagus
coat become more longitudinal, especially dor- and forms a continuous layer only near the
sally as they approach the cardia. About 3 cm. stomach.
from the cardia, ventrally, many of the oblique The mucous coat (tunica mucosa) is com-
666 C h ap ter 13. T h e D ig e s tiv e S y ste m and A bdom en
C ornicu la te
tub e rcle ~
B T h y r o i d cart.
Cr i co- esophageal lig.
- - T hy r oi d car t . Fibers form in g _
-Cricothyroideus o u t e r d o r s a l l o op s
-Cricopharyngeus' Cricopharyngeus m
------ T r a c h e a - -
Spiral de cussation
— O u te r d o r s a l loops - begins
- V e n t r a l l o n g i t u d i n a l m.
M id -v e n tra l-
L a te ra l longitudinal m
M id -d o rs a
S p ira l decussation-
M id-ventral-
M i d - d o r s al -
G reate r curvature
C ircular fibe rs
of s t o m a c h
of s t o m a c h -
O biigue fibers _
o f stom ach
F ig . 1 3 - Musculature of esophagus.
A. Outer layer, cranial end, lateral ventral aspect.
B. Outer layer, cranial end, lateral dorsal aspect.
C. Inner layer, cranial end. (Esophagus opened on left side.)
D. Outer layer, caudal end, ventral aspect.
E. Outer layer, caudal end, dorsal aspect.
F. Inner layer, caudal end. (Esophagus opened on right side.)
A bdom en 667
posed of a superficially cornified stratified squa in the amount of the cervical part supplied by
mous epithelium which contains the openings, the pharyngoesophageal and recurrent laryn
at about 1 mm. intervals, of the ducts of the geal nerves. It is further evident from physio
esophageal glands. In the collapsed esophagus logical experiments that these two nerves are
it forms large and numerous longitudinal rugae reciprocal in the amount of the cervical portion
or folds. Cardiac glands exist in the distal part of the esophagus that they supply.
of the esophagus. Hwang, Grossman, and Ivy also found that a
small branch from the pharyngeal branch of the
Vessels of the Esophagus vagus passed under the pharyngeal muscles to
reach the esophagus. Stimulation of this branch
The arteries to the cervical portion of the produced contraction of the cranial half of the
esophagus are primarily twigs from the cranial cervical portion. It is probable that the recur
and caudal thyroid arteries. The glandular man rent laryngeal nerves carry the afferent (Chau-
tle underlying the annular fold is richly supplied veau 1886) and some motor fibers to the cervical
by branches from the cranial thyroid artery. In portion of the esophagus as well as providing
the region of the thoracic inlet on the left side, both the motor and sensory nerve supply to the
a long but small descending branch from the thoracic part as far caudally as the heart. The
left caudal thyroid artery anastomoses with an dorsal and ventral branches of the vagi and
ascending branch from the bronchoesophageal. the vagal trunks they form supply the esophagus
From this small anastomotic trunk, branches go caudal to the heart.
to the esophagus. The esophageal portion of the
bronchoesophageal artery is the main source of ABDOMEN
blood to the cranial two-thirds of the thoracic
portion of the esophagus. The remaining part is The abdomen (abdominal region, or regio ab
supplied by esophageal branches of the aorta dominis) (Figs. 13-9, 13-10, 13-11) is that part
and/or dorsal intercostal arteries and the termi of the trunk which extends from the diaphragm
nal portion is supplied by the esophageal branch to the pelvis. It contains the largest cavity in the
of the left gastric artery. body, the abdominal cavity (cavum abdominis).
The veins which drain the esophagus are es Caudally, the abdominal cavity is continuous
sentially satellites of the arteries which supply with the pelvic cavity, the division between the
it. Those veins from the thoracic portion empty two being a plane through the pelvic inlet, or
largely into the azygos vein, with the vein which brim of the pelvis. The abdominal cavity is a
accompanies the esophageal branch of the left muscle- and bone-bounded cavity. It is lined
gastric artery being a tributary of the portal internally by the transversalis fascia, which in
system. Adjacent veins, like the arteries, anasto turn is covered in most places by the perito
mose with each other on the esophagus. Ac neum.
cording to Baum (1918), lymph vessels from the The peritoneum forms the potential perito
esophagus drain into the medial retropharyn neal cavity (cavum peritonei), which is con
geal, deep cervical, cranial mediastinal, bron tained largely, but not exclusively, within the
chial, portal, splenic, and gastric lymph nodes. abdominal cavity. In both sexes the pelvic
cavity contains the pelvic portion of the perito
neal cavity. Vaginal processes, always well
Nerves of the Esophagus developed in the male and usually present in the
female, also exist as extra-abdominal portions.
The cricopharyngeal muscle and the cervical The abdominal cavity is truncate and cone-
portion of the esophagus are supplied with mo shaped, with a concave cranial end or base. It
tor fibers from the small pharyngoesophageal contains the abdominal viscera, which include
nerve which leaves the vagus just above the primarily the flexuous alimentary canal and its
nodose ganglion (Hwang, Grossman, and Ivy two associated, indispensable glands, the liver
1948). These investigators also found that a and the pancreas. A large portion of the genito
variable portion of the cervical part of the urinary system is located here, as well as por
esophagus contracted when the peripheral ends tions of the endocrine, vascular, and nervous
of the vagus or the recurrent laryngeal nerves systems, including the spleen, adrenal glands,
were stimulated at the base of the neck. Most and many nerve plexuses, vessels, and lymph
nerve twigs to the esophagus are too small to be nodes. It is bounded cranially by the diaphragm;
seen in gross. There is probably an overlapping dorsally by the lumbar vertebrae, the sublum-
F ig . 13-9. Viscera of the dog. (The location of the diaphragm is indicated by a dotted circle.)
A. Viscera of male dog, left lateral aspect. B. Viscera of female dog, right lateral aspect.
1. Left lung 1. Right lung
2. Heart 2. Heart
3. Liver 3. Liver
4. Stomach 4. Stomach
5. Left kidney 5. Right kidney
6. Ureter 6. Ureter
7. Bladder 7. Bladder
8. Urethra 8. Urethra
9. Rectum 9. Rectum
10. Greater omentum covering small intestine 10. Greater omentum covering small intestine
11. Spleen 11. Cecum
12. Descending colon 12. Descending duodenum
13. Ductus deferens 13. Right uterine hom
14. Left testis 14. Right ovary
15. Prostate 15. Vagina
16. Thymus
A bdom en 669
Costal arches
X i p h o i d regio n
L . h g p o c h o n d r i o c r e g i on
U mb i I i c us
jh' l ~ L . l a t e r a l r e g i o n
!~ \ A ~ U m b i l i c a l reqion
j / \ \ ~ L. i n g u i n a l re g io n
P u b ic re g io n
bar muscles, and the crura of the diaphragm; middle gluteal muscles bilaterally, and the
bilaterally by the diaphragm, the two oblique ischial arch ventrally. The floor is formed in part
and transverse abdominal muscles, and a small by the coccygeal muscles but mainly by the
portion of the shaft of the ilium on each side. pubes and ischii, which lie largely peripheral to
Ventrally, the right and left rectus abdominis these thin muscular sheets. The pelvic cavity
muscles and their sheaths form the abdominal contains the rectum and the urethra in both
wall. sexes, the vagina and part of the vestibule in the
There are three unpaired apertures in the female, and a part or all of the prostate in the
diaphragm: the esophageal hiatus, for passage male.
of the esophagus, vagal nerve trunks, and esoph The abdominal and pelvic cavities are lined
ageal vessels; the postcaval foramen, for passage by fascia throughout. In most places this fascia
of the postcava; and the aortic hiatus, for pas attaches to muscles or bones peripherally and
sage of the aorta, lumbar cistern, and the azygos blends with the subserous areolar tissue cen
and hemiazygos veins. Paired slitlike openings trally. The fascia was previously named accord
existing dorsal to the diaphragm are formed ing to the region or parts it covered so that parts
ventrally by the dorsal edge of the diaphragm, of it were known by such names as diaphrag
and dorsally by the psoas muscles. At these sites matic, transversalis, iliac, internal spermatic,
the pleura and peritoneum are separated only and pelvic fascia. The term transversalis fascia
by the fused endothoracic and transversalis fas (fascia transversalis) is used to include all of
ciae. The sympathetic trunk and splanchnic these fascial divisions. The subserous areolar
nerves pass dorsal to the lumbocostal arch on tissue forms the medium whereby the perito
each side. neum is united with the transversalis fascia. In
Caudally, the abdominal cavity communi obese specimens the large fat deposits around
cates freely with the pelvic cavity at the pelvic the kidneys, in the falciform ligament, in the
inlet. In the fetus, there is a relatively large pelvis, and around the vaginal rings are located
opening, the umbilical aperture, located mid- in the subserous fascia.
ventrally, which serves for the passage of the
umbilical blood vessels, the small vitelline duct, Regions of Abdomen
and the stalk of the allantois. After these struc
tures are disrupted at birth, this opening rapidly For convenience of description the abdomen
closes, forming a faint scar, the umbilicus, on is divided into nine regions by two transverse
the mid-ventral line. There is a passage on each and two sagittal planes (Fig. 13-10).
side in the caudoventral part of the abdominal The more cranial of the two transverse planes
wall, called the inguinal canal, for the passage of passes through the most caudal border of the
the vaginal process and the spermatic cord in the costal arch. This plane therefore passes through
male and the round ligament in the female. In the caudal portion of the second lumbar verte
both sexes the external pudendal vessels and the bra. The more caudal of the two transverse
genital nerve pass through the caudal part of the planes crosses the body at a level through the
inguinal canal. Another pair of abdominal open ventral cranial iliac spines, or the most cranial
ings in the caudal part of the abdominal wall are parts of the wings of the ilia. This plane bisects
the right and left vascular lacunae. The femoral the caudal part of the sixth lumbar vertebra. By
artery, vein, lymphatics, and saphenous nerve, means of these two planes the abdomen is di
surrounded by transversalis fascia, pass through vided into three segments which are named,
each vascular lacuna. from before backward, the epigastric, mesogas-
The pelvic cavity (cavum pelvis) begins at tric, and hypogastric segments (Sisson and
the pelvic inlet as a continuation of the abdomi Grossman 1953). Of these three segments, the
nal cavity, and in a broad sense is regarded as a epigastric is by far the largest, as it extends far
division of it. It is bounded dorsally by the sa forward in the thoracic cage, where it is limited
crum and first coccygeal vertebra; bilaterally, it cranially by the diaphragm. The hypogastric
is bounded in front by the ilia and behind these segment is smallest, as it ends caudally at the
by the coccygeus, levator ani and middle pelvic inlet.
gluteal muscles. It ends at the pelvic outlet, The two imaginary sagittal planes, which
which is bounded by the first coccygeal verte further divide the abdomen into the nine
bra dorsally, the sacrotuberous ligaments and regions, pass on each side midway between the
A bdom en 671
Live r
-Stomach
Descendi ng d u o d e n u m - -Spieen
Ricjht kidney -
Left kidney
L o c a t i o n of c e c u m
- G r e a t e r omentum
covering small intestin es
U reter
Rectum - - - T e s t i c u i a r a. v v.
Bladder - -
Lat. I /'g. of b l a d d e r
M edian umbi I ic a l fo ld - C a u d a l d e e p e p i g a s t r i c a v v.
ventral cranial iliac spine and the median sagit with the dorsal part of the right sublumbar
tal plane. The three parts of the epigastric seg region. The urinary bladder, nestled in the
ment are the right and left hypochondriac greater omentum but not covered ventrally by
regions (regio hypochondriaca dextra et sinistra) it, is the only visceral organ which lies in contact
and the median xiphoid or epigastric region with the abdominal wall in the pubic region.
(regio epigastrica). The middle zone or meso- All abdominal organs vary normally in size
gastric segment includes the unpaired, middle and position. The stomach, uterus, urinary blad
ventral region of the abdomen, known as the der, and spleen vary more than the other organs
umbilical region (regio umbilicalis), and the because of their ability to undergo marked
right and left lateral regions (regio lateralis dex changes in size and shape. The gravid uterus
tra et sinistra). The lateral regions are also alters the position of all the other movable ab
known as the right and left flan k (latus), and dominal organs more markedly than do any of
each contains an expansive sublumbar fossa the others. It always occupies the most ventral
(fossa sublumbalis), which forms a ventral arc position in the abdomen because it contains no
and a dorsal, straight base located ventrolateral gas and is therefore the heaviest freely movable
to the transverse processes of the lumbar verte abdominal organ. In advanced pregnancy it
brae and the iliocostalis muscle. The three parts nearly completely fills the ventral half of the
of the caudal abdominal segment are the right abdominal cavity. It is impossible to assign a
and left inguinal regions (regio inguinaiis dextra constant position for any one of the abdominal
et sinistra) and the median unpaired pubic organs, especially those which are attached by
region (regio pubica). In addition to the nine peritoneal folds. Roentgenograms and hardened
abdominal regions already named the lumbar anatomical preparations clearly reveal the nor
region (regio lumbalis) bounds the dorsal part mal variations which exist from time to time and
of the abdominal cavity, and the sacral region in different specimens.
(regio sacralis) bounds the dorsal part of the
pelvic cavity. T he P e r it o n e u m
ads, do develop in this way. The abdominal The cavities enclosed by serous membrane
part of the alimentary canal arises very early in are closed cavities, except for the peritoneal
the embryo between the two layers of perito cavity in most female animals. In the bitch there
neum which form a mid-sagittal partition sepa is an opening at the abdominal end of each
rating the abdominal part of the celom into uterine tube and thus, through the genital tract,
right and left parts. Most of the ventral part of to the outside. No organs or tissues are located
this fold becomes obliterated shortly after it in the peritoneal cavity. In life this is an almost
forms. The parts which are left go to the liver, non-existent cavity containing only enough
the stomach, and the beginning of the duode lubricating fluid to moisten the apposed perito
num. Because most of the ventral part of the neal surfaces, both between different organs and
mesentery (that ventral to the digestive tube between the organs and the parietal perito
before it has rotated) becomes obliterated, the neum.
definitive appearance of the tube suggests that Organs which lie against the walls of the ab
it migrated ventrally from the dorsal abdominal dominal or pelvic cavities and which are covered
wall; actually it develops between the two peri only on one surface by peritoneum are said to be
toneal layers. retroperitoneal. Most of these organs are small
The peritoneum serves to reduce friction be and are embedded in fat. Organs which project
tween parts. A small amount of viscous fluid is freely into the abdominal, pelvic, and scrotal
produced for this purpose. Whenever two peri cavities and receive a nearly complete covering
toneal surfaces in contact fail to move for an of peritoneum are termed intraperitoneal.
appreciable time, the mesothelium of the ap The common dorsal mesentery (mesenterium
posed surfaces is absorbed and the connective dorsale commune) is the peritoneal fold which
tissue re-enforcement, the stroma, ceases to be leaves the dorsal abdominal wall and reflects,
differentiated from that of adjacent parts; thus, directly or indirectly, around most of the freely
peritoneal sheets become obliterated and vis movable organs of the abdominal cavity. It can
cera adhere to neighboring structures. From a be demonstrated by grasping the abdominal
relatively simple disposition in the embryo, the part of the digestive tube along with the pan
arrangement of the peritoneum in an adult be creas and spleen and moving them ventrally. In
comes a complicated maze of mesenteries and thin dogs it can be seen to leave the aorta, pro
ligaments. viding a route by which the celiac, cranial, and
The peritoneum may be divided into: caudal mesenteric arteries, autonomic nerves,
The parietal peritoneum (peritoneum parie- lymphatics, and radicles of the portal vein pass
tale), which covers in large part the inner sur to or from the intestine and other organs. It is
face of the walls of the abdominal, pelvic, and the retained part of the serous partition which
scrotal cavities. divided the celom dorsal to the alimentary
The visceral peritoneum (peritoneum vis- canal into right and left halves. At an early stage
cerale), which covers the organs of the abdomi in development the alimentary canal is rela
nal, pelvic, and scrotal cavities, wholly or in tively straight, thus allowing the common dorsal
part. mesentery to exist as a simple median partition.
The connecting peritoneum, which consists When the parts of the digestive tube differenti
of double sheets of peritoneum extending be ate, those parts of the common dorsal mesen
tween organs or connecting them to the parietal tery going to the several parts receive specific
peritoneum. These peritoneal folds are referred names, as follows: to the stomach, mesogas-
to as mesenteries, omenta, or ligaments. A mes trium; to the duodenum, mesoduodenum; to the
entery (mesenterium), in a restricted sense, jejunoileum, mesojejunoileum; to the colon,
passes from the abdominal wall to the intestine. mesocolon; and to the rectum, mesorectum. In
It is wide and contains many vessels. In a fat dogs, the apposed surfaces of the two layers
broader sense, a mesentery is any wide serous of peritoneum of the common dorsal mesentery
fold which attaches organs to a wall and serves are separated by fat which is deposited along
as a route by which the nerves and vessels reach the arteries, and because of this, neither the
the organs. An omentum passes from the stom abdominal aorta nor the caudal part of the post
ach to other organs or to a wall. A ligament cava is visible.
passes from a wall to an organ, or from an organ A transverse section of the abdomen just
to an organ, and is usually narrow and contains cranial to the tuber coxae reveals the disposi
few vessels. tion of the peritoneum in its simplest form.
674 C h apter 13. T h e D ig e s tiv e S y s te m and A bdom en
Tracing the peritoneum to the right from the the first 2 cm. after entering the pelvic cav
inner surface of the left rectus abdominis, one ity the peritoneum reflects dorsally on the neck
finds it is closely united by transversalis fascia of the bladder of the female or on the prostate
to the inner sheath of that muscle. At the linea gland of the male, forming the shallow pubo
alba it is thrown into an extremely delicate plica vesical excavation (excavatio pubovesicalis).
about 3 cm. high, the middle umbilical fold After reflecting around the cranial surface of
(plica umbilicalis medialis). In the fetus, this the bladder the peritoneum leaves the dorsal
fold contained the stalk of the allantois, but in surface of its neck in the female or the prostate
the adult no trace of this fetal structure exists. gland in the male and, forming an acute angle
After covering the right rectus sheath the in the male caudal to the farthest caudal exten
peritoneum runs dorsally on the lateral abdomi sion of the pubovesical excavation, reflects
nal wall. Dorsolaterally, it sends a thin, narrow cranially on the ventral surface of the rectum to
plica ventromedially, which surrounds the ex form the rectovesical excavation (excavatio
ternal spermatic vessels in the male. It covers rectovesicalis).
the variable amount of fat occupying the groove The caudal extension of the rectovesical ex
lateral and ventral to the sublumbar muscles cavation is not as great in the female as in the
and continues medially in the male to approxi male, and it differs also in that it is divided into
mately the mid plane. In its course across the ventral and dorsal parts by the presence of the
sublumbar muscles it loosely covers the ureter vagina and uterus and the right and left broad
as the latter crosses the external iliac vessels. ligaments, which attach these organs to the pel
Near the mid plane it covers the right sympa vic walls laterally. In this way, the vesicouterine
thetic trunk, and opposite the body of the sev excavation (excavatio vesicouterina) is formed
enth lumbar vertebra it leaves this bone and ventrally, and the rectouterine excavation (ex
runs ventrally to the descending colon. cavatio rectouterina) is formed dorsally. The
After nearly completely encircling the colon caudal angles of the reflections of the perito
the peritoneum runs back to the vertebra so neum from one pelvic organ to the next dorsal
that, between the dorsal abdominal wall and to it are located progressively farther caudally
the colon, it forms a fold, consisting of two lay in the male when they are traced in parasagit
ers of peritoneum. Between these layers is lo tal planes from the pubis to the tail. A line
cated a small amount of transversalis fascia, the drawn from a point about 1 cm. caudal to the
caudal mesenteric vessels, and the lumbar cranial border of the pubis to the transverse
colonic nerves. This fold and its contents con process of the third coccygeal vertebra indicates
stitute the descending mesocolon (mesocolon their most caudal extensions in the male. In the
descendens). The peritoneum, in running from female, the rectouterine excavation extends
the descending mesocolon across the left ab farther caudally than the vesicouterine, but
dominal wall to the starting point ventrally, neither extends as far caudally as does the undi
covers the same structures on the left side as it vided vesicorectal excavation which takes their
does on the right. In the female the broad liga place in the male.
ment o f the uterus (lig. latum uteri) leaves each To understand the definitive positions of the
side of the sublumbar region as a peritoneal connecting peritoneum to the alimentary tract
fold to surround the uterine horn. At the level and the liver, it is helpful to know about the
under consideration, a lateral fold leaves the primary rotations the tube makes and the dif
broad ligament to surround the round ligament ferential growth that certain parts undergo.
of the uterus. Early in development, the dorsal part of the
stomach grows faster than the ventral part,
Pelvic Peritoneal Excavations with the result that it rotates on its axis to the
left. Simultaneously with this rotation the stom
The peritoneum of the pelvis will now be ach comes to lie in nearly a transverse position
traced by following it in a sagittal section just with the outlet located on the right while the
lateral to the median plane. By starting ven inlet remains in nearly the median plane. Distal
trally at a transverse plane passing through the to the stomach the anlage of the liver and pan
tuber coxae and advancing caudally, one finds creas appear as diverticula of the mucosa of the
the peritoneum runs on the sheath of the rectus duodenum. The liver grows ventrally between
abdominis and passes over the ventral part of the peritoneal layers forming the ventral mes
the pelvic inlet into the pelvic cavity. Within entery, and the pancreas grows into the dorsal
A bdom en 675
mesentery. The cecum develops as a diverticu In the postmortem specimen the omentum
lum of the lateral wall of the proximal part of appears as a fenestrated membrane.
the ascending colon. The bursal portion (pars bursalis) of the
The abdominal part of the alimentary canal greater omentum attaches cranioventrally to
grows many times the length of the abdominal most of the greater curvature of the stomach.
cavity. Therefore, the parts with long mesen Always closely related to the ventral abdominal
teries, mainly the jejunum and to a lesser extent wall, it extends caudally as far as the urinary
the ileum, become greatly folded and occupy bladder. Regardless of the fullness of the blad
no constant location in the abdominal cavity. der, the bursal portion passes dorsally between
During development a part of the tube, in the that organ and the intestinal coils, and then re
form of a U-shaped loop, is accommodated in a flects upon itself and returns to the region dor
normal umbilical hernia which lasts but a short sal to the stomach, intimately covering the
time. The cranial mesenteric artery lies in the jejunal coils in its course. Because of this folding
mesentery which connects the two limbs of the of the greater omentum, the intestinal coils are
loop. As the loop forms it rotates about three- covered by two layers of the greater omentum.
fourths of a circle in a counterclockwise direc The more superficial, ventral layer is the
tion, as viewed ventrally, around the axis of the parietal part (pars parietalis), and the deeper,
artery. Because of this rotation, the proximal dorsal layer is the visceral part (pars visceralis).
part of the large intestine hooks around the Between these two layers is a potential cavity,
cranial mesenteric artery from the left and the the lesser peritoneal cavity, or the omental
small intestinal mass moves caudal to the axis bursa. Except for the numerous fenestrae or
of the artery by passing to the right of it. windows in each layer of the wall of the omen
The various derivatives of the common dor tal bursa and the large constant opening, the
sal mesentery and the ventral mesentery will epiploic foramen, the omental bursa is a closed
now be described as they run between the ab sac. The epiploic foramen is bounded ventrally
dominal wall and the various parts of the ali by the peritoneum covering the portal vein and
mentary canal. dorsally by that covering the postcava. The
The peritoneal folds which, in the adult, greater omentum is considerably longer and
leave approximately the greater and lesser wider than the abdomen; to accommodate this
curvatures of the stomach are known as the bulk, it is folded in around the margin of the
greater and the lesser omentum, respectively. intestinal coils. These infoldings usually occur
The greater omentum, the most specialized as follows: caudally, cranial to the bladder; on
serous fold in the body, is derived from the the right, medial to the descending portion of
dorsal mesogastrium. The lesser omentum, the duodenum; and on the left, ventral to the
small and relatively simple, is that portion of sublumbar muscles, left kidney, and the fat
the ventral mesogastrium which extends be which surrounds these structures.
tween the liver and the lesser curvature of the The splenic portion (pars lienalis) is also
stomach and the initial part of the duodenum. called the gastrosplenic ligament (lig. gastro-
lienale). Although Zietzschmann (1939) con
Greater Omentum siders this portion as extending beyond the ends
and hilus of the spleen, in this description the
The greater omentum (omentum majus) term will be limited to that portion of the
(Figs. 13-11, 13-13) usually reaches as a lacy greater omentum which extends from the dia
apron from the stomach to the urinary bladder, phragm, fundus, and greater curvature of the
covering the intestinal coils ventrally and on stomach to a line which parallels the hilus of
the sides. It is unequally divided into the large the spleen. The spleen is not located between
bursal portion and the smaller splenic and veil the two peritoneal layers which form the gastro-
portions (Zietzschmann 1939). Each portion is splenic ligament but is interposed in an out-
composed of a double peritoneal sheet in which pocketing of the superficial peritoneal layer
are streaks of fat, largely located around the which forms the parietal sheet. When taut, the
various-sized arteries which run through it and gastrosplenic ligament is about 5 cm. wide ven
forming one of the major fat storehouses in trally. The large splenic vessels and nerves
obese specimens. Between the finer streaks of which approach the spleen near the middle of
fat the peritoneum is transparent because of its its hilus give origin to the left gastroepiploic
thinness.
676 C h apter 13. T h e D ig e s tiv e S y s te m and A bdom en
V e n t r a l l e a f of Lesser om en tu m
gr. o m e n t u m
Ventral l e a f a f -
gr. o m e n t u m D o rs a l le a f of
D orsal le a f of
gr. o m e n t u m
gr. o m e n t u m ^ ---- S p le e n
\ Gastrosplenic -
Pancreas
M esoduodenum __ lig. L o c a tio n o f
(in d o rs a l le a f)
p o r t a l v.
f S p le n o c o lic lig .
Pancreas -
Du o d en o jeju n al
Root of
fle xu re
M e se n te ry - m esentery
c o m m o n to
P a n c rea s --
duad. * colo n M e s o c o lo n
( in m e s o d u o d .)
x Cut edge a f
D u o d en o c o lic Cut edge of m es o je ju n o ile u m
m e s o je ju n o ile u m
A B
G o llb la d d e r C o r o n a r y li g .
D ia p h ro g m ~
Cut edge of
L ive r le s s e r omen. F a lc ifo rm lig - _ - L e s s e r omen.
Common — P o stcava «- - L. t r i a n g u l a r
Esophagus lig . o f l i v e r
b ile duct
C e l i a c a. R t. t r i a n g u l a r
P o r t o l v. - - D orsal
li g . o f l i v e r
C ranial m eso g astriu m
H e p a tic a . ' \ m e s e n t e r i c o.
" Mesoduodenum
L o ca tio n o f ' \v
e p ip lo ic fo ra m e n Root of H e p a t o r e n a l lig. •M e so je jun oileu m
m esentery
P o stcava ■■ - M e s o c o l o n
Caudal
m e s e n t e r i c a.
D
F ig . 13-12. Peritoneum.
A. Plan of visceral and connecting peritoneum, ventral aspect. The greater omentum is transected caudal to the
stomach. Arrow in omental bursa.
B. Plan of peritoneum with greater omentum reflected cranially. The transverse colon is displaced caudally.
C. Plan of the dorsal reflections of the connecting and parietal peritoneum. The stomach and intestines removed.
D. Plan of the dorsal reflections of the connecting and parietal peritoneum. All abdominal viscera removed.
Abdo m en 677
Spleen
Pancreas'
D esc. colon
R. k i d n e y
I \
Epiploic foramen' j f R e n a l v.1 / ^ S \ L. k i d n e y
Postcava/ L P ostcava/ yR o o t o f
A orta
A z y g o s vei n B H m esentery
talis) is the main cavity of the omental bursa. dorsal wall of the body of the stomach just caudal
It is enclosed by the bursa portion of the greater to the attachment of the lesser omentum.
omentum and extends caudally and laterally The various peritoneal folds which attach the
from the stomach to the urinary bladder. The liver and digestive tube to the abdominal wall
two laminae of the greater omentum do not or to other viscera are treated in the descrip
fuse with each other caudally, thus reducing the tions of the several organs.
size of the omental bursa, nor does any portion
of it attach to the colon as it does in some other
species (e.g., man and horse). STOMACH
The omental bursa is closed caudally by the
parietal wall reflecting upon itself to form the The stomach (ventriculus [gaster]) (Figs.
visceral wall. Cranially, the closure of the omen 13-14, 13-15), the largest dilatation of the ali
tal bursa is more involved. The dorsal wall of mentary canal, is a musculoglandular organ
the stomach and the liver largely fill the interposed between the esophagus and the small
“mouth” of the bursa. At the cardia, the inner intestine. An axis through it is shaped some
peritoneal layer of the greater omentum be what like the letter C rotated 90 degrees in a
comes continuous with the similar layer of the counterclockwise direction. It varies greatly in
lesser omentum as the two layers are connected size. It stores and partly mixes the food, while
by the visceral peritoneum covering the dorsum its intrinsic glands intermittently add enzymes,
of the cardia. On the right side the inner peri mucus, and hydrochloric acid (Alvarez 1940),
toneal layers of the two omenta converge on the The stomach lies largely in a transverse posi
medial surface of the cranial part of the duo tion, more to the left of the median plane than
denum, thus closing the bursa at this site. At to the right of it. It forms an extensive concavity
other places around its periphery the inner peri in the caudal surface of the liver, and when it
toneal layers of the visceral and parietal sheets is empty it is located completely cranial to the
become continuous. The outer peritoneal layers thoracic outlet (Zietzschmann 1938). The inlet
of the two walls are also continuous except on of the stomach is called the cardia, and the out
the left, where the veil portion of the greater let the pylorus. The major divisions of the stom
omentum is formed. ach are the cardiac portion, fundus, body, and
The splenic recess (recessus lienalis) is largely pyloric portion. It possesses dorsal and ventral
non-existent unless the abdominal cavity is walls, and greater and lesser curvatures.
opened and the spleen displaced. In such in The dorsal wall (facies visceralis) presents a
stances, it is a recess of the omental bursa oppo convex outer surface which faces mainly dor
site the hilus of the spleen. sally, but also caudodextrally. It lies in contact
Three folds invaginate the inner peritoneal with the left lobe or limb of the pancreas and is
sheet of the omental bursa or the whole bursa separated from the intestinal mass and left
wall. These are formed by the three branches of kidney by the visceral or dorsal leaf of the
the celiac artery and the nerve plexuses which greater omentum. The ventral wall (facies pari-
surround them. The right gastropancreatic fo ld etalis) faces to the left and cranially as well as
(plica gastropancreatica dextra) is a low peri ventrally. In the contracted state the stomach
toneal fold which contains the common hepatic lies in contact with the liver, in which it pro
artery. It extends obliquely across the medial duces an extensive gastric impression. The di
face of the portal vein just within the vestibule lated stomach extends beyond the liver chiefly
from the epiploic foramen. The largest fold is to the left and ventrally.
formed by an upward displacement of the bursa
wall produced by the splenic artery and nerves Curvatures of Stomach
and their continuation opposite the hilus of the
spleen as the left gastroepiploic artery and The greater curvature (curvatura ventriculi
nerve plexus. The left gastropancreatic fo ld major) is convex, and extends from the cardia
(plica gastropancreatica sinistra) is formed by to the pylorus. It is approximately 30 cm. long
the left gastric artery and nerve plexus. It is in a stomach moderately filled. The parietal leaf
short and extends from the celiac artery to the of the greater omentum attaches to the greater
left extremity of the lesser curvature of the curvature except on the left, where its line of
stomach. Surrounded by fat, it continues on the attachment runs obliquely across the dorsal wall
680 C h ap ter 13. T h e D ig e s tiv e S y ste m and A bdom en
/ Dorsal l e a f
g r e a t e r o m e n t u m (cut )
Common b i le duct
P o s i t i o n of ve n tra l
d u o d e n a l p a p i I la
Dorsal p a n c r e a t i c d u c t
~ ~Stomach
P o s itio n of d o r s a l -
duodenal p a p illa
- L e f t l o b e of
R t lobe of p a n c r e a s ~ pancreas
c o v e rin g r t kid ney
■ Tr ansver se c o l o n
Ascendinq colon-' W
Cecum
D e s c e n d i n g colon
Duodenum k i dnetj
R o o t o f m e s e n t e r y (cut)
F ig . 13-14. Abdominal viscera, ventral aspect. The pancreas in situ; the position of the kidneys indicated by a dotted line.
Esophagus-
Common b i le d u c t
H epatic ducts-
P y lo r ic ostium
I n t r a m u r a l p o r t i o n of d u c t -
Dorsal duodenal p a p i l l a -
of the stomach to form, with the lesser omen distal third is contracted and bent so that the
tum at the cardia, a closure of the omental greater curvature (caudal side) is three or four
bursa at this site. times longer than the opposite side. It is largely
The iesser curvature (curvatura ventriculi surrounded by a heavy, double sphincter which
minor) also runs from the cardia to the pylorus, forms the narrowest part of the cavity of the
and is the shortest distance between these two stomach. The passage through the pyloric part
parts. It does not form an even concavity but is is known as the pyloric canal (canalis pyloricus).
in the form of a 50 to 70 degree angle, the angu
lar notch (incisura angularis). Lying within this Shape, Position, and Capacity
angle is the papillary process of the liver. The
pyloric antrum and pylorus lie to the right of The empty, contracted stomach is not only
the papillary process, and the body of the stom contained within the caudal part of the thoracic
ach lies to the left of it. The caudal edge of the cage, but it is also nestled within the caudal con
lesser omentum attaches to it. cavity of the liver, being completely separated
from the abdominal wall. When the stomach
Regions of Stomach increases in size as the result of filling, the
fundus enlarges first and pushes caudodorsally.
The cardiac portion (pars cardiaca) of the It tends to displace the liver ventrally as the
stomach is that portion which blends with the stomach comes in contact with the left lateral
esophagus. The four coats of both organs blend abdominal wall and diaphragm. The body of the
with each other. The greatest differences occur stomach is the second part to fill and expand.
in the muscular and mucous layers as these are It is the largest division of the organ, as well as
traced from the esophagus to the stomach. the part capable of the greatest dilation. During
The fundus of the stomach (fundus ventric filling, it migrates caudoventrally and makes
uli) is the rather large blind outpocketing extensive contact with the abdominal wall. It
located to the left and dorsal to the cardia. The is particularly distendable in puppies, and
esophagus joins the stomach in such a way that when maximally expanded it may extend from a
on the right its surface continues with that of transverse plane through the eighth thoracic
the lesser curvature of the stomach without vertebra to a plane caudal to the umbilicus.
definite demarcation. On the left the cardiac This necessitates an expansion of the abdomen
notch (incisura cardiaca) is formed between the and a crowding of the intestinal mass and spleen
cardia and the bulging fundic part. The body caudally and slightly dorsally. According to
of the stomach (corpus ventriculi) is the large Grey (1918) the normal stomach possesses a
middle portion of the organ. It extends from the striking capacity to adjust its size to the volume
fundus on the left to the pyloric portion on the of its contents with only minimal changes in
right. If two transverse planes are projected intragastric pressure.
through the stomach at right angles to its curved Secord (1941) states that food remains in the
long axis, one through the caudal face of the dog’s stomach from 10 to 16 hours. Zietzsch
cardia above and the other through the angular mann (1938) states that the pylorus varies least
notch below, these planes will limit the begin in position as the stomach fills. It is always more
ning and end of the body of the stomach. The cranial and ventral than the cardia. The pyloric
shortest path which ingesta can take in passing portion is the last part to expand, the antrum
from the cardia to the pyloric portion of the expanding more than the canal. The pyloric por
stomach is known as the gastric groove (sulcus tion functions chiefly as an ejection mechanism
ventriculi). This path follows the lesser curva by which the partly digested stomach contents,
ture of the stomach. the chyme, is forced through the pyloric canal
The pyloric part (pars pylorica) (Figs. 13-15, and squirted into the duodenum. The empty or
13-16) is approximately the distal third of the the partly filled stomach is shaped like a C, with
stomach as measured along the lesser curvature. its convex surface facing caudoventrally and to
It is always somewhat sacculated as it unites the the left.
body of the organ to the duodenum. The pyloric The capacity of the stomach varies from 0.5
part is irregularly funnel-shaped toward the to 8 liters. Greater ranges in relative size are
pylorus, which is directed cranially. The initial present in puppies than in adult dogs. Ellen
two-thirds is thin-walled and expanded to form berger and Baum (1943) cite Neumayer, who
the pyloric antrum (antrum pyloricum). The gives the capacity as 100 to 250 cc. per kilogram
of body weight.
682 C h ap ter 1 3 . T he D ig e s t iv e S y stem and A bd o m en
Ci r c u l a r f i b e r s Esophagus
!
t Cardia
Angular in cisu re , '
P y /orus-
-L o n g itu d in a l fibers
The average empty stomach of a 15-kg. dog the angular incisure there are no longitudinal
weighs about 100 gm. fibers so that the circular fibers of the inner
coat become superficial. As the longitudinal
fibers cover the pyloric portion they are particu
Coats of Stomach larly heavy on the sides between the curvatures,
but no definite pyloric ligaments are formed, as
The serous coat (tunica serosa) completely in man.
covers the stomach except for an extremely The inner circular layer (stratum circulare) of
narrow line on the distal half of the greater the stomach is more complete and specialized
curvature which continues obliquely across the than is the longitudinal layer. At the cardia the
dorsal surface of the stomach to the cardia. A circular layer is thickened to form the feeble
second similar line on the stomach which is not cardiac sphincter (m. sphincter cardiae). This
covered by peritoneum extends from the cardia sphincter is augmented on the greater curva
along the lesser curvature to the duodenum. ture by the acquisition of a condensation of the
The smooth peritoneal sheets of serous mem transversely running inner oblique fibers. At
brane which intimately cover the dorsal and the approximate junction of the fundus and the
ventral walls of the stomach fuse just distal to body on both the dorsal and ventral walls the
these lines to form the greater and lesser circular coat enters into the formation of a
omenta. In obese specimens a small irregular muscular fiber interchange with the longitu
strip of fat and the nerves and vessels serving dinal coat. The circular coat is not covered by
the organ widen the line along which the the longitudinal coat in and adjacent to the
omenta arise. The two peritoneal layers which angular incisure so that in this region it receives
form these folds separate at the cardia to extend a peritoneal covering, superficially. Deeply,
forward on the abdominal portion of the esoph throughout the length of the lesser curvature
agus. At the duodenum the peritoneal leaves are from the cardia to the pyloric antrum, there is
similarly forced apart by the duodenum. The a muscular trough, about 2 cm. wide, which
peritoneal leaves of the lesser omentum become is formed on the sides by the parallel longi
continuous with those of the mesoduodenum tudinal parts of the oblique fibers and deeply by
over the common bile duct. The serosa of the the circular fibers.
stomach is extremely thin and elastic. It ad Surrounding the pyloric canal, the inner circu
heres closely to the stomach musculature by a lar layer is well developed, as Torgersen (1942)
scanty amount of subserous tissue. has pointed out. The musculature is thickest as it
The muscular coat (tunica muscularis) of the crosses the greater curvature. The pylorus, which
stomach (Fig. 13-16) consists essentially of an opens into the duodenum, is also surrounded by
outer longitudinal and an inner circular layer a circular muscle termed the pyloric sphincter
of smooth muscle fibers. To these layers are (m. sphincter pylori).
added oblique fibers over the body of the stom The oblique fibers (fibrae obliquae) are adja
ach. cent to the submucosa. They appear to arise
The outer longitudinal layer (stratum longi- from a heavy transverse stratum which is arched
tudinale) is continuous with the essentially across the dorsal (greater curvature) boundary
outer longitudinal layers of both the duodenum of the cardiac orifice. These fibers run distally
and the esophagus. The longitudinal fibers on and outward toward the pylorus and the greater
and adjacent to the lesser curvature as traced curvature in each wall. The oblique fibers thus
from the esophagus spray out and end before are spread like a fan; in a moderately distended,
reaching the angular incisure. Those fibers on medium-sized stomach, those bundles nearest
the dorsal and ventral walls of the body of the the lesser curvature are essentially parallel. The
stomach end before reaching the middle of the fibers next peripheral to the parallel ones fan
body, whereas the longitudinal fibers on out and end on the inner surface of the circular
the greater curvature continue uninterruptedly fibers of the distal part of the body of the stom
from the esophagus to the duodenum. On both ach; the most proximal oblique fibers become
the dorsal and ventral walls of the stomach at almost transverse proximally and blend with
about the junction of the fundus and body there those of the circular layer in augmenting the
is a kind of muscular whorl formed by the size of the dorsal part of the feeble cardiac
bundles of fibers of the longitudinal layer chang sphincter.
ing position and direction. At and adjacent to The submucous coat (tela submucosa) con
684 C h apter 1 3 . T he D ig e s t iv e S ystem and A bd o m en
sists of a strong but thin elastic layer of areolar lamina muscularis mucosae and the blind ends
tissue which more firmly attaches to the mucosa of the glands. According to these authors the
than to the muscularis. It contains the finer lamina propria contains the gastric glands, and
branches of the gastric vessels and nerves. In in certain areas, exclusive of the cardiac gland
the contracted organ it is thrown into folds zone, the glands are divided into groups by
which occupy the centers of the relatively in heavier strands of supporting tissue which con
elastic plicae of the mucous coat. tain muscle fibers from the lamina muscularis
The mucous coat (tunica mucosa) consists of mucosae. Three types of gastric glands are
a columnar surface epithelium, a glandular recognized in the dog. These are the cardiac
lamina propria, and a lamina muscularis mu glands, gastric glands proper, and the pyloric
cosae consisting of muscular fibers which may glands.
be irregularly interwoven or stratified (Traut- The cardiac glands (glandulae cardiacae),
mann and Fiebiger 1957). In the contracted according to Haane (1905), are found in a nar
empty or even a moderately distended organ the row zone around the cardia. Cardiac glands are
mucosa and much of the underlying submucosa also scattered along the lesser curvature, ac
are thrown into folds, the plicae gastricae. These cording to Ellenberger (1911).
folds are largely longitudinal in direction and The gastric glands proper (glandulae gastricae
very tortuous except adjacent to the lesser [propriae]), or fundic glands, occupy about
curvature, where the folds are less crowded and two-thirds of the gastric mucosa. This includes
are relatively straight. In a strongly contracted the left extremity, or fundus, and the body of
stomach the mucosal folds, which may be 1 cm. the stomach. It is exclusive of the pyloric part
high, lie closely adjacent to each other. The nor and the cardiac gland region.
mal color of the mucosa in the body and fundus The pyloric glands (glandulae pyloricae) are
of a fresh stomach is pink to grayish red. In the found in the pyloric part of the stomach. Be
pyloric region it is lighter in color. The color tween the pyloric and gastric glands proper,
varies with the amount of contained blood, as according to Bloom and Fawcett (1962), there
well as the freshness of the material. exists in the dog a zone of intermediate glands
Under magnification the mucosa is seen to which reaches a width of 1 to 1.8 cm. Harvey
possess about 40 raised areas for every cubic (1906) states that when the stomach is flattened
millimeter. These are the areae gastricae. In the out the intermediate zone is 2 to 3 cm. wide.
pyloric region these areas are elongated in a The difference between the various gland zones
longitudinal direction, but elsewhere they are is in the type of cells they contain, and there
polygonal and rendered distinct by small sur fore in the nature of the secretion they produce.
rounding furrows. Each of the small gastric The mucosal regions of the stomach are not co
areas- on their surfaces and sides is stippled extensive with the gross divisions of the organ
with numerous minute openings, the foveolae with the same or comparable names. There is
gastricae. The foveolae are longest, about 0.68 considerable intermixing of the glands of each
mm. in the pyloric region. They gradually gland area with those of adjacent areas. The
shorten toward the cardia and disappear, so intermediate gland zone is formed in this man
that none exist in the cardiac gland region ner. The cardiac glands of the stomach are
which is adjacent to the esophagus (Mall 1896). similar to the cardiac glands in the distal portion
According to Mall, there are 1,000,000 foveolae of the esophagus. The pyloric glands imper
in the stomach of the dog, and each of these has ceptibly blend with those of the duodenum and
about 16 gastric glands opening into it. as they do they come to lie in the submucosa.
Both the type of cells in the gastric glands and
their morphology and location have influenced
Glands of Stomach the naming of the mucosal areas of the stomach.
According to Bloom and Fawcett (1962), four
The glands of the stomach are known as the types of glandular cells are found in the stomach
gastric glands (glandulae gastricae). They are mucosa. These are: (1) chief, or zymogenic,
branched tubular glands with necks and bodies cells, which contain granules that are believed
which reach nearly to the lamina muscularis to contain pepsinogen, the precursor of the
mucosae. According to Trautmann and Fiebiger chief gastric enzyme, pepsin; (2 ) parietal cells,
(1957), in older carnivores a double-layered which are spherical or pyramidal cells lying next
lamina subglandularis intervenes between the to the basement membrane, and are considered
S m a l l I n t e s t in e 685
the source, probably indirectly, of the hydro tric nodes. A few lymphatics from the stomach
chloric acid of the gastric juice; (3) mucous drain into the right hepatic node after first hav
neck cells, which are located in the necks of the ing passed through the duodenal node.
gastric glands, filling the spaces between the
parietal cells, and which produce mucus; (4)
argentaffin cells, which are moderately abun Nerves of Stomach
dant in the proper gastric glands and less fre
quent in the pyloric glands, and which are The stomach is supplied by parasympathetic
numerous in the first part of the small intestine. fibers from the vagi and by sympathetic fibers
The glands of the cardiac, intermediate, and from the celiac plexus. The ventral vagal trunk,
pyloric regions function mainly to produce after passing through the esophageal hiatus,
mucus; the proper gastric glands produce hy immediately sends two to four small branches
drochloric acid indirectly and the enzyme pep to the pylorus and liver. Other branches go to
sin. As in most other areas of the alimentary the lesser curvature of the stomach. The dorsal
canal, lymph nodules are scattered throughout vagal trunk sends branches to the lesser curva
the mucosa of the stomach. Some of these ex ture also, and to the ventral wall of the stomach.
tend through the lamina muscularis mucosae According to Jemerin and Hollander (1938) the
into the submucosa (Bensley 1902). vagi of the dog do not form observable esoph
ageal or gastric plexuses. The sympathetic fibers
to the stomach reach it by traveling on the
numerous gastric branches of the celiac artery.
Vessels of Stomach They come from the celiac plexus.
through the tuber coxae, makes a U-shaped to the deep circumflex iliac vessels. It is con
turn, and runs obliquely craniosinistrally to be tinued by a rounded angle which is somewhat
continued by the jejunum to the left of the root less than a U-turn as the ascending portion of
of the mesentery. This loop formed by the duo the duodenum diverges slightly from the de
denum is more marked in puppies than in adults scending portion. The caudal portion of the
(Mitchell 1905). Both the pancreatic and bile duodenum is related to the terminal portion of
ducts open into the duodenum. The acid chyme the ileum and to the jejunum ventrally.
which enters it from the stomach is mixed with As the ascending portion (pars ascendens) of
the alkaline secretions from the liver, pancreas, the duodenum runs obliquely forward and to
and small intestinal glands. Because of the high the left from the caudal flexure, it crosses ob
nutritive content of the material ingested, most liquely the dorsally lying ureters, sympathetic
free-living intestinal parasites are found in the trunks, postcava, aorta, and lumbar lymphatic
duodenum. For descriptive purposes the duo trunks. Ventrally, the ascending portion is re
denum is divided into four portions. These are lated to the coils of the jejunum. On the left it
the cranial, descending, caudal, and ascending approaches the descending colon, and makes
portions. a sweeping curve ventrally to form the duo
The cranial portion (pars cranialis) of the denojejunal flexure (flexura duodenojejunalis).
duodenum arises from the pylorus in such a way At this flexure the jejunum continues the duo
that its right wall is considerably longer than denum ventrally, caudally, and to the left, and
its left wall. This configuration brings about the enters into the formation of numerous coils and
acute cranial duodenal flexure (flexura duodeni kinks (festoons), which constitute most of the
cranialis) formed by the cranial portion of the intestinal mass.
duodenum. The pyloric part of the stomach Attachments and peritoneal relations. The
runs cranially, whereas the cranial portion of first two parts or right portion of the duodenum
the duodenum runs essentially caudally. The is located in the free border of the mesoduo
cranial portion of the duodenum is also known denum. The ventral peritoneal layer of the
as the duodenal cap or bulb. It lies opposite the mesoduodenum, near the stomach, is continu
ninth and tenth ribs and the intervening inter ous with the ventral layer of the lesser omentum
costal space. Ventrally, it is separated from the over the bile duct. The dorsal peritoneal layer
stomach by the greater omentum. Dorsally and is coextensive with the dorsal layer of the lesser
laterally it lies in contact with the liver, and omentum as traced over the portal vein and
medially it is in contact with the pancreas. through the epiploic foramen. Upon leaving the
The descending portion (pars descendens) of right portions of the duodenum the two peri
the duodenum, about 15 cm. long, runs caudally toneal leaves which form the mesoduodenum
from the cranial portion nearly to the pelvic as traced to the left extend directly to the right
inlet. Its lateral surface lies in contact with the lobe of the pancreas or merge for a short dis
parietal peritoneum of the upper flank, and tance before covering this portion of the gland.
with the right lateral and the right medial lobes The peritoneal leaves merge again upon leaving
of the liver, cranially. Dorsally, the right lobe or the left side of the right pancreatic lobe and pass
limb of the pancreas lies in contact with it. to and cover the ascending part of the duo
Medially, it is related primarily to the cecum denum. Upon leaving the ascending duodenum
caudally and the ascending colon cranially, but the mesoduodenum becomes continuous with
it is separated from these by the infolded the right peritoneal leaf of the descending meso
greater omentum. colon. Caudally, the two peritoneal layers of the
The caudal portion of the duodenum is also mesoduodenum form a triangular fold with a
known as the caudal flexure of the duodenum free caudal border which runs from the meso
(flexura duodeni caudalis). It connects the de colon in the region of the pelvic inlet obliquely
scending and ascending portions from right to craniodextrally to the caudal duodenal flexure.
left and is about 5 cm. long. It lies in a frontal This triangular attachment of the initial part of
(horizontal) plane. The caudal portion usually the ascending duodenum is called the duodeno-
lies ventral to the body and right transverse colic ligament (lig. duodenocolicum).
process of the sixth lumbar vertebra. A full The duodenal fossa (fossa duodenalis) (Fig.
bladder and colon would force it cranially. The 13-17) is comparable to the inferior duodenal
initial segment of this portion, at the right, lies fossa of man. It may be absent, or, when pres
in contact with the parietal peritoneum of the ent, it may admit the end of the little finger. It
sublumbar region and the uterine hom ventral varies in length up to 4 cm. Its opening faces
Sm a l l I n t e s t i n e 687
cranially and is usually situated about 2 cm. sympathetic nerve trunks, and lumbar lym
caudal to the duodenojejunal flexure. The fossa phatics. The jejunum rarely extends into the
runs caudally along the attached border of the pelvic cavity caudally because the urinary blad
ascending duodenum. When maximally devel der and the rectum largely fill the pelvic inlet.
oped it reaches a transverse level through the The spleen, through the greater omentum, is
most cranial extension of the duodenocolic liga related to the craniosinistral part of the jejunum.
ment. This simple peritoneal pocket is bounded The ileum is the terminal part of the small in
by the duodenum ventrally and on the right, testine. In man, the distal three-fifths of the
and by the left leaf of the descending mesocolon jejunoileum is regarded as ileum, the proximal
dorsally and on the left. two-fifths as jejunum. Most veterinary anat
omists regard only the short, terminal, usually
Jejunum and Ileum contracted part of the small intestine as ileum.
In the dog, unlike man, the ileum contains
The jejunum and ileum compose the bulk of fewer aggregated lymph follicles than the more
the small intestine. The jejunum begins at the proximal part of the small intestine, including
left of or caudal to the root of the mesentery the duodenum (Titkemeyer and Calhoun 1955).
at the duodenojejunal flexure, and the ileum Its terminal part usually crosses the ventral sur
ends by opening into the initial portion of the face of the descending colon. Its termination at
ascending colon to form the ileocolic orifice the ileocolic valve is located within the duode
(ostium ileocolicum). The orifice lies usually nal loop or ventral to the ascending part of the
between the descending and ascending portions duodenum. The ileum is readily identified by
of the duodenum. In contrast to the relatively the ileocecal fold and its contained antimesen-
fixed duodenum the jejunoileum is the most teric ileal vessels.
mobile and free part of the entire alimentary
canal. It is suspended by the long mesentery Mesentery of Small Intestine
from the cranial part of the sublumbar region
and is therefore also known as the mesenteric The ileocecal fo ld (plica ileocecalis) is a nar
portion of the small intestine (pars intestinum row but usually long, variably developed plica
tenue mesenteriale). No definite gross, micro of peritoneum which continues proximally on
scopic, or developmental manifestations mark the ileum from the area of adhesion of the ce
the division between jejunum and ileum. This cum to the ileum. It is rarely over 1 cm. wide at
division was made by early investigators from its origin, and when it is double at this site it is
the gross appearance of this portion of the considerably narrower. It varies in length from
bowel. According to Field and Harrison (1947) 2 to 30 cm. A streak of fat located in its ileal at
Galen applied the term “jejunum” to the middle tachment surrounds the antimesenteric ileal
portion of the small intestine because it is vessels. Its distal end is reduced to a low ridge
usually empty or appears emptier than the rest. of peritoneum formed by the underlying ves
The first known use of the term “ileum” was by sels and fat. Frequently the vessels can be
an anonymous author. It is applied to the rela traced proximally on the ileum beyond the end
tively short, contracted terminal portion of the ing of the fold.
small intestine in the domestic animals. Al The mesentery (mesenterium) (Fig. 13-12) is
though there are distinctive differences, particu also known as the great or proper mesentery, or
larly in the mucosa, between a typical portion the mesojejunoileum, to differentiate it from the
of the jejunum and that of the ileum, there are various other portions which are derived from
no sufficiently marked gross differences in the the common dorsal mesentery. In the adult it is
character of the walls to differentiate the jeju continuous in front with the mesogastrium
num from the ileum. (deep sheet of the greater omentum) and be
hind with the descending mesocolon. Embry-
Position of Small Intestine onically, the great mesentery is continuous with
the mesoduodenum in front and with the as
The jejunum is located ventrocaudal to the cending mesocolon behind. Through rotation,
empty stomach. It is separated from the ventral torsion, and differential growth of the various
and lateral abdominal wall only by the deep and portions of the alimentary canal the definitive
superficial sheets of the greater omentum. It is continuations of the great mesentery have been
related dorsally to the large intestine, duo altered. The great mesentery is in the form of a
denum, pancreas, kidneys, postcava, aorta, large fan hanging from the cranial part of the
688 C h apter 1 3 . The D ig e s tiv e S y s te m and A bdom en
sublumbar region. In its free distal border is lo of the wall opposite the mesentery. Titkemeyer
cated the convoluted jejunum and ileum. It is and Calhoun also describe a special connective
about 20 cm. wide and 5 mm. thick. Its length tissue layer about 30 fi thick between the intes
or greatest dimension varies greatly, depending tinal glands and the lamina muscularis mucosae.
on where the measurement is taken. It is only This layer, found only in the dog, is apparently
about 1.5 cm. long at its parietal attachment to similar to the layer in a comparable location in
the aorta and diaphragmatic crura opposite the the stomach. The lamina muscularis mucosae,
second lumbar vertebra. This portion of the according to the previously mentioned investi
great mesentery is known as the root o f the mes gators, was found to be three times thicker in
entery (radix mesenterii). It is the thickest por the dog than it was on an average in the other
tion because it includes the cranial mesenteric domestic animals studied. It is definitely divided
artery, intestinal lymphatics, and the large mes into inner circular and outer longitudinal layers.
enteric plexus of nerves which surround the In the dog, the duodenal glands (glandulae duo-
artery. The free or intestinal border of the great denales) differ from the intestinal glands in that
mesentery is its longest part. It extends from they closely resemble the pyloric glands of the
the duodenojejunal flexure, proximally, to the stomach. They are located only around a nar
ileocolic junction, distally. It is as long as the row zone of the duodenum adjacent to the py
jejunoileum, or about 250 cm. in life. It is lorus. The main portion of each gland is located
greatly folded or ruffled as it follows the turns in the submucosa and a portion may extend into
of the intestine. The peripheral part of the mes the lamina propria of the mucosa. Warren
entery is much thinner than the root. Even in (1939) estimates that the ratio of mucosal area
obese specimens the great mesentery does not to serosal area for the whole small intestine of
contain a great amount of fat, and it is deposited the dog is 8.5 to 1. The villi account for most of
most abundantly only along the larger vessels, this large surface area, since the dog has no cir
so that large translucent areas are present be cular mucous folds.
tween the vascular branches and arcades. The submucous coat (tela submucosa) resem
bles that of the stomach and large intestine. It
loosely binds together the mucous and muscular
Coats of Small Intestine layers. The smaller blood vessels, lymphatics,
and the submucous nerve plexus are located in
The small intestine, like the other parts of the it. Trautmann and Fiebiger (1957) state that
alimentary tract, is composed of mucous, sub- the aggregated follicles are located mainly in
mucous, muscular, and serous tunics. the submucosa, with only a small portion in the
The mucous coat (tunica mucosa), through lamina propria of the mucosa.
out the small intestine of the dog, presents a free The muscular coat (tunica muscularis) con
surface which is velvety owing to the presence sists of a relatively thin outer longitudinal layer
of innumerable intestinal villi (villi intestinales). (stratum longitudinale) and a thicker inner cir
The single-layered surface cells are of two cular layer (stratum circulare). According to
types. One type consists of the columnar cells Titkemeyer and Calhoun (1955), a definite ileo
which function in absorption, the other type colic sphincter (sphincter ileocolicum) exists as
consists of the goblet, mucus-producing cells. a thickening of the inner circular coat, guarding
The deeper part of the mucosa is occupied a definite constriction of the lumen of the gut at
largely by the intestinal glands (glandulae in this level, which is called the ileocolic orifice
testinales) and diffuse lymphoid tissue and single (ostium ileocolicum). As described below, the
follicles. In about 22 areas throughout the small cecum communicates only with the large intes
intestine of the dog the lymphoid follicles are tine in the dog.
grouped together to form the aggregated fo lli The serous coat (tunica serosa) of the small
cles (noduli lymphatici aggregati). Titkemeyer intestine is composed of the peritoneum, which
and Calhoun (1955) found the aggregated folli completely covers the duodenum except along
cles to be circumscribed elevations measuring the lines where it is attached, including the
about 2 cm. by 1.5 cm. They are more numer duodenocolic ligament, and a small elongated
ous in the proximal portion of the small intestine area where it leaves the pancreas to reflect
than in the ileum, many being found in the duo around the duodenum. The jejunum and ileum
denum. In the distended bowel they are visible are also truly intraperitoneal organs. The only
through the serosa and are more numerous in parts not covered by peritoneum are along the
the side walls of the intestine than in that part lines of the mesenteric attachment and on the
L a r g e I n t e s t in e 689
antimesenteric side of the terminal portion of only slightly larger in diameter than the small
the ileum where the cecum is loosely fused to intestine. Its most important function is the
the ileum and where this attachment is con dehydration of its fecal contents. The large in
tinued by the ileocecal fold. testine is divided into cecum, colon, rectum,
and anal canal. It begins at the ileocolic sphinc
Vessels of Small Intestine ter and ends at the anus.
C RAN I AL
.Transverse colon
Opening into
duodenal fassa
Mi ddl e col i c a.
Descending
colon
-Duodenocolic
ligam e nt
- Duodenum
A Je ju nu m
CRANIAL
Inner c irc u la r
Oute r - m u s c l e of
loncji t u d i n al m — a s c e n d i n g colon
I l eo c o l i c -
s phi nc t er - Cecocolic
sphincter
Ileum—
Openings f o r - - -Cecum
0 a.,v. r n.
A s c e n d i n g colon
S o l i t a r y lymph foil i d e s -
C e c o c ol i c o r i f i c e -
Accessory ileocecal fo ld - _
Ileocolic o rific e -
v s p h i n c t e r m.
I l e o c e c a l fold - - ^ j
7- - S o l i t a r y lymph f o l l i c l e s
Recfum - - Re ct o c o c c y y e u s m,
- S p h i n c t e r ani e xt er nus
Sp h i n c t e r ani i n t e r n u s
Anorectal line
-Circumanal ylands
Op en i ng to a n a l s a c
Cutaneous zone
C o l u m n a r zone
Anocutaneous line '' Cutaneous r i d g e
of i n t e r m e d i a t e zone
F ig . 1 3 -1 9 . T h e rectum and anal canal, opened to the left of the mid-dorsal line.
692 C h apter 13. T h e D ig e s tiv e S y s te m and A bdom en
that the variations of its flexures are formed by tinal mass lies adjacent to the ascending colon
its not having a wide mesentery. As the cecum ventrally and on the left. Cranially the ascending
grows out from the colon it is apparently re colon lies in contact with the stomach unless this
strained from growing in a straight line by its viscus is greatly distended, in which case the
attachment to the ileum. The flexures develop stomach displaces the small intestinal mass cau
quite irregularly. In its simplest form the cecum dally and lies ventral to the relatively fixed as
is sigmoid in shape, but more often it is in the cending colon.
form of an irregular corkscrew with a large U- The transverse colon (colon transversum)
shaped kink extending to the left from its ileal forms an arc which runs from right to left cra
attachment. The cecum is located to the right nial to the cranial mesenteric artery and the dor
of the median plane, usually within the duo sal part of the mesojejunoileum or root of the
denal loop. It lies dorsal to and occasionally mesentery. Like the ascending colon, with
partly surrounded by the coils of the jejunum, which it is continuous at the right colic flexure,
ventral to the right transverse processes of the it may fail to form or be placed to the left of the
second to fourth lumbar vertebrae. In rare in mesenteric root owing to the failure of the gut
stances it contacts the right lateral abdominal to rotate sufficiently. It is related cranioventrally
wall. to the stomach and craniodorsally to the left
limb of the pancreas. Ventrally and caudally it
Colon lies in contact with the coils of the small intes
tine. It is about 7 cm. long.
The colon (Fig. 13-14) is divided into ascend The descending colon (colon descendens) is
ing, transverse, and descending portions and the longest segment of the colon. It extends
their connecting flexures. The colon lies in the from the left colic flexure to a transverse plane
dorsal part of the abdominal cavity and is passing through the pelvic inlet, where it is con
shaped like a shepherd’s crook or question mark. tinued by the rectum without demarcation. It
The hooked part of the colon lies cranial and is about 12 cm. long and usually quite straight.
to the right of the root of the mesentery. The It follows the curvature of the left lateral ab
cranial part of the crook is the transverse dominal wall, usually covered by the greater
colon, the short right portion is the ascending, omentum, from the dorsal part of the left costal
or right, colon. The flexure which unites these arch to a point ventral to the promontory of the
two parts is known as the right colic flexure sacrum. It lies closely applied dorsally to the
(flexura coli dextra), or hepatic flexure. The iliopsoas muscle, but at its beginning it lies in
transverse colon is continued by the descend contact with the left lateral or occasionally with
ing colon at the flexure located to the left of the the ventral surface of the left kidney. The left
root of the mesentery. This bend is called the ureter lies dorsal to its medial border initially,
left colic flexure (flexura coli sinistra), or splenic but farther caudally this tube obliquely crosses
flexure. The colon measures about 2 cm. in di the dorsal surface of the colon and curves
ameter throughout its length and is approxi around the caudal part of its lateral border be
mately 25 cm. long in a beagle-type preserved fore emptying into the bladder. Usually the as
specimen. Because of its shorter mesentery the cending portion of the duodenum lies adjacent
colon does not vary as much in position or in to its medial or mesenteric border and the
length as does the small intestine. spleen crosses it laterally; elsewhere it is
The ascending colon (colon ascendens) be bounded by the small intestinal mass. The
gins at the ileocolic sphincter, runs cranially, uterus and the bladder lie ventral to its terminal
and ends at the right-angled right colic flexure. part. The body of the uterus always lies in con
It usually is about 5 cm. long, but this varies tact with it, and the bladder, if it is distended
greatly. In rare instances the ascending colon sufficiently to extend cranial to the uterine body,
is lacking, or it may lie cranial or even to the lies in contact with it at a more cranial level. In
left of the mesenteric root. In these specimens the male an enlarged prostate gland replaces
apparently the whole gut failed to rotate around the uterus as a ventral boundary.
the cranial mesenteric artery as it usually does.
Typically, the ascending colon is related to
the mesoduodenum and the right limb of the Mesentery of Colon
pancreas dorsally, where it lies ventral to the
right kidney. The descending part of the duo The mesocolon (Fig. 13-12) is divided into
denum bounds it on the right. The small intes the same parts as the colon which it suspends.
L a r g e I n t e s t in e 693
No part of the colon of the dog is retroperito mesocolon represents the common dorsal mes
neal, nor does any part of the greater omentum entery and attaches to the aorta approximately
attach to it. As the proximal part of the colon in the median plane. Both its right and left peri
hooks around the cranial side of the root of the toneal leaves are interrupted in a frontal plane
mesentery, the mesocolon of this part attaches about 1 cm. from their aortic attachments. The
to it. The two peritoneal sheets which largely secondary fold which blends with the right peri
compose the mesocolon do not run uninter toneal sheet is the duodenocolic ligament. It ex
ruptedly at all places to their central attach tends as far caudally as the pelvic inlet before it
ments. Therefore, the various parts will be is effaced. On the left side, a loose, fat-streaked
described separately. The ascending mesocolon plica, a fold from the greater omentum and
(mesocolon ascendens) is the mesentery of the spleen, blends with the left peritoneal sheet of
ascending colon. It is shortest at its origin, the descending mesocolon. It attaches at the
where the ileum, colon, and cecum come to same dorsoventral level as does the duode
gether. In some specimens these parts are di nocolic ligament on the right side and fades
rectly attached by areolar tissue to the left completely a few centimeters cranial to the pel
mesenteric lymph node, but usually a short vic inlet. In addition to carrying the caudal
mesentery exists. Cranially, this mesentery is the mesenteric artery and the large cranially cours
beginning of the ascending mesocolon; caudally, ing caudal mesenteric vein and fat, one to sev
it is the end of the great mesentery or the meso- eral left colic lymph nodes are located around
ileum. The medial peritoneal sheet of the as the terminal branches of the caudal mesenteric
cending mesocolon attains a width of 2 to 3 cm. artery.
at the right colic flexure. Its length varies with
the length of the ascending colon. It is continu
ous centrally with the great mesentery which Rectum
covers the right mesenteric lymph node, cranial
mesenteric vessels, and nerves. A small colic The rectum (Fig. 13-19) is arbitrarily said to
fossa is usually formed by a thin, circular plica begin at the pelvic inlet (Sisson and Grossman
of peritoneum that bridges between the ascend 1953), where it is continuous cranially with the
ing and transverse colon at the right colic flex descending colon; it ends caudally, ventral to
ure. The left sheet of the ascending mesocolon the second or third coccygeal vertebra, at the
is interrupted by the mesoduodenum attaching beginning of the anal canal. It is straight, about
to it. It becomes coextensive with the right 5 cm. long, and 3 cm. in diameter. Dorsally, the
peritoneal leaf of the root of the mesentery by rectum is attached to the ventral surface of the
continuing on the large cranial mesenteric vein. sacrum by the thin, 1 cm. wide mesorectum. Cau
The transverse mesocolon (mesocolon trans- dally, the mesorectum becomes narrow and ends
versum) runs directly to the mesenteric root. It at a point usually opposite the second coccygeal
is about 3 cm. wide at the right colic flexure, but vertebra. The peritoneal sheets of the mesorec
as it crosses the median plane it increases in tum are continued on the sides of the pelvis as
width to about 5 cm. at the left colic flexure. the parietal peritoneum; caudally the visceral
The length of the transverse mesocolon, deter peritoneum from the rectum reflects forward
mined by the length of the transverse colon, is at an acute angle in a frontal plane to become
usually about 7 cm. It is continuous at the right coextensive with the parietal peritoneum which
colic flexure with the ascending mesocolon and is derived from the lateral portion of the meso
at the left colic flexure with the descending rectum. In this manner, a pararectal fossa
mesocolon. The descending mesocolon (meso (fossa pararectalis) is formed on each side, right
colon descendens) is continuous without de and left of the terminal portion of the rectum.
marcation with the mesorectum at the pelvic Ventrally, the peritoneum blends with that of
inlet. The width of the descending mesocolon the rectovesical excavation of the male or with
is greatest at the left colic flexure, where it the rectouterine excavation of the female. The
measures about 5 cm. Its width decreases cau rectum is bounded dorsally by the right and left
dally, so that it is only about 2 cm. wide as it is ventral sacrococcygeal muscles. Laterally it is
continued in the pelvis by the mesorectum. Its bounded primarily by the levator ani (medial
parietal attachment is about 12 cm. long, and coccygeal) muscle. The visceral branch of the
its visceral attachment is approximately twice internal iliac artery obliquely crosses the lateral
this length. The dorsal part of the descending side of the initial portion of the rectum. After
694 C h apter 13. T h e D ig e s tiv e S y s te m and A bdom en
the bifurcation of this vessel the urogenital gland tissue located under it. The shorter ven
artery continues the direction of the parent ves trolateral borders converge in forming a V with
sel, and the internal pudendal runs parallel to the apex pointing ventrally. A low ridge, about
the rectum for about 2 cm. before passing lat 2 mm. wide and 8 mm. long, continues from the
eral to the levator ani muscle. In a craniocaudal apex of the V toward the pudenda. The inner
sequence the rectum is crossed laterally first by portion of the cutaneous zone is about 4 mm.
the obturator, then by the ischiatic, pelvic, and wide, and in life its surface is moist. The duct
anal nerves. The pelvic plexus lies lateral to the from the anal sac opens on this zone, about 2
middle portion of the rectum. The hypogastric mm. from its cranial limit, and in the depths of
nerve enters it from in front, and the pelvic the lateral angle of the anus. The outer cutane
nerve or nerves enter it from above. Ventrally, ous zone may be defined as the relatively hair
the rectum is bounded by the vagina in the fe less zone peripheral to the anus. It varies
male, and by the urethra in the male. When the greatly in width, particularly in adult male dogs,
prostate gland is small, it lies within the pelvis owing to the varied development of the under
or at the pelvic brim, and bounds the rectum at lying circumanal glands. Because these glands
that place. When it is large, the prostate lies probably grow throughout life in the male
largely cranial to the pelvic inlet. The most (Parks 1950) and their full development tends
prominent feature of the rectal mucosa is the to result in a loss of hair, the outer cutaneous
presence of approximately 100 solitary lymph zone may attain a width of 4 cm. in large, old
nodules (noduli lymphatici solitarii). These nod male dogs. The cutaneous zone of the dog is
ules are each about 3 mm. in diameter and 1 studded by small elevations on the crests of
mm. high. The free surface of each is umbili- which open the ducts of the circumanal glands.
cated, forming a crater, or rectal pit. The intermediate zone (zona intermedia) is
usually less than 1 mm. wide, and is in the form
of an irregular, sharp-edged scalloped fold,
Anal Canal which is divided into four arcs. Known as the
anocutaneous line (linea anocutanea), it com
The anal canal (canalis analis) (Fig. 13-19) pletely encircles the anal canal. Its mucosa,
is the terminal, specialized portion of the ali like the surface of the cutaneous zone, is also
mentary canal. It is about 1 cm. long and ex stratified squamous epithelium.
tends from the termination of the rectum to the The columnar zone (zona columnaris) is so
anus. The anal canal lies ventral to the fourth named because it contains longitudinal or
coccygeal vertebra and is surrounded by both oblique ridges, or anal columns (columnae ana-
the smooth and the striated anal sphincter mus les), which run forward from the anocutaneous
cle. The mucosa of the anal canal is divided into line for about 7 mm. Caudally, the adjacent anal
cutaneous, intermediate, and columnar zones. columns are united by the fold which forms the
The cutaneous zone (zona cutanea), the most anocutaneous line. In this way, a large number
caudal of the three zones of the anal canal, is of pockets are formed, called anal sinuses
divided into external and internal portions (sinus anales). The anal sinuses are contained
(Martin 1923). The anus, the terminal opening within the four arches of the anocutaneous line,
of the alimentary canal, may be located in a thus producing its scalloped appearance. The
plane separating the two portions of the cuta smaller of the four arches are dorsal and ven
neous zone. Thus the outer cutaneous zone is tral; the larger ones are located laterally. The
not properly a zone of the anal canal, because columns are not uniform in either length or di
it lies outside of the canal. It is feasible, how rection. Some disappear after running only a
ever, to describe the two zones together as the few millimeters cranially; most end in a line
division between them varies with the move which encircles the anal canal, known as the
ment of the tail and the degree of fullness of the anorectal line (linea anorectalis).
rectum. Except during defecation, the anus is The anal sacs (sacci anales) (Fig. 13-20), one
closed. In an animal’s normal position, with the on each side of the anal canal, are approximately
tail hanging, the anus is indicated by a trans spherical sacs which are located between the
verse groove; however, if the tail is slightly inner smooth and the outer striated sphincter
raised, the boundaries of the anus form an ir muscle of the anus. The anal sacs vary in size
regular isosceles triangle. The dorsal or longer from a pea to a marble, the average diameter
border is not straight in old male dogs but is being a little less than 1 cm. The excretory duct
ventrally arched owing to the large mass of of each sac is about 5 mm. long and 2 mm. in
L a r g e I n t e s t in e 695
diameter, and opens near the cranial end of the tion being a foul-smelling, serous to pasty liquid.
furrow between the dorsal and lateral parts of According to Montagna and Parks (1948), both
the inner cutaneous zone of the anus adjacent the anal sac and its duct are lined by comified
to the intermediate zone. In about 10 per cent stratified squamous epithelium, subjacent to
of dogs the opening of the anal sac is located in which lies a thick mantle of glandular tissue em
the broad depression formed by the lateral arch bedded in a connective tissue stroma rich in dif
of the anocutaneous line on each side. The anal fuse lymphatic tissue.
sacs are of considerable clinical importance.
They frequently become enlarged, owing to ac Special Muscles of Rectum and Anal Canal
cumulated secretion, or they may become ab
scessed and painful, causing constipation. In The internal anal sphincter (m. sphincter
frequently they rupture to the outside, lateral ani internus) is the caudal thickened part of the
to the anus, producing anal fistulas. For a clini circular coat of the anal canal. It is composed
cal discussion see Grau (1935). of smooth, and therefore involuntary, muscle.
Glands of anus. Three gland areas are located Its size, particularly its width, is much less than
in relation to the anus. These comprise the cir that of the external anal sphincter. It is lined by
cumanal glands, the anal glands, and the glands submucosa on its inner surface, and is separated
of the anal sacs. from the external anal sphincter by a small
The circumanal glands (glandulae circum- amount of fascia. On its lateral external surface
anales) are located around the anus in a sub on each side lies the anal sac, which is inter
cutaneous zone which may reach a radius of 4 posed largely between the two sphincter mus
cm. and a thickness of 8 mm. Circumanal gland cles. The duct from the anal sac crosses the
elements are also found in the walls of the anal caudal border of the internal sphincter, which
sac ducts (Parks 1950), and may extend periph extends slightly farther caudally than does the
erally a short distance under the skin which con external anal sphincter.
tains abundant hair. Parks found that the The external anal sphincter (m. sphincter
circumanal gland is a bipartite structure consist ani externus) averages about 1.5 cm. in width
ing of a superficial sebaceous portion and a deep and 0.5 to 1.5 mm. in thickness. It is more than
non-sebaceous part. Since the non-sebaceous 2 cm. wide dorsally, and usually about 1 cm.
cell is capable of transforming and in certain wide ventrally. On the sides, because of the
cases does transform into a sebaceous cell, underlying anal sacs, it forms variably devel
Parks concluded that the circumanal gland is oped bulges. It is largely a circular band of
potentially a sebaceous gland. striated, and therefore voluntary, muscle, and
Scattered among the circumanal glands are is the chief guardian of the lumen of the anal
apocrine glands, and sweat glands are located canal. It lies largely subcutaneously on the
in a zone 2 to 4 mm. wide directly peripheral to sides; ventrally, its fibers decussate somewhat
the anal orifice. Probably because the circum and may spread out and end on the urethral
anal glands continue to grow throughout life muscle and the origin of the bulbocavernous
in the unaltered male, adenomas of this region muscle of the male. In the female, the com
are common in old male dogs. parable fibers blend with the constrictor vulvae.
The anal glands (glandulae anales), accord About half of the deeper fibers of this sphincter
ing to Trautmann and Fiebiger (1957), are tubu- from each side continue across to the other side,
loalveolar glands which open to the outside in ventrally; in both sexes the fibers of the super
the intermediate zone. Their secretion is fatty ficial half blend with the muscles of the external
in the dog. Bradley and Grahame (1948) state genitalia. On the sides the cranial border of the
that the microscopic anal glands are laterally lo external anal sphincter is united by fascia to
cated, cranial to the circumanal glands. Ellen the caudal borders of the levator ani muscles.
berger and Baum (1943) describe these as a Dorsally the muscle becomes wider and at
band of grape-shaped glands, 5 mm. wide. taches mainly to the coccygeal fascia opposite
The glands of the anal sac (glandulae sacci the third coccygeal vertebra. This attachment
anales) lie in the wall of the sac and open into it. is made in such a way that a cranially facing,
They are composed of large, coiled, apocrine, concave fascial arch is formed through which
sudoriparous tubules. Similar tubules lie in the pass the smooth, paired rectococcygeal mus
wall of the duct, which also contains sebaceous cles, the extremities of the arch being continued
acini. The anal sac therefore is a reservoir for by small tendons to the coccygeal fascia.
the secretion of the glands of its wall, the secre A band of smooth muscle fibers about 3 mm.
696 C h ap ter 13. T h e D ig e s tiv e S y ste m and A bdom en
- - Ilium
_ - Gl ut e us p r o f u n d u s /
- - Gluteus m ed i us m.
- U r o g e n i t a l a v v.
— S c i a t i c n.
1----- Caudal g l u t e a l arv.
Gluteal fascia
-Sacrotuberous lig.
" " P i r i f o rm is m.
Gluteus s u p e r f i c i a l i s m.
Fa t in Coccijcjeus m.
i schiorectal'
f ossa P e r i t o n e a I c a v i ty
x Levator ani m.
Br a nc he s
c a ud al r e c t a l a
Sphi ncter ani exter nus
Perineal a w . '
xAnal sac
S p h i n c t e r a n i i nt e r nu s / , C i rc u ma na l gl ands
Openincj of duc t of anal s a c1 1Duct of a n a l sac
C o l u m n a r zone of anus Cutaneous zone of anus
F i g . 13-20. Section through anus in horizontal plane, dorsal aspect. The right side is cut at a lower level, through the duct of
the anal sac.
L a r g e I n t e s t in e 697
wide and less than 1 mm. thick arises on each blood to the anal canal. Thus the blood from the
side from the ventral surface of the sacrum or anal canal returns to the heart both by the por
first coccygeal vertebra and sweeps caudoven- tal system, through the cranial rectal, caudal
trally. As it obliquely crosses the rectum it con mesenteric, portal, and hepatic veins and post
tributes some fibers to it and then fans out be cava, and by the systemic system, through the
tween the anal sac and the internal anal caudal rectal and perineal veins, which are trib
sphincter. This band of muscle fibers constitutes utaries of the internal pudendal or the perineal
the coccygeoanal muscle (see Fig. 3-41). The and caudal gluteal veins. The internal pudendal
major portion of this muscle appears to end near and caudal gluteal veins unite to form the in
the duct of the anal sac, with some fibers insert ternal iliac vein. The internal and external iliac
ing in the external anal sphincter. The minor veins unite to form the common iliacs, which in
ventrolateral portion of the coccygeoanal mus turn unite to form the postcava. The venous
cle, in combination with fibers from the external blood is returned from the anal canal through
anal sphincter, continues distally as the mixed the veins which are the satellites of the arteries
(smooth and striated) retractor penis muscle of of supply. The skin of the perineum and the
the male. underlying circumanal glands are served largely
The rectococcygeal muscle (m. rectococcyg by the perineal vessels, usually from both the
eus) is a paired, smooth muscle at its origin, internal pudendal and caudal gluteal arteries.
composed of a condensation of many of the The lymph vessels from the anal canal drain
outer longitudinal fibers from each side of the into the sacral lymph nodes, if present, and the
rectum. These fibers sweep caudodorsally internal iliac lymph nodes. The external anal
from the sides of the rectum and pass dorsally sphincter muscle, being striated and voluntary,
through the fascial arch formed by the attach is supplied by the anal branch of the pudendal
ment of the external anal sphincter to the fascia nerve. The involuntary, smooth internal anal
of the tail. Right and left portions lie closely to sphincter and rectococcygeus are supplied by
gether at this site, ventral to the third coccygeal autonomic fibers from the pelvic plexuses. The
vertebrae, and fuse. Posteriorly the muscle lies parasympathetic portion comes to it through
on the bodies of successive coccygeal vertebrae, the pelvic nerves, branches from usually the
in the groove formed by the apposed ventral first and second sacral nerves, and the sympa
sacrococcygeal muscles, and runs caudally to thetic portion is derived from the hypogastric
attach on the bodies of the fifth and sixth coc nerves which arise from the caudal mesenteric
cygeal vertebrae. The attachment of the recto ganglion.
coccygeus on the tail serves as an anchorage
whereby the muscle can stabilize the anal canal
and rectum and prevent their being pulled Coats of Large Intestine
cranially by a peristaltic wave, or, by its con
traction, it can move the anal canal and rectum All except the terminal part of the large intes
caudally during defecation. The movement of tine has the usual four coats as found in the
the tail during the act of defecation has a direct small intestine. These are the mucous, submu
influence in evacuating the rectum not only cous, muscular, and serous tunics or coats.
through the action of the rectococcygeus but The mucous coat (tunica mucosa) of the large
also by the action of the coccygeus and levator intestine differs from that of the small intestine
ani muscles. The coccygeal muscles cross in that there are no aggregated lymph nodules
the rectum laterally and tend to compress the or intestinal villi. Solitary lymph nodules, how
tube while the rectococcygeus, by shortening, ever, are numerous, and can be counted from
aids the circular muscle coat in moving the fecal the outside in the dilated gut. Although present
column to the outside. throughout the large intestine they are most
numerous in the rectum. The mucosa of the
Vessels and Nerves of Anal Canal large intestine, except in the anal canal, con
tains no folds which cannot be effaced by dis
The mucosa of the anal canal and the sphinc tention. Many folds which appear similar to
ter muscles which surround it receive their those of the contracted stomach are present in
blood mainly from the right and left caudal rec the contracted large intestine. Depending on
tal arteries which anastomose in the anal the type of contraction, these plicae are either
sphincters. The long cranial rectal artery may longitudinal or circular. The intestinal glands
extend far enough caudally to furnish some (glandulae intestinales) of the large intestine
698 C h apter 1 3 . T he D ig e s t iv e System and A bd o m en
are longer, straighter, and richer in goblet cells The bones, muscles, blood vessels, nerves, and
than are those of the small intestine (Trautmann glands of the pelvic outlet are described in the
and Fiebiger 1957). They are lined by a colum appropriate chapters, so that at this place only
nar epithelium which is continuous with the the ischiorectal fossa, and the perineal and, to
columnar epithelium of the mucosal surface of a lesser extent, the pelvic fasciae will be de
the lumen of the gut. The lamina muscularis scribed. The perineal region holds much inter
m ucosae consists of muscle fibers which are est for the surgeon because of the frequency of
poorly arranged in two strata. occurrence of adenomas and perineal hernias
The submucous coat (tela submucosa) does in old male dogs.
not differ appreciably from that of the small The ischiorectal fo ssa (fossa ischiorectalis) is
intestine. Many of the solitary lymph nodules the deep, wedge-shaped depression located lat
are located partly within it. This tunic contains eral to the terminal pelvic portions of the diges
the submucous nerve plexuses and many vessels tive and urogenital tubes. The lateral boundary
in the meshes of loose connective tissue. of the fossa is the levator ani and coccygeus
The muscular coat (tunica muscularis) is muscles which obliquely cross the lateral sur
uniform in thickness. The stratum longitudinale face of these tubes as the muscles pass essen
is not concentrated in muscular bands as it is in tially sagittally from the os coxae to the coccyg
man, horse, and pig and no haustra are present. eal vertebrae. The most caudal part of the
The fibers forming the longitudinal stratum medial boundary is the external anal sphincter
sweep dorsocaudally from the sides of the rec dorsally, and the constrictor vulvae of the fe
tum and, opposite the first or second coccygeal male and the retractor penis of the male ven
vertebra, leave the dorsum of the rectum to trally. The lower lateral and ventral boundary
form the smooth m. rectococcygeus, which is formed by the internal obturator muscle, the
passes dorsal to the external anal sphincter and dorsal and upper lateral boundary by the super
attaches to the bodies of the fifth and sixth coc ficial gluteal muscle. Cranially and ventrally, the
cygeal vertebrae. The stratum circulare of the fossa forms a narrow fornix where the medially
large intestine resembles that of the small intes located coccygeal muscles arise adjacent to the
tine, except that it is heavier. Its caudal portion origin of the laterally located internal obturator
forms the internal anal sphincter muscle. muscle. This angle is located opposite the bodies
The serous coat (tunica serosa) resembles of the ilium and ischium and along the entire
that of the small intestine. It covers the colon, pelvic symphysis. The ischiorectal fossa in well-
cecum, and much of the rectum, as the visceral nourished dogs is filled with fat which is
peritoneum. The anal canal and the caudal por bounded peripherally by the skin.
tion of the rectum are retroperitoneal. See the The perineal fa sc ia (fascia perinei) is a term
description of the pararectal and rectovesical or used by veterinary anatomists to include those
rectouterine excavations, in the discussion of parts of the adjacent fasciae from the tail, rump,
the peritoneum, above. and thigh which converge at the anus, enclos
ing the pelvic outlet. It is divided into superficial
and deep strata.
The superficial perineal fascia (fascia perinei
superficialis) forms the feeble matrix in which
PERINEUM the fat of the ischiorectal fossa is elaborated. It
is a single-layered, loose fascia, but it is abun
The perineum is the region of the pelvic out dantly developed. The numerous small perineal
let. On the surface of the body of the dog it is vessels and nerves which stream caudally over
limited by the tail above, the scrotum or begin the ventral part of the pelvic outlet lie in the
ning of the vulva below, and by the skin which superficial perineal fascia. It is continuous with
covers the paired superficial gluteal and inter the superficial fascia of the rump, thigh, and
nal obturator muscles and the tubera ischiorum tail.
on the sides. Deeply, the perineum is bounded The deep perineal fa scia (fascia perinei pro
by the third coccygeal vertebra above, the funda) covers the dorsomedial surface of the in
sacrotuberous ligaments on the sides, and by ternal obturator muscle. It firmly attaches to the
the arch of the ischium below. The digestive dorsal subcutaneous portion of the tuber ischii
system terminates in the anus. The urogenital ventrally, and the adjacent portion of the
system in both sexes continues distal to the obliquely running sacrotuberous ligament cau
perineal region to occupy the pudendal region. dolaterally. Craniolaterally it becomes continu
L iv e r 699
ous with the deep gluteal fascia of the superficial faces bilaterally and dorsally than faces cranially
gluteal muscle. and ventrally. Therefore, this surface can be
In the male the deep perineal fascia is heavy logically subdivided into a right part (pars dex
and tightly applied to the dorsal surfaces of the tra), a left part (pars sinistra), a cranial part (pars
right and left ischiocavernosus muscles and to cranialis), a dorsal part (pars dorsalis), and a. ven
the unpaired bulbocavernosus muscle which tral part (pars ventralis). The right part is larg
lies between them. In both sexes the deep est, as it extends from the cranial part, which
perineal fascia, but no muscle fibers, extends lies opposite the sixth intercostal space to the
cranially between the digestive and urogenital last, or twelfth, intercostal space; on the left
tubes at the pelvic outlet. Developed in the cau side the caudal border usually lies opposite the
dal part of this frontally located fascial partition tenth intercostal space. The cranial part is
is the retractor penis muscle of the male and the marked by a shallow, broad, indistinct cardiac
constrictor vulvae muscle of the female. Just impression (impressio cardiaca) which is located
cranial to the muscles which pass from the ex largely to the left of the median plane. The dor
ternal anal sphincter to the pudenda in both sal part is deeply notched, approximately in the
sexes, collagenous fibers directly unite the com median plane, by the postcava and the esoph
plex musculature between the anal canal and agus. The postcava lies to the right of the esoph
the vagina or the bulb of the penis. This median agus as they groove the liver.
fascial union, if sufficiently heavy or differenti The visceral surface (facies visceralis) of the
ated as a fibromuscular node, is spoken of as the liver is irregularly concave and faces mainly
“perineal body” (centrum tendineum perinei). caudoventrally and to the left. It lies in contact
with the stomach, duodenum, pancreas, and
LIVER right kidney. All but the pancreas produce im
pressions on the organ hardened in situ. The
The liver (hepar) (Figs. 13-21, 13-22) is the liver is completely invested with peritoneum
largest gland in the body. It is both exocrine and except at the hilus and where the gall bladder
endocrine in function. The bile, which is its is fused to it. The centrally located papillary
exocrine product, is largely stored in the gall process (processus papillaris), which protrudes
bladder before being poured into the descend caudally from its dorsal part, is the most prom
ing portion of the duodenum. Its endocrine inent feature of the visceral surface.
substances released into the blood stream func The gastric impression (impressio gastrica)
tion in the intermediary metabolism of fats, occupies the whole left half of the visceral sur
sugars, and some nitrogenous products. face when the stomach is moderately full. The
pyloric part, owing to its relatively fixed posi
Physical Characteristics tion, produces an oblique impression across the
middle portion of the gland, where it lies in con
The average weight of the liver in 91 dogs
tact with the caudal face of the gall bladder.
was 450 gm. The average weight of these dogs When the stomach is empty, the coils of small
was 13.3 kg. In this sampling of adult mongrel intestine contact the visceral surface of the liver
dogs of both sexes the weight of the liver aver through the greater omentum but leave no im
aged 3.38 per cent of the body weight. Thus,
pressions on it.
the liver weighed approximately a pound in
The duodenal impression (impressio duo-
those dogs whose average weight was about 27
denalis) begins at the junction of the right and
pounds. The liver is relatively much heavier in
quadrate lobes as the most cranial indentation
the puppy than in the aged dog. The fresh liver
of the visceral surface. This impression at first
is a deep red color, firm in consistency, but fri
runs to the right, then it makes a sweeping arch
able. In a 30-pound, hound type dog, its dorso-
caudoventrally and finally runs caudodorsally,
ventral dimension is 14 cm., its width 12 cm.,
essentially paralleling and lying dorsal to the
and its thickness 6 cm.
right ventral border of the organ. The right
limb of the pancreas lies dorsomedial to the
Surfaces, Borders, and Relations
cranial and descending portions of the duo
The diaphragmatic surface (facies dia- denum in contact with the liver, but leaves no
phragmatica), or parietal surface, of the liver impression on it.
is strongly convex in all directions as it lies The renal impression (impressio renalis) is a
mainly in contact with the diaphragm. It is so deep, nearly hemispherical fossa formed by the
strongly convex that more of this surface cranial pole of the right kidney projecting into
700 C h ap ter 13. T h e D ig e s tiv e S y s te m and A bdom en
Q u a d r a t e l obe
Gal Ibladd er
' H e p a t i c v.
F alciform
L. t r i a n g u l a r I ig.
/ I 1 /
H e p a t i c vv.' /
1 '/ / /
/ /
L e s s e r omerfum' ^
Caudat e labe ‘ I I I
I — Caudat e p r o c e s s of
Postcava1 {1 j1 R. t r i a n g u l a r l i g . c a u d a t e l obe
C o r o n a r y lig. ' Bare a r e a o f l i v e r
xLesser omenium
xLeft t r i a n g u l a r lig.
\ \ ^ \ NP r o p e r h e p a t i c aa.
C ommon h e p a t i c a.
. , v P o r t a l v.
' i > '
Hepatorenal lig. | J | c ommon b i l e duc t
Rt l a t lobe ; 'Rt g a s t r i c a
Postcava ' ' Ga s t r o d u o d e n a l a.
the most caudodorsal portion of the liver. Ven becomes wider, until it reaches a width of 4.5 to
trally, the liver covers more than half of the 5 cm. in its middle, after which it gradually be
kidney. Because of the close proximity of the comes narrower and ends in a point dorsal to the
postcava to the right adrenal gland, this gland last sternebra. The lateral border may protrude
does not leave an impression on the liver. as much as 2 cm. caudal to the ventral portion of
The porta of the liver (porta hepatis) is the the costal arch, but in some specimens it is com
hilus of the organ. The hepatic vessels and pletely contained within the rib wall. The dorsal
nerves and the bile duct communicate with the portion partially caps the body of the stomach.
gland through the porta. The nerves and arter The visceral surface of the left lateral lobe is con
ies enter the porta dorsally, the biliary duct cave peripherally as it lies on the fundus and
leaves ventrally, and the portal vein enters be body of the stomach. Centrally, it is partly cov
tween the two. It is located on the dorsal third ered by the papillary process of the caudate lobe.
of the visceral surface, ventrodextral to the at This central portion of the lobe is slightly con
tachment of the papillary process. From the vex, forming the omental tuber (tuber omen-
porta, the deep fissures which subdivide the tale). It lies adjacent to the lesser omentum
organ diverge toward the lateral and ventral covering the papillary process and is formed by
surfaces, so that the liver, opposite and dorsal the moldable hepatic tissue protruding toward
to the porta, is solid. the lesser curvature of the stomach.
Ventral, right and left lateral borders (margo The left m edial hepatic lobe (lobus hepatis
ventralis, lateralis dexter et lateralis sinister) are sinister medialis) varies from being nearly tri
recognized. They are sharp-edged and continu angular to oval in outline as seen from the dia
ous around the periphery of the organ except phragmatic surface. The fissure which separates
dorsally where this circumferential margin is it from the left lateral lobe begins from 1.5 to 3
effaced by the deep, broad notch which con cm. from the most caudoventral portion of the
tains in its depths the postcava and esophagus. organ. It exists as a deep, curved cleft, which
In addition to the main clefts which subdivide usually completely separates the two portions
the liver into lobes, there are a few short fissures of the left lobe of the liver. It extends to the
which cut into the borders of the organ. In the porta. Dorsally, in some specimens, the two por
dog, the fissure fo r the round ligament (fissura tions are joined by a narrow but deep bridge of
lig. teretis) is the caudoventral portion of the liver tissue; otherwise the two portions are
interlobar fissure between the quadrate lobe on joined together only by the intrahepatic vessels
the right and the left lateral lobe on the left. In and nerves. The two portions of the left lobe
the puppy, the round ligament (umbilical vein are separated from the quadrate and right lobes
of the fetus) runs from the umbilicus to by a deep fissure, nearly mid-sagittal in location,
the porta of the liver in or just caudal to this which extends to the porta and nearly to the
cleft. esophageal notch.
The quadrate lobe (lobus quadratus) is a deep
Lobes and Processes wedge of liver tissue which lies essentially in
the median plane, where it is interposed in the
The liver is divided into four lobes and four fissure which separates the right medial and
sublobes, as well as two processes, by deeply the left lobes, being fused to a certain extent to
running fissures. the former. Its diaphragmatic surface is fusi
The left hepatic lobe (lobus hepatis sinister) form and it extends neither to the ventral bor
is that portion of the liver which lies entirely, or der nor to the notch for the esophagus and
almost entirely, to the left of the median plane. postcava. The middle of its right surface is
This lobe forms from a third to nearly a half of smoothly excavated by the left half of the fossa
the total liver mass. Its parenchyma is usually for the gall bladder (fossa vesicae felleae). In
completely divided into two sublobes, as fol nearly half of the specimens examined the quad
lows: The left lateral hepatic lobe (lobus hepatis rate lobe did not reach the visceral surface of the
sinister lateralis) begins dorsally under the left liver.
crus of the diaphragm, where it is about 3 cm. The right hepatic lobe (lobus hepatis dexter)
wide. Traced ventrally, it crosses under the left is smaller than the left hepatic lobe and lies
portion of the tendinous center and then under completely to the right of the median plane. It
the left portion of the muscular periphery of the lies between transverse planes passed through
diaphragm. Its diaphragmatic surface gradually the upper portions of the sixth and the tenth
L iv e r 703
intercostal spaces. Like the left hepatic lobe it its diaphragmatic surface forms nearly an
is divided into medial and lateral sublobes. equilateral triangle as it lies mainly ventral to
The right m edial hepatic lobe (lobus hepatis the right kidney. The parenchyma of the cau
dexter medialis) is fused to the medially lying date lobe is usually partly fused to the right
quadrate lobe. The degree of fusion varies; in lateral lobe, but occasionally the two portions
some specimens only the dorsal portions of these are completely separated, or other variations
always closely adjacent lobes are fused; in others may exist.
the fusion extends nearly to the fossa for the gall
bladder, leaving only a short fissure extending Peritoneal Attachments and Fixation
dorsally from the fossa to separate the two
lobes. The right medial lobe is always longer The liver is almost completely enveloped by
than the right lateral lobe and is the portion peritoneum, which forms its serous coat (tunica
which extends caudally beyond the ventral por serosa). The serous coat is fused to the under
tion of the costal arch if any portion of the right lying fibrous capsule (capsula fibrosa perivascu
lobe protrudes beyond it. Its diaphragmatic laris), a thin but strong layer, composed mainly
surface is in the form of a curved triangle, with of collagenous tissue, which closely invests the
its base dorsomedial and its apex ventromedial surfaces of the liver and sends interlobular
in location. It is also triangular in cross section trabeculae into the gland substance. At the
as it is wedge-shaped, possessing a concave, porta the fibrous coat becomes heavier and is
slightly fissured medial border which extends to continued into the interior of the liver in as
the visceral surface of the organ. The right half sociation with the vascular and nervous struc
of the fossa fo r the gall bladder is located on its tures which serve the gland. The only parietal
medial face opposite the comparable excavation attachment of the liver is to the diaphragm by
on the quadrate lobe. means of continuations of its serous and fibrous
The right lateral hepatic lobe (lobus hepatis coats in the form of the coronary ligament and
dexter lateralis) is shaped roughly like a laterally several small folds which radiate from it. These
compressed hemisphere with a slightly concave folds are the two right triangular, the usually
base. Cranially it is overlapped by the right single left triangular, and the falciform liga
medial lobe; caudally it overlaps the caudate ments. The hepatorenal ligament and the lesser
process of the caudate lobe and is usually fused omentum also attach to the liver.
to it, lateral to the postcava. Its most ventral The coronary ligament o f the liver (lig. coro-
extension lies opposite the distal portion of the narium hepatis) is not a true peritoneal liga
middle third of the caudal border of the right ment since the two sheets of peritoneum which
medial lobe. form it are not in the form of a fold but are ir
The caudate lobe (lobus caudatus) is com regularly separated. The term refers to the line
posed of the caudate and papillary processes of peritoneum which reflects around a tri
and the isthmus of liver tissue which connects angular “bare area” of the liver, about 2 cm.
them. This isthmus is compressed between the long on each side, and is continued on the dor
postcava dorsally and the portal vein ventrally. sal surface of the postcava and the tributaries
It is a bridge of hepatic tissue which is about 1.5 which enter it from the diaphragm. The coro
cm. long, 1 cm. wide, and 0.5 cm. thick. nary ligament is irregular in outline. It reflects
The papillary process (processus papillaris) is the close embryonic relationship between the
pyramidal to tongue-shaped, and is usually diaphragm and liver. Its stellate border gives
partly subdivided by one or two fissures. The rise to the three or more triangular ligaments
more constant fissure separates the frenular and is coextensive with the dorsal part of the
part from the body of the process. This process falciform ligament.
is loosely enveloped by the lesser omentum and The right triangular ligament (lig. triangulare
lies in the lesser curvature of the stomach. It dextrum) is a plica of peritoneum which extends
projects by an acute angle to the left and for between the diaphragm and the dorsal part of
ward from its attachment to the caudate lobe. the right lateral lobe. Its free lateral border is
The caudate process (processus caudatus) 1 to 5 cm. wide. As it passes medially it becomes
forms the most caudal portion of the liver as it progressively narrower until its two formative
extends to a plane through the twelfth inter peritoneal layers become continuous with the
costal space or last rib on the right side. Its right peritoneal leaf of the coronary ligament as
caudolateral portion is deeply recessed by the this bounds the bare area of the liver on the
cranial half of the right kidney. The outline of right. It is usually longer than it is wide. A sec
704 C h ap ter 13. T h e D ig e s tiv e S y s te m and A bdom en
ond, smaller right triangular ligament regularly equally thin fibrous capsule which sends septa
goes from the diaphragm to the diaphragmatic into the gland. On close observation the surface
surface of the right medial lobe. Other smaller of the liver presents a fine mottled appearance.
but similar plicae are present. The delicate dappling is due to the contrast in
The left triangular ligament (lig. triangulare color between the dark, small, polygonal units
sinistrum), like the comparable right ligament, of liver parenchyma and the lighter connective
may also be double or triple. If there are two, tissue surrounding them. These units, called
the caudal member is larger than the cranial and hepatic lobules (lobuli hepatis), are the smallest
contains the fibrous appendix o f the liver (ap grossly visible functional divisions of the organ.
pendix fibrosa hepatis), when this is present. Each lobule is about 1 mm. in diameter and is
This fibrous appendix is a narrow, thin tapering composed of curved sheets of cells which en
band of atrophic hepatic tissue located in or close numerous, blood-filled cavities known as
near the free border of the ligament. It is the liver sinusoids. According to Elias (1949),
present in only a small number of adult speci the sinusoids of the dog are intermediate in
mens. When a second triangular ligament is form between the saccular and tubular types
present on the left side, it runs from the left which are found in some other mammals. The
medial lobe to the diaphragm. sheets or plates of cells which form their walls
The falciform ligament o f the liver (lig. falci- are one cell thick and contain openings which
forme hepatis) is a remnant of the ventral allow free passage of the intersinusoidal blood.
mesentery which extends between the liver and
the diaphragm and ventral body wall caudally
Blood and Lymph Vessels
to the umbilicus. The middle portion of the
falciform ligament in the dog usually becomes In the centers of the lobules there are typi
wholly or partly obliterated before birth so that cally the single central veins (venae centrales).
the umbilical vein, which in early fetal life is These constitute the beginning of the efferent
located in its free border, usually has no peri or outgoing venous system of the liver. Adjacent
toneal attachment immediately before birth. central veins fuse to form the interlobular veins
The proximal end of the falciform ligament— (venae interlobulares). The interlobular veins
that part extending from the umbilicus to the unite with each other to form finally the hepatic
diaphragm—remains as a fat-filled irregular veins (vv. hepaticae), which empty into the
fold which may weigh several pounds in obese postcava. Arey (1941) and others have shown
specimens. The distal portion of the falciform that in the dog, but not in the cat, there are
ligament may disappear completely, but usually spiral and circular muscle fibers in the walls of
it remains as a thin, avascular fold which extends the central and (sublobular) interlobular veins.
from the dorsal end of the fissure between the The sphincter action produced by these muscle
right and left lobes to the coronary ligament. fibers restricts venous drainage, producing pre
When present, the left peritoneal sheet of the cisely the effects of experimental shock. Prinz
falciform ligament becomes coextensive with metal et al. (1948) have shown that glass beads
the left portion of the coronary ligament, and measuring 50 to 180 n did not pass from an af
the right peritoneal sheet becomes coextensive ferent vessel to the efferent hepatic vessel drain
with the ventral portion. ing the area supplied by the injected afferent
The hepatorenal ligament (lig. hepatorenale) vessel. The hepatic veins convey to the post
is a delicate peritoneal fold which extends from cava all the blood which the liver receives
the medial portion of the renal fossa to the ven through the portal vein and proper hepatic
tral surface of the right kidney lateral to the fat arteries.
which fills its hilus. It is not constant. The lesser The portal vein (v. portae) brings the func
omentum (omentum minus) is a thin, lacy, fat- tional blood to the liver from the stomach, intes
streaked, loose peritoneal fold which is that tines, pancreas, and spleen. About four-fifths of
remnant of the ventral mesentery which extends the blood entering the liver reaches it by the
from the liver to the lesser curvature of the portal vein (Markowitz et al. 1949). The proper
stomach and cranial part of the duodenum. hepatic arteries (aa. hepaticae propriae) furnish
the liver with the blood which nourishes its cells
Structure —the nutritional supply. The parenchymal cells
are bathed by mixed blood from the portal vein
The free surface of the liver is firmly covered and proper hepatic arteries so that they receive
by the thin peritoneum superficially and the nutrition from both. The proper hepatic arteries
L iv e r 705
supply primarily the liver framework, including liver through the splanchnic nerves, celiac
its capsules and the walls of the blood vessels, ganglia, and celiac plexus, and continue on the
the intrahepatic biliary duct system, and the common and proper hepatic arteries as the
nerves. plexuses of these arteries. Alexander (1940)
Although only about one-fifth of the blood states that in some specimens the biliary system
coming to the liver reaches it through the receives afferent fibers from the phrenic nerves.
proper hepatic arteries, their occlusion usually He also confirmed that the hepatic artery re
results in death if such occlusion is not accom ceives only sympathetic fibers.
panied by massive doses of penicillin. Marko
witz et al. (1949) found that, without antibiotic Bile Passages and Gall Bladder
treatment following ligation of the arteries at
the portal fissure, gangrene results. In research The bile, produced by the sheets of liver cells
on the arterial blood supply to the liver it surrounded by the blood sinuses, is discharged
should not be overlooked that there are never into the minute bile canaliculi, or bile capil
less than two and in some specimens there are laries, which lie between these cells. The
as many as five proper hepatic arteries which canaliculi unite to form the plexiform interlobu
leave the common hepatic artery, and most of lar ducts (ductuli interlobulares), which lie in
these arteries branch before they enter the lobes the interstitial tissue between the lobules.
of the liver. See the treatise of Payer et al. (1956) Finally the interlobular ducts of various sizes
on the surgical anatomy of the arteries to the unite to form the lobar or bile ducts (ductuli bilif-
liver of the dog. eri), which are variable in number and termina
In the fetal pup there is a shunt from the tion. The extrahepatic bile passages consist of
umbilical vein to the hepatic venous system, the hepatic ducts (ductuli hepaticae) from the
known as the ductus venosus. The ductus liver, the cystic duct (ductus cysticus) to the gall
venosus becomes fibrotic after birth and is bladder, and the common bile duct (ductus
known as the ligamentum venosum. In the still hepaticus communis) to the duodenum. The
born pup it is several millimeters long and about right and left hepatic ducts may be single or
2 mm. wide. It extends obliquely from left to double. One of the many possible patterns of
right in the porta hepatis, where it lies ventral hepatic duct termination is illustrated by Figure
to the attachment of the papillary process. The 13-23.
lymph vessels from the liver freely anastomose The gall bladder (vesica fellea) (Fig. 13-23)
with those of the gall bladder (McCarrell et al. stores and concentrates the bile; the function of
1941). They drain into the hepatic and splenic its mucoid secretion, like that of mucus gen
lymph nodes. erally, is for lubrication and protection (Ivy
1934). Although the vagal nerve fibers are motor
Nerves to its musculature, Winkelstein and Aschner
(1924) state that the gall bladder displays vari
The liver is supplied by both afferent and ef ations in tonicity but seems to possess little con
ferent fibers through the vagi and by sympa tractile power. Intra-abdominal pressure chiefly
thetic fibers from the celiac plexus. The vagal due to the inspiratory phase of respiration ef
fibers reach the abdomen by passing through the fects a large variation in pressure within the gall
diaphragm with the esophagus as the dorsal and bladder. The gall bladder is a pear-shaped
ventral vagal (esophageal) nerve trunks. Chiu vesicle which lies between the quadrate lobe
(1943) has shown that in a representative dog medially and the right medial lobe laterally.
two branches leave the ventral vagal trunk and When distended, it extends through the thick
one leaves the dorsal at the level of the cardia. ness of the liver to its diaphragmatic surface and
They pass obliquely to the right in the lesser contacts the diaphragm. Its capacity in a beagle
omentum toward the porta and supply the liver sized dog is 15 ml. (Mann, Brimhall, and Foster
parenchyma and biliary system. McCrea (1924) 1920). It is about 5 cm. long and 1.5 cm. in its
describes the abdominal distribution of the vagi greatest width in such specimens. The blind,
in the rabbit, cat, and dog. Possibly the liver rounded, cranial end of the gall bladder is
also receives vagal fibers through the portion of known as the fundus (fundus vesicae felleae),
the dorsal vagal trunk which joins the celiac the large middle portion, as the body (corpus
plexus. Chiu (1943) also mentions the pos vesicae felleae), and its slender, tapering,
sibility of a coronary nerve reaching the liver caudodorsally directed extremity, as the neck
in the dog. The sympathetic fibers reach the (collum vesicae felleae).
706 Chapter 13. The D ig e s t iv e System an d A bd o m en
The cystic duct (ductus cysticus), in a topo muscle. As such, the musculus proprius forms a
graphical sense, may be regarded as the begin variable ring of muscle which surrounds the
ning of the biliary duct system. It extends from terminations of the bile and minor pancreatic
the neck of the gall bladder to the site of its ducts to form the m. sphincter ampullae hepato-
junction with the first tributary from the liver. pancreaticae. Ensheathing the remaining intra
From this level distally to the duodenum the mural portion of the bile duct the musculus
main excretory channel which receives bile proprius constitutes the m. sphincter ductus
from the hepatic ducts is known as the common choledochi. Eichhorn and Boyden point out that
bile duct (ductus choledochus). Higgins (1926) in the dog the downgrowth of a septum of the
has recorded double cystic and common bile tunica muscularis on the mucosal side of the
ducts in the dog. In the dog, the lobar ducts do duct creates a muscular funnel through which
not unite to form the hepatic duct as they do in the bile duct must pass. This feature makes the
man, but enter the main trunk of the excretory discharge of bile dependent to a large degree
tree which, as stated above, may be regarded as upon the activity of the duodenum.
beginning with the cystic duct. The distal or Apparently great variation exists in the
caudal portion of the common bile duct enters amount of musculature which is present at the
the dorsal or mesenteric wall of the duodenum. termination of the common bile duct. Halpert
This portion of the common bile duct may be (1932) found proper muscle present in only 1
known as the free portion, in contrast to the of 25 dogs he examined. Casas (1958) states
intramural portion, which extends obliquely that in the dog the sphincter of Oddi consists of
through the duodenal wall. The free portion is three layers of muscle.
about 5 cm. long and 2.5 mm. in diameter as it
courses through the lesser omentum. The intra PANCREAS
mural portion of the duct and its mode of
emptying into the lumen of the duodenum The pancreas (Fig. 13-14) is yellowish gray
have been studied by many investigators. when preserved and pinkish gray in life. It is a
Eichhorn and Boyden (1955) have analyzed rather coarsely lobulated, elongate gland. The
the structure of the choledochoduodenal junc lobules, as Revell (1902) has pointed out, pro
tion in the dog. They illustrate and describe duce a nodular surface with irregularly crenated
from wax reconstructions and maceration speci margins. The pancreas is located in the dorsal
mens the intramural portion of the bile duct and part of both the epigastric and the mesogastric
its musculature in both the fetus and adult. In abdominal segment, caudal to the liver. Like
their review of the literature they call attention the liver, the pancreas has both an exocrine and
to the early work by Oddi (1887), which in an endocrine function. Its exocrine secretion,
cluded studies on the dog. The bile duct has an the pancreatic juice, the most important of the
intramural length of 1.5 to 2 cm. It terminates digestive secretions, is conveyed to the descend
on a small hillock located at the end of a low ing portion of the duodenum by one or several
longitudinal ridge representing its intramural ducts, usually two. It is a clear alkaline secretion
course. The bile duct opens in the center of a containing three principal enzymes, one of
small rosette upon the hillock, and to one side which reduces proteins, one fats, and the third
is the slitlike opening of the minor (ventral) pan carbohydrates. Insulin, a protein hormone, is
creatic duct. The site of this combined opening the endocrine secretion produced by the islet
of the bile duct and the minor pancreatic duct cells. This hormone keeps the sugar content of
is spoken of as the major duodenal papilla. the blood at a constant level, and in its absence
About 3 cm. distal to this opening lies a second a fatal sugar diabetes sets in.
low hillock, upon which the major pancreatic The weight of the pancreas averaged 31.3
duct from the dorsal pancreas opens. gm. in 76 dogs with an average weight of 13.8
Eichhorn and Boyden have verified the exist kg., the pancreas thus accounting for an aver
ence of a double layer of smooth muscle around age of 0.227 per cent of total body weight in
the intramural portion of the bile duct. The this group of adult mongrel dogs of both sexes.
outer layer is formed by the tunica muscularis The pancreas therefore weighs about 1 ounce
of the duodenum. The inner layer, or musculus in a dog whose weight is approximately 30
proprius, begins in the infundibular portion of pounds. These data compare favorably with
the bile duct and extends in the submucosa to Mintzlaff’s (1909) findings in 30 dogs, although
the termination of the duct as the sole investing his specimens were on an average larger. The
P a n crea s 707
708 Chapter 13. T he D ig e s t iv e S ystem and Abdom en
average total length of the pancreas in a 30- the pyloric region, which forms a large concave
pound dog is approximately 25 cm., or 10 impression on the cranial portion of the pan
inches. creatic angle. Caudal to this impression the
The pancreas, when hardened in situ, is in pancreas is about 1 cm. thick and 3 cm. wide.
the form of a V which lies in a frontal plane with The portal vein crosses the dorsal portion of
the apex pointing forward. The gland is basi the pancreatic angle. As the gastropancreatic
cally divided into a thin, slender right lobe, and artery and gastroduodenal vein disappear into
a shorter, thicker and wider left lobe. The two the pancreas at this place, they are crossed on
lobes are united at the pancreatic angle, which their right side by the bile duct, which lies
lies caudomedial to the pylorus. The two lobes adjacent to the duodenum.
of the pancreas are also commonly known as the The left lobe of the pancreas (lobus pancre
limbs of the gland. atis sinister, or cauda pancreatis [N.A.]) lies in
the dorsal sheet of the greater omentum. It be
Lobes and Relations gins at the pancreatic angle and runs caudo-
sinistrally. It is about two thirds as long and
The right lobe of the pancreas (lobus pan- half again as wide as the right lobe, measuring
creatis dexter, or caput pancreatis [N.A.]) lies 10 cm., or 4 inches, in length, and 4 cm., or 1.6
in the mesoduodenum near or in contact with inches, in width. Its dorsal surface (facies dor
the dorsal portion of the right flank. It extends salis), on the right, is related to the caudate
from a transverse plane through the middle of process of the liver and then, in succession on
the ninth intercostal spaces to one through the the left, to the portal vein, postcava, and aorta.
fourth lumbar vertebra. The right lobe varies It ends in the left part of the sublumbar region
in width from 1 to 3 cm. and in thickness up to in close relation to the cranial pole of the left
1 cm. Its length is approximately 15 cm., or 6 kidney and the middle portion of the spleen. A
inches, in a beagle-type dog. The right lobe is full stomach alters these relations. The ventral
positioned in the mesoduodenum in such a way surface (facies ventralis) of the left lobe of the
that its round, flat, caudal extremity lies in the pancreas is related ventrocaudally to the trans
concavity of the duodenal loop. By traction the verse colon and ventrocranially to the dorsal
gland can be separated for a distance of about wall of the stomach.
3 cm. from the various parts of the duodenum An accessory pancreas (pancreas accesso-
which forms the loop since the mesoduodenum rium) is occasionally found in the dog. Baldyreff
at this place is loose. (1929) cites cases in which the aberrant gland
As the right lobe runs obliquely cranially was located in the wall of the gall bladder and
toward the pylorus it becomes narrow and flat in the caudal part of the great mesentery. Pan
tened dorsoventrally, so that dorsal and ventral creatic bladders have been described by various
surfaces are formed. Upon contacting the initial authors as occurring in the cat, but none have
part of the descending duodenum it becomes been recorded in the dog (Boyden 1925).
molded to this organ. The caudal part of the
right lobe of the pancreas is related to the sub
lumbar fat containing the ureter and to the ven Ducts of Pancreas
tral surfaces of the right kidney and the caudate
process of the liver. The right lobe of the pan The pancreas nearly always has two excre
creas is related ventrally to the ileum and cecum tory ducts (Figs. 13-14, 13-15), in conformity
caudally, and to the ascending colon cranially. with the dual origin of the gland, one anlage
Loops of the jejunum contact those portions of arising dorsally from the duodenum and the
its ventral surface which are not already in con other ventrally at the termination of the bile
tact with more fixed viscera. In some specimens duct (Revell 1902). These two ducts usually
the right lobe of the pancreas and the adjacent intercommunicate within the gland, since the
descending part of the duodenum have gravi parenchyma of the whole gland is elaborated
tated lateral and even ventral to the jejunal coils. around them. In the adult, the two portions of
The pancreatic angle (angulus pancreatis, or the gland are fused without any demarcation to
corpus pancreatis [N.A.]) unites the two lobes indicate their dual origin. Revell, however,
of the pancreas in an angle of about 45 degrees, points out that, when the two ducts do not com
which is open sinistrocaudally. Cranially, it lies municate within the gland, the ventral pan
closely applied to the caudosinistral portion of creatic duct drains the right lobe and the dorsal
P a n crea s 709
duct drains the left lobe. Although this is the the two lobes. In type 5 (8 per cent), there were
basic pattern by which the pancreatic ducts three orifices into the duodenum from the ducts
form in the domesticated mammals, great varia from the pancreas and in one specimen there
tions exist among the different species and were two additional small ducts, one emerging
within the same species. on either side of the minor duodenal papilla.
The larger or main excretory duct of the pan Other variations of the ducts of the canine
creas of the dog is the ventral pancreatic duct pancreas exist, as Revell (1902) and Mintzlaff
(ductus pancreaticus ventralis). The ventral (1909) have shown. The main duct from each
duct is the smaller one in man, and is known lobe occupies the approximate center of the
as the ductus pancreaticus accessorius. From lobe and is joined at right angles by tributaries
its formation at the union of the ducts from the from the adjacent parenchyma. Because the
two lobes in the dog to the site where it perfo gland is ribbon-like, the small ducts from the
rates the intestinal wall, the ventral pancreatic adjacent parenchyma enter largely on opposite
duct is about 3 to 4 mm. long and 2 mm. wide. sides, the openings being spaced at 0.5 to 1.5
The union of the two lobar ducts to form the cm. intervals.
main duct may occur at any level up to the The opening of the smaller, or ventral, pan
intestinal wall, or, rarely, the two ducts may creatic duct is closely associated with that of
open separately (Revell 1902). The dorsal pan the bile duct. In two out of three specimens
creatic duct (ductus pancreaticus dorsalis) is Eichhom and Boy den (1955) found the slitlike
the smaller duct in the dog. In man it is the orifice of the ventral pancreatic duct located
larger pancreatic duct of the two, and is known distal to that of the bile duct; others have de
simply as ductus pancreaticus. According to scribed this opening as proximal to that of the
Bottin (1934), the two pancreatic ducts open bile duct. The main or dorsal pancreatic duct
separately into the duodenum in about 75 per usually opens into the duodenum 28 mm. from
cent of dog specimens and they always com the opening of the bile duct into the duodenum,
municate with each other in the gland. or approximately 8 cm. from the pyloric sphinc
In the 50 dogs of all ages and breeds and of ter (Nielsen and Bishop 1954). Its entry into
both sexes which Nielsen and Bishop (1954) the duodenum resembles that of the bile duct
studied by the use of radiopaque mediums, the in that a ridge of mucosa is formed, with a slight
duct system of the canine pancreas could be elevation at its distal end on which the opening
divided into five main types. In type 1 (46 per is located.
cent) a single main duct, formed by a tributary The dorsal pancreatic duct, like the ventral
from each lobe uniting in a Y junction, entered duct, but unlike the bile duct, runs through the
the duodenum at the (minor) ventral duodenal duodenal wall rather directly. The opening
papilla. In this group there was also an addi through the mesenteric wall of the proximal
tional duct, arising from the left lobar duct, portion of the descending duodenum is fre
which frequently followed a most indirect and quently located to the left of the cranial pan
tortuous route and entered the duodenum at or creaticoduodenal vessels, whereas the bile and
near the major duodenal papilla. Type 2 (22 ventral pancreatic ducts open to the right of
per cent) was similar to type 1, except that the these vessels. Eichhom and Boyden (1955) have
small dorsal duct arose from the right lobar duct reported on the musculature of the pancreatic
instead of the left, and crossed over the duct of and bile ducts of the dog, and Mann, Foster,
the left lobe before entering the duodenum. In and Brimhall (1920) have described the rela
type 3 (16 per cent), each lobe had its own ex tions of the common bile duct to the pancreatic
cretory duct. The ducts crossed, the one from ducts in 15 species of common laboratory and
the right lobe emptying into the dorsal duo domestic mammals.
denal papilla with the bile duct and that from
the left lobe emptying on the ventral duodenal
papilla. A fine and often tortuous shunt con
nected the two ducts. In type 4 (8 per cent), the Blood and Lymph Vessels
ducts from the two lobes anastomosed in the Y
formation, but there was no other duct leading The main vessels to the right lobe of the pan
into the duodenum or into the bile duct. In two creas are the pancreatic branches of the cranial
of the four specimens there was a small anasto and caudal pancreaticoduodenal arteries which
mosis within the pancreas between the ducts of anastomose in the gland. The left extremity of
710 Chapter 13. T he D ig e s t iv e Sy stem and A bd o m en
the left lobe of the pancreas is primarily sup Arnall, L. 1961. Some aspects of dental development in the
plied by the pancreatic branch of the splenic dog; calcification of crown and root of the deciduous
dentitions. J. small Anim. Pract. 1: 169-173.
artery. It also receives small branches from the Baldyreff, E. B. 1929. Report of an accessory pancreas on the
common hepatic artery as this vessel may ileum of a dog. Anat. Rec. 43: 47-51.
groove the dorsal surface of the organ, and the Baum, H. 1918. Das Lymphgefasssystem des Hundes. Berlin,
left limb regularly receives, near the pancreatic Hirschwald.
angle, one or two branches from the gastroduo Bennett, G. A. 1944. The lyssa of the dog. (Abstr.) Anat. Rec.
88: 422.
denal artery. Small pancreatic branches directly Bennett, G. A., and R. C. Hutchinson. 1946. Experimental
from the celiac may supply a small portion of studies on the movements of the mammalian tongue; the
the left limb of the pancreas near its free end. protrusion mechanism of the tongue (dog). Anat. Rec.
The caudal pancreaticoduodenal vein, a satel 94: 57-83.
Bennett, G. A., and A. J. Ramsay. 1941. Experimental studies
lite of the artery of the same name, is the princi on the movements of the mammalian tongue; move
pal vein from the right pancreatic lobe. It is the ments of the split tongue (dog). Anat. Rec. 79: 39-51.
last tributary to enter the cranial mesenteric Bensley, R. R. 1902. The cardiac glands of mammals. Amer.
vein and, unlike the intestinal veins which J. Anat. 2: 105-156.
Blakeley, C. L. 1957. Otorrhea and Surgical Drainage. Pp.
empty into it, it enters the larger vessel from the 309-320 in Canine Surgery, edited by K. Mayer, J. V.
cranial side. The left lobe of the pancreas is Lacroix, and H. P. Hoskins. 4th Ed. Evanston, 111., Amer
drained primarily by two veins which terminate ican Veterinary Publications, Inc.
in the last 2 cm. of the splenic vein. The satellite Bloom, W., and D. \V. Fawcett. 1962. Textbook of Histology.
of the small branch of the cranial pancreatico 8th Ed. Philadelphia, W. B. Saunders Co.
Bottin, J. 1934. Contribution a l’Etude de l’Anatomie des
duodenal artery which supplies the left lobe near Canaux Excreteurs du Pancreas chez le Chien. C. R.
the pancreatic angle drains this part of the Soc. biol. (Paris) 117: 825-827.
gland. Boyden, E. A. 1925. The problem of the pancreatic bladder.
The lymphatics from the pancreas drain into Amer. J. Anat. 36: 151-183.
Bradley, O. C., and T. Grahame. 1948. Topographical Anat
the duodenal lymph node, if present, and into omy of the Dog. 5th Ed. London, Oliver & Boyd.
the hepatic, splenic, and mesenteric lymph Brown, M. E. 1937. The occurrence of arteriovenous anasto
nodes. moses in the tongue of the dog. Anat. Rec. 69: 287-292.
Casas, A. P. 1958. Contribution a 1’etude du sphincter d’Oddi
Nerves chez Canis familiaris. Acta anat. (Basel) 34: 130-153.
Chauveau, A. 1886. The Comparative Anatomy of the Do
mestic Animals. New York, D. Appleton & Company.
Most sympathetic fibers come from the celiac Chiu, S. L. 1943. The superficial hepatic branches of the vagi
plexus and reach the organ by following the and their distribution to the extrahepatic biliary tract in
pancreatic branches of the cranial pancreatico certain mammals. Anat. Rec. 86: 149-155.
Colyer, F. 1936. Variations and Diseases of the Teeth of Ani
duodenal and celiac arteries. It is probable that mals. London, J. Bale Sons and Danielson, Ltd.
the caudal part of the right lobe receives sympa Dechambre, M. 1912. Absence totale des dentes chez un
thetic fibers from the cranial mesenteric plexus chien. Rec. Med. v6t. 89: 67-68.
which follow the caudal pancreaticoduodenal Dyce, K. M. 1957. The muscles of the pharynx and palate of
the dog. Anat. Rec. 127: 497-508.
artery and its pancreatic branches. McCrea
Eichhorn, E. P., Jr., and E. A. Boyden. 1955. The choledo-
(1924) states that, in the dog, vagal (parasympa choduodenal junction in the dog—a restudy of Oddi’s
thetic) fibers reach the pancreas as fine twigs sphincter. Amer. J. Anat. 97: 431-451.
which run with the splenic branch of the celiac Elias, H. 1949. A re-examination of the structure of the mam
artery and with the cranial mesenteric artery, malian liver; parenchymal architecture. Amer. J. Anat.
84: 311-333.
presumably along the caudal pancreaticoduo Ellenberger, W. 1911. Handbuch der vergleichenden mikro-
denal branch. These findings accord with those skopischen Anatomie der Haustiere. Vol. 3. Berlin, Paul
of Richins (1945) in the cat. Parey.
Ellenberger, W., and H. Baum. 1943. Handbuch der ver
gleichenden Anatomie der Haustiere. 18th Ed. Berlin,
Springer.
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Dis. 16: 344-348. tic Animals. 4th Ed. Philadelphia, W. B. Saunders Co.
Martin, P. 1923. Lehrbuch der Anatomie der Haustiere. Vol. St. Clair, L. E., and N. D. Jones. 1957. Observations on the
4. Stuttgart, Schickhardt and Ebner. cheek teeth of the dog. J. Amer. vet. med. Ass. 130:
McCarrell, J. D., S. Thayer, and C. K. Drinker. 1941. The 275-279.
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McCrea, E. D. 1924. The abdominal distribution of the vagus. 19. The Wistar Inst. Anat. Biol., Philadelphia.
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mit besonderer Beriicksichtingung der Zahnaltersbestim- New York, Paul B. Hoeber.
mung und der Zahnanomalien. Vet. Diss., Zurich. Tims, H. W. 1902. On the succession and homologies of the
Michel, G. 1956. Beitrag zur Topographie der Ausfuhrungs- molar and premolar teeth in mammalia. J. Anat. Physiol.
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animals. Amer. J. vet. Res. 16: 152-157.
712 Chapter 13. T he D ig e s t iv e S y stem and Abd o m en
Torgersen, J. 1942. The muscular build and movements of the Williams, T. 1935. The anatomy of the digestive system of
stomach and duodenal bulb. Acta radio! (Stockh.) Suppl. the dog. Vet. Med. 30: 442-444.
45: 1-191. Wmkelstein, A., and P. W. Aschner, 1924. The pressure fac
Trautmann, A., and J. Fiebiger. 1957. Fundamentals of the tors in the biliary duct system of the dog. Amer. J. med.
Histology of Domestic Animals. (Translated and revised Sci. 168: 812-819.
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Habel and E. L. Biberstein.) Ithaca, N.Y., Comstock genetic significance of the presence of a third upper
Publishing Assoc. molar m the modem dog. Amer. Midland Nat. 14:36-48.
Warren, R. 1939. Serosal and mucosal dimensions at different Zietzschmann, O. 1938. Lage und Form des Hundemagens.
levels of the dog’s small intestine. Anat. Rec. 75: 427- Berl. Munch, tierarztl. Wschr. 1 0 :138-141, and Vet. Rec.
437. 50: 984-985.
Webb, R. L., and P. H. Simer. 1940. Regeneration of the --------------- 1939. Das Mesogastrium dorsale des Hundes mit
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Williams, R. C. 1961. Observations on the Chronology of De Jb. 83: 325-358.
ciduous Dental Development in the Dog. Thesis, Cornell
University.
C H A P T E R 14
TH E RESPIRA TO RY SYSTEM
The respiratory system consists of the lungs brachycephalic breeds, the shortened muzzle
and of the air passageways which lead to the often is the cause of respiratory difficulties.
sites of gaseous exchange within the lungs.
Various structures associated with these pas External Nose
sageways modify or regulate the flow of air,
serve as olfactory receptors, facilitate water and The external nose consists of a fixed bony case
heat exchange, and make phonation possible. and a movable cartilaginous framework. The
The nasal cavity and turbinates warm and mois cartilaginous portion is movable or distensible
ten the air, and remove foreign material from it. by virtue of several skeletal muscles associated
The pharynx serves as a passageway for both with the muzzle. The short hair on the skin of
the respiratory and the digestive system. The the nose is directed caudally on the mid-dorsal
larynx guards the entrance to the trachea, func surface and gradually slopes in a posteroventral
tions in vocalization, and regulates both the in direction laterally, where it is continued on the
spiration and expiration of air. The trachea is a lips. The apical portion of the nose is flattened
non-collapsible tube, lined by ciliated epithe and devoid of hair. It is called the nasal plane
lium. It divides into the principal bronchi, and (planum nasale). The integument of the nasal
the air passageways continue in the two lungs plane presents epithelial elevations or papillary
as lobar bronchi, segmental bronchi, bronchi ridges which result in patterns characteristic for
oles, alveolar ducts, alveolar sacs, and alveoli. each individual. For this reason nose prints may
The terminal divisions are located in the elastic, be used as a means of identification in the dog,
well vascularized lungs, which passively expand similar to the way finger prints are used in man
and collapse in response to changes in intratho (Homing et al. 1926).
racic pressure, created by action of the muscles The lateral walls of the bony portion of the
of the diaphragm and thoracic wall. nose are formed by the incisive bones and max
illae, whereas the roof is formed by the paired
nasal bones. The concave anterior ends of the
NOSE AND NASAL PORTION nasal bones, dorsally, and the incisive bones,
OF THE PHARYNX laterally and ventrally, bound the largest open
ing into the skull. This opening, called the piri
N o se form aperture (apertura piriformis), is wider
ventrally than dorsally and lies in an oblique
The nose (nasus), in a broad sense, refers to plane. In life this opening is bounded anteriorly
the external nose (nasus externus) and its associ by the nasal cartilages. The aggregate of these
ated nasal cartilages (cartilagines nasi), as well cartilages, with their ligaments and covering
as to the internal nose, or nasal cavity (cavum skin, comprises the movable portion of the nose
nasi). In terms relating to nasal structures or dis (pars mobilis nasi). The movable part of the nose
eases the root rhin-, from the Greek rhinos for ends in a truncated apex (apex nasi).
nose, is frequently employed. The facial portion
of the respiratory system and the anterior por Cartilages of the Nose
tions of the upper and lower jaws collectively
constitute what is called the muzzle. In dolico- The mobile part of the external nose has a
cephalic breeds the muzzle is long and may ac framework composed entirely of the nasal carti
count for half of the total length of the skull. In lages (Fig. 14-1). These include the unpaired
713
714 Chapter 14. T he R e s p ir a t o r y S y s t e m
septal cartilage, the paired dorsal parietal and parietal cartilage is greatly thickened anteriorly
ventral parietal cartilages, and the paired acces and contains a plexus of blood vessels which
sory cartilages. Related to the ventral part of form a meshwork in the collagenous tissue
the septal cartilage is the vomeronasal cartilage. directly posterior to the nostril. It becomes
The septal cartilage of the nose (cartilago much thinner posteriorly. At a transverse plane
septi nasi) is a perpendicular median plate through the piriform aperture it blends with the
which separates most of the nasal cavity into anterior extremity of the maxilloturbinate. The
right and left nasal fossae. It is an anterior con free medial border of the dorsal parietal carti
tinuation of the perpendicular plate of the eth lage curves ventral to the thicker free lateral
moid bone which fails to ossify. In the region of border of the ventral parietal cartilage posterior
the piriform aperture of the osseous skull, it is to a transverse plane through the posterior an
lacking over a distance of about 1 cm. so that gle of the mid-lateral slit of the nostril.
the nasal septum in this region is formed of the The ventral parietal cartilage (cartilago parie-
membranous nasal septum (pars membranacea talis ventralis) is a continuation of the anterior
septi nasi), which connects the cartilaginous im portion of the lateral half of the septal cartilage.
movable posterior part with the mobile anterior Posteriorly its origin moves obliquely dorsad on
part. the lateral surface of the ventral part of the sep
The caudal part of the cartilaginous nasal tal cartilage. It is slightly shorter and about one-
septum is thicker ventrally, where it lies in the fourth as wide as the dorsal parietal cartilage.
septal groove of the vomer, than it is dorsally, As it rolls dorsally it is neither of uniform thick
where it blends with the thin conjoined ventral ness nor of uniform curvature. Anteriorly, it
processes of the nasal bones. It presents a promi runs forward into the apex of the nose. It ends
nent caudal process which occupies the space posterior to the lateral leaf of the septal carti
between the osseous nasal septum dorsally and lage adjacent to the articulation of this cartilage
the groove in the vomer ventrally. with the accessory cartilage. Posteriorly it as
The anterior part of the cartilaginous nasal sumes a sigmoid shape, in cross section, being
septum continues a median course forward from bent in such a way that its free border is added
the membranous portion of the nasal septum to the free border of the dorsal parietal carti
(Fig. 14-2). It lies between the right and left lage in forming the cartilaginous basis of the
nasal vestibule. The anterior border of this fold which continues the maxilloturbinate bone
portion of the septum is divided into right and to the vestibule.
left laminae. The cleft between the two leaves The accessory cartilage (cartilago accessoria)
is deeper and much wider ventrally than it is is a laterally convex leaf which articulates with
dorsally. It forms a depressed triangular area the ventrolateral angle of the wide ventrally
ventrally. The accessory cartilage is united by divided portion of the septal cartilage and ex
collagenous tissue to the ventral parietal carti tends dorsoposteriorly to the lateral surface of
lage. The dorsal portion of each lamina is rolled the expanded portion of the dorsal parietal car
laterally to form the dorsal parietal cartilage. tilage. For a considerable portion of its length
Arising from the ventral portion of the anterior it lies directly under the integument that covers
part of the septal cartilage is the small ventral the ventral surface of the mid-lateral slit in the
parietal cartilage, which turns upward and in nostril.
ward toward the dorsal parietal cartilage (Fig. A second small accessory cartilage is occa
14-1 D). sionally located directly dorsal to the septal
The dorsal parietal cartilage (cartilago parie- cartilage in the groove formed by the origins of
talis dorsalis) is the most expansive of the carti the right and left dorsal parietal cartilages. Its
lages in the mobile part of the external nose. On position is only a few millimeters anterior to the
each side it is a continuation of half of the dorsal intemasal suture.
portion of the septal cartilage. From this dorsal The mobile part of the nose is moved and the
origin it is rolled into a tube by curving out shape of the nostrils is altered by the action of
ward, downward, and inward. It is about 2.5 intrinsic muscles and of the nasal part of the
cm. long. Its widest portion is its anterior half, maxillonasolabialis and of the levator nasolabi
which in large heads attains a width of 1 cm. alis muscles which are inserted on these car
Posteriorly, it joins the dorsal part of the piri tilages.
form aperture, to which it is attached by fibrous The vomeronasal cartilage (cartilago vomero-
tissue along the concave border of the nasal nasalis), which encloses the vomeronasal organ,
bone. The free rolled-in border of the dorsal is not a complete cartilaginous tube in the dog.
N o se and N a s a l P o r t io n of the P h aryn x 715
/D orsa l n a s a l lig.
/ L o c a t io n of s e s a m o i d cart.
L a t na sal lig.
Dorsal p a ri e ta l
c a r t i lage
-Cartilagin ous
s e p t um
'Accessory
c a r t i lage
NVentral p a r i e t a l
cartilage
Cartilaginous septum
- -Dors, p a r i e t a l cart.
- Ven t , p a r i e t a l c a r t .
•Accessory c a r t i l a g e
According to Negus (1958), the cartilage lies at The alar fold (plica alaris), which is an exten
first on the lateral side of the vomeronasal or sion of the maxilloturbinate, terminates within
gan, forms a curved plate which arches over the the vestibule by a bulbous enlargement that
organ, and then continues along its medial sur fuses to the wing of the nostril. The wing of the
face. nostril (ala nasi) is the thickened dorsolateral
portion of the nostril. The wing of the nostril
Ligaments of the Nose contains much of the dorsal parietal and acces
Three ligaments, comprised of one paired and sory cartilages. It is the most mobile portion of
one unpaired, attach the mobile part of the nose the nostril, because it receives the terminal fi
to the dorsal portion of the osseous muzzle (Fig. bers of the nasal portions of the mm. maxillo-
14-1 B). The dorsal nasal ligament (lig. nasale nasolabialis and levator nasolabialis.
The nasal vestibule (vestibulum nasi) is not
dorsale) is a single band of collagenous tissue
an empty antechamber, as it is in man, but
which runs from the dorsal accessory cartilage
rather it is largely obliterated by the large bul
to the dorsum of the nasal bones. The lateral
bous end of the alar fold which extends into it.
nasal ligament (lig. nasale laterale), one on
Because the end of the alar fold is fused to the
either side, is a collagenous band which runs
inner surface of the wing of the nostril, it acts
from the mid-lateral surface of the dorsal parie
to divert the incoming air. Upon entering the
tal cartilage to the border of the piriform aper
vestibule through a nostril, air is diverted medi
ture directly dorsal to the end of the nasomaxil
ally and ventrally into the largest meatus of the
lary suture. The ligaments of the nose are best
nose. The nasolacrimal duct (ductus nasolacri-
developed in old dogs of the working breeds. In
malis), which conducts the lacrimal secretion
small, young dogs they cannot be isolated satis
from the eye, opens into the vestibule by a
factorily.
minute orifice located at the anterior end of the
The Nasal Cavity attached margin of the alar fold. No cilia are
present on the mucosa of the vestibule.
The nasal cavity (cavum nasi) is the internal After the inhaled air leaves the nasal vesti
nose, or facial portion of the respiratory pas bule it traverses the longitudinal nasal meatuses
sageway (the cavity of the external nose). It to reach the nasal part of the pharynx.
extends from the nostrils to the choanae, being The dorsal nasal meatus (meatus nasi dorsalis)
divided into right and left halves by the nasal is a passage through the dorsal part of the nasal
septum. Each half of the nasal cavity is known fossa. It lies between the dorsal surface of the
as a nasal fossa. nasal concha, or turbinate, and the ventral sur
Each nasal fossa (fossa nasalis) begins at the face of the nasal bone. Laterally it is limited by
nostril with the nasal vestibule and ends with the nasoturbinate crest, from which the nasal
the nasopharyngeal meatus and choana. The concha or nasoturbinate bone arises. Medially
nasal fossa is divided into four principal air the dorsal nasal meatus becomes confluent with
channels and several smaller ones (Fig. 14-3). the common nasal meatus.
During development the growth of laminae The middle nasal meatus (meatus nasi me
from the lateral and dorsal walls of the nasal dius) lies between the ventral part of the nasal
fossa results in the formation of turbinate scrolls turbinate bone dorsally and the dorsal part of
which largely fill the cavity and restrict the flow the numerous scrolls composing the maxillotur
of air (see pages 24 to 31). The air passages thus binate bone ventrally. Throughout the long
created between the turbinates are called the middle portion of the middle nasal meatus its
nasal meatuses. width is approximately 1 mm. At its anterior
The nostril (naris), the opening into the nasal end it presents a dilatation, and posteriorly its
vestibule, is a curved opening which is much lateral portion is divided into several parts.
wider dorsomedially than it is ventrolaterally. The atrium of the middle meatus (atrium
It possesses more than usual importance, be meatus medii) is an ellipsoidal dilatation which
cause in some brachycephalic dogs the opening connects the nasal vestibule with the middle
is too restricted and interferes with respiration. nasal meatus. The atrium is formed ventrally by
Leonard (1956) devised an operation whereby the narrow handle of the club-shaped mucosal
the transverse diameter of the nostril may be alar fold which runs forward and upward from
increased. Satisfactory results were obtained in the maxilloturbinate bone. Dorsally it is
all dogs operated upon. bounded by the relatively straight anterior por-
N o se and N a s a l P o r t io n of th e P harynx 717
iFnon fa I bone
bone
fro n ta l sinus
, , Cri bri form p lo fe of e th m o id bone
tion of the nasal turbinate fold. In large mesati on the ethmoturbinates, which lie in the pos
cephalic heads it is approximately 5 mm. deep, teromedial and posterodorsal parts of the nasal
5 mm. wide, and 2 cm. long. cavity. The olfactory epithelium is characterized
Laterally, the posterior part of the middle by three cell types: receptor cells, supporting
nasal meatus is divided by the scrolls of the sec cells, and basal cells. For a review of the mor
ond endoturbinate from the ethmoid bone into phology of the olfactory system in vertebrates,
several air passages which lie between these see Allison (1953).
scrolls. Under normal conditions of inspiration, the
The ventral nasal meatus (meatus nasi ven respiratory and olfactory currents of air are as
tralis) is located between the maxilloturbinate sociated. When a dog deliberately wants to
scrolls and the dorsal surface of the hard palate. sample the environment, the nostrils are dilated,
It is narrow anteriorly as it leaves the nasal ves and with a forced inspiration the dog sniffs the
tibule. It gradually widens posteriorly, and at air. This act provides a greater volume of in
tains a width of 1 cm. at the large nasomaxillary spired air, which takes a more dorsal course
opening, where it continues ventral to the around the ethmoturbinates, where the olfac
transverse lamina or floor plate of the ethmoid tory receptors are most numerous.
bone as the nasopharyngeal meatus. This wide Stratified squamous epithelium continuous
portion of the nasal fossa is located in a trans with that of the naris extends into and lines the
verse plane through the posterior portions of nasal vestibule. It then gradually changes into
the fourth upper premolar teeth, where the stratified columnar and finally into ciliated pseu-
middle, ventral, and common nasal meatuses dostratified epithelium, which is characteristic
converge. for the respiratory passage (Trautmann and Fie-
The common nasal meatus (meatus nasi com biger 1957).
munis) is a longitudinal narrow space on either
side of the nasal septum. Laterally, it is bounded Glands of the Nose
by the nasal and maxilloturbinate bones. Above,
below, and between these bones it is coexten The lateral nasal gland (glandula nasalis later
sive with the dorsal, middle, and ventral nasal alis) is a serous gland which is located in the
meatuses, respectively. mucosa of the maxillary recess near the open
The nasopharyngeal meatus (meatus naso- ing of this recess into the nasal fossa. Its duct or
pharyngeus) extends on either side from the ducts open into the middle nasal meatus. The
caudal dilated portion of the ventral nasal meatus lamina propria of the mucosa of the respiratory
to the choana. It is a short passage with a much part also contains serous, mucous, and mixed
longer lateral than medial wall. It is bounded tubuloalveolar glands. These glands are also
laterally by the maxillary and palatine bones, present in the mucosa of the nasal vestibule.
dorsally by the transverse lamina of the ethmoid Goblet cells are present throughout the respira
bone, ventrally by the palatine bone, and medi tory region, and olfactory glands which contain
ally by the vomer. yellow pigment granules are located in the
The choanae are the openings of the two olfactory epithelium (see Chapter 17).
nasopharyngeal meatuses into the nasal portion The nasolacrimal duct (ductus nasolacrimalis)
of the pharynx. They are oval in shape and carries the serous secretion from the conjunc
oblique in position. tival sac to the nasal vestibule. A characteristic
The paranasal sinuses, which are also con of a healthy dog is a moist nose, yet there are
nected with the respiratory passageways, are no glands under the integument which covers
described with the skeletal system. the nasal plane. It is probable that this fluid rep
resents the combined secretions of the vestibu
lar and lateral nasal glands.
Nasal Mucosa
Function of Internal Nose
The various types of epithelia which line the
nasal cavity and coat its associated structures The internal nose functions to moisten and
are spoken of collectively as the nasal mucosa. warm the inspired air. According to Negus
The transition from the more peripheral non (1958), the mucosa of the maxilloturbinate is
olfactory type of epithelium to the deeper-lying the chief participant in these functions. Al
olfactory epithelium is not abrupt. The recep though no cilia are present in the vestibule, for
tors for the sense of smell are located primarily eign particles are trapped by the mucus from
L arynx 719
the goblet cells. The submucosa is extremely to the atlas. It is about 6 cm. long in a medium
vascular. sized dog, nearly half of this length being oc
cupied by the epiglottic cartilage, which lies in
front of the laryngeal opening. The intrinsic
N a s a l P o r t io n of P harynx
muscles of the larynx (Figs. 14-8, 14-10, 14-11)
control the size of the laryngeal inlet, the size
The nasal portion of the pharynx (pars nasalis
and shape of the glottis, and the positions of the
pharyngis), also called the nasal pharynx or
laryngeal cartilages.
nasopharynx, extends from the choanae to the
pharyngeal isthmus. The pharyngeal isthmus
(Fig. 14-4) is formed cranial to the larynx by Cartilages of the Larynx
the crossing of the digestive passageway (oral
pharynx) and the respiratory passageway (nasal The laryngeal cartilages (cartilagines laryngis)
pharynx). The anterior part of the nasal pharynx (Figs. 14-6 to 14-13) are the epiglottic, thyroid,
is bounded by the hard palate ventrally, the vo cricoid, arytenoid, sesamoid, and interarytenoid
mer dorsally, and the palatine bones bilaterally. cartilages. Only the arytenoid cartilage is paired.
Although the middle and posterior portions of The epiglottic cartilage (cartilago epiglottica)
the pharynx are bounded dorsally by the base forms the basis of the epiglottis. In outline the
of the skull and the muscles which attach to it, cranial margin of the cartilage forms a thin, dor
its ventral boundary is the mobile, long soft pal sally concave triangle with its apex pointing for
ate (Fig. 14-5). At each act of swallowing the ward. The epiglottis resembles a sharp-pointed
cavity of the posterior part of the nasal pharynx spade. Its dorsocaudal surface, known as the
is obliterated by the pressure of the material aboral surface (facies aboralis), is concave. The
swallowed and the root of the tongue forcing opposite surface, called the oral surface (facies
the soft palate dorsally. oralis), because it faces the oral pharynx, is con
On each lateral wall of the nasal pharynx, vex. The oral surface is attached to the middle
above the middle of the soft palate, is an oblique of the body of the hyoid bone by the short, stout
slitlike opening, about 5 mm. long, which is the hyoepiglottic muscle. On either side of the me
pharyngeal opening of the auditory tube (ostium dian mucous fold which covers this muscle is a
pharyngeum tubae auditivae). The opening is deep pocket of mucosa, called the vallecula,
located directly posterior to the posterior bor which may attain a depth of 1.5 cm. Each val
der of the pterygoid bone and faces anteroven- lecula is limited laterally by a small fold of strati
trally. fied squamous epithelium running from the oral
The auditory tube (tuba auditiva), or eusta- surface of the epiglottis near its caudolateral
chian tube, extends between the cavity of the angle to the lateral wall of the laryngeal part of
middle ear and the cavity of the nasal pharynx. the pharynx. The stalk of the epiglottis (petiolus
It serves to equalize the atmospheric pressure epiglottidis) is in the form of a thickened handle
on the two sides of the tympanic membrane. of fibrous tissue which unites the mid-caudal
portion of the epiglottis and the dorsal cranial
surface of the thyroid cartilage.
LARYNX The thyroid cartilage (cartilago thyroidea) is
the largest cartilage of the larynx. It forms the
The larynx is a musculocartilaginous organ, middle portion of the laryngeal skeleton and is
leading to the trachea, which serves for vocal open dorsally. It consists of right and left lami
ization and prevents the inspiration of foreign nae (lamina dextra et sinistra), which are united
material. The valvular function of the larynx, ventrally to form a short but deep trough. An
by means of the epiglottis, is vital, since it is inconspicuous oblique line (linea obliqua) serves
across its inlet that all substances swallowed primarily for the insertion of the sternothyroid
must pass in their course from the oral pharynx muscle. Each lamina is expanded dorsally to
through the laryngeal pharynx to the esophagus. form transversely thin processes, the cranial and
The function of vocalization is most important caudal cornua (cornu cranialis et caudalis). Sep
in the hound breeds of dogs. Negus (1949) has arating the cranial and caudal thyroid cornua
described and illustrated the comparative anat from the thyroid laminae on each side are the
omy of the larynx from fish through mammals. cranial and caudal thyroid notches (incisura
The larynx is located directly posterior to the thyroidea cranialis et caudalis), respectively.
root of the tongue and the soft palate, ventral The cranial laryngeal nerve and the laryngeal
720 Chapter 14. T he R e s p ir a t o r y System
--------
NASAL
NASAL PHARYNX
C A V IT Y
I W w p a l a t T - '
TRACHEA
ESOPHAGUS
'p h a r y n x
1 ORAL PHARYNX O R A L C A V IT Y
Fic. 14—
4. Diagram showing relation of portions of pharynx to esophagus and trachea.
A. During normal respiration.
B. During deglutition.
M .p a lo t i n u s
t p i g l o f t is
CornaSSiol to o th .
- M u c o s a c o v e r i n g ker ot ohyoi d
Under surface
of mucosa o f -
nasal p h a ry n x Tongue pulled back
^Posterior b o r d e r o f s o f t p a l o t e
artery pass through the cranial thyroid notch. varies greatly in different species of mammals,
Where the two laminae fuse ventrally a slight so that what may appear as a process of the ary
laryngeal prominence (prominentia laryngea) is tenoid in one species may be a separate car
formed. The laryngeal prominence, known as tilage in another. In the dog, the arytenoid
the “Adam’s apple” in man, is not visible ex cartilage embodies the comiculate cartilage and
ternally in the dog, but it can be palpated. the cuneiform cartilage of other mammals. As
The caudal border of the thyroid cartilage a result, this compound cartilage may be de
possesses the median deep caudal thyroid notch scribed as possessing a comiculate process, a
(incisura thyroidea caudalis), whereas the cra muscular process, a vocal process, and a cunei
nial border is slightly convex from side to side. form process.
The caudal border of the thyroid cartilage is The articular surface (facies articularis) is a
united to the ventral arch of the cricoid carti slightly oval, concave facet on the caudal border
lage by the cricothyroid ligament (ligamentum of the arytenoid, which faces caudomedially
cricothyroideum). The cranial border is joined and receives the cricoid articular facet, to form
to the thyrohyoid bones by the hyothyroid the cricoarytenoid articulation (articulatio cri-
membrane (membrana hyothyroidea). coarytenoidea).
The cricoid cartilage (cartilago cricoidea) is The muscular process (processus muscularis)
the only cartilage of the larynx which forms a is a relatively thick, rounded process which is
complete ring. The dorsal portion is approxi located directly lateral to the articular surface.
mately five times wider than the ventral portion. The m. cricoarytenoideus dorsalis inserts on this
The expanded dorsal part is the lamina of the process.
cricoid cartilage (lamina cartilaginis cricoideae). The comiculate process (processus comicu-
It possesses a median crest (crista mediana) for latus) is the longer and more caudal of the two
muscle attachment. Occasionally a pair of vas dorsal processes which form the dorsal margin
cular foramina are located in the lamina, one of the laryngeal inlet. In man and several other
on each side of the cranial portion of the crest. mammals it exists as a separate cartilage (car
The arch of the cricoid cartilage (arcus carti tilago comiculata), sometimes spoken of as
laginis cricoideae) extends ventrally from the Santorini’s cartilage.
lamina and completes the enclosure of the cau The vocal process (processus vocalis) is a cau
dal part of the cavity of the larynx. It is bilater dal ventral projection of the arytenoid cartilage.
ally concave in a transverse direction. The It is about 3 mm. thick and 5 mm. long at its
cricoid cartilage possesses two pairs of articular base. The vocal ligament and the m. vocalis,
facets. An indistinct pair of facets, for articula from the thyroid cartilage, attach to the vocal
tion with the apices of the caudal cornua of the process (Fig. 14-13).
thyroid cartilage, are located at the junction of The cuneiform process (processus cunei-
the lamina and the arch about 1 mm. from the formis) is the most cranial portion of the aryte
caudal border. A more prominent pair of facets, noid cartilage. It is connected by a narrow neck
for articulating with the arytenoid cartilages, (Fig. 14-6) to the main portion of the arytenoid
are located on the cranial border of the lamina cartilage, and is considered by some authors to
on each side of the median crest. Both pairs of be a separate cartilage. It exists as an inconstant
facets are enclosed in articular capsules and separate cartilage in man (cartilago cunei-
form synovial joints with the cartilages with formis), sometimes spoken of as Wrisberg’s car
which they articulate. The sides of the ventral tilage. In the horse it is fused to the epiglottic
arch are gradually reduced in width ventrally. cartilage rather than to the arytenoid, and in
Mid-ventrally, in a medium-sized dog, the nar the ox it is lacking. In the dog, the cuneiform
rowest part of the arch is only 5 mm. long; in process is roughly triangular in shape. The ven
such a dog the mid-dorsal lamina would be ap tral portion lies in the aryepiglottic fold, and
proximately 2 cm. long. the dorsal portion aids in forming the laryngeal
The arytenoid cartilage (cartilago aryte- inlet. Attached to the cuneiform process are the
noidea) is an irregular cartilage, one on either ventricular ligament and the m. ventricularis
side, which articulates with the craniodorsal (Fig. 14-11). Duckworth (1912) suggested that
border of the cricoid cartilage. When the laryn the cuneiform cartilage arose in mammals from
geal cartilages are viewed laterally the aryte the lateral margin of the epiglottis.
noid is largely hidden from view by the thyroid The sesamoid cartilage (cartilago sesamoi-
lamina. dea) (Fig. 14-7) is an oval or dumbbell-shaped
The morphology of the arytenoid cartilage nodule located cranial to the cricoid lamina and
722 C h ap ter 14. T he R e s p ir a t o r y System
Thyrohyoid T r a ch ea
Keratohyoi d -
Basihyoid
Thyroid c ar t il age —
*1 - S t al k
- - Cricothyroid
articulation
'St J - A r c h
C o r ni c ul at e p r o c e s s ,Cricoarytenoid
Cuneiform p r o c e s s ~ articulation
i Muscular
process
Vocal p r o c e s s -
H
F ig . 14-6. Laryngeal cartilages.
A. Scheme of laryngeal cartilages and
hyoid apparatus, lateral aspect.
B. Epiglottis, dorsal aspect.
C. Thyroid cartilage, lateral aspect.
D. Cricoid cartilage, lateral aspect.
E. Arytenoid cartilage, lateral aspect.
F. Arytenoid cartilage, medial aspect.
G. Interarytenoid cartilage.
H. Sesamoid cartilage, dorsal aspect.
L arynx 723
E p i hy o i d -
Keratohyoid
Basihyoid - Thy r oi d c a r t i l a g e
Thyrohyoid - i | j!j - Epiglottis
St y l oh yo i d -
Cran. cornu of thyroid c art i l age- - V o c a l process
Tymponohyoid c art i lage- - - Cune i f or m p r o c e s s A r y t e n o i d
+ -Epiglottis
L a t ventricle -
- - Laryngeal s a c cu l e
Cra ni al cornu of thyroid c a r t i l a g e - - III - -M. v e n t r i c u l a r i s
1 Vi- - M. t hy ro ar y t e n oi d e us
Laryngeal s a c c u l e " '
" M. a rytenoi deus tronsversus
Corniculate c a rtila g e ' -M. c r i c o o r y t e n o i d e u s dorsalis
Cricoid cartiloge ' ' ' Caudal cornu of thyroid c ar t i l a g e
Trachea'
F ig . 14-8. Laryngeal muscles, dorsal aspect. (The right corniculate cartilage has been cut
and the right laryngeal saccule reflected.)
724 C h ap ter 14. T he R e s p ir a t o r y System
between the arytenoid cartilages. It is occasion close together to form the interarytenoid groove
ally paired, in which case an intersesamoid liga (incisura interarytenoidea). This groove com
ment or fibrous union joins the two. Primarily, municates with the middle cranial portion of
the sesamoid cartilage appears to be intercal the laryngeal pharynx. The cuneiform tubercle
ated in the transverse arytenoid muscle. There (tuberculum cuneiforme) forms a relatively
is frequently a small contact surface with the heavy cone-shaped projection which is united
dorsal portion of each arytenoid cartilage. in a sagittal plane with the comiculate tuber
The interarytenoid cartilage (cartilago inter- cle; it is also united in a transverse plane to each
arytenoidea) is small, flat, and easily overlooked. lateral angle of the epiglottic cartilage by means
It lies cranial to the cricoid lamina and caudo- of a loose plica of mucosa. The channel lying
dorsal to the transverse arytenoid muscle and external to each aryepiglottic fold is called the
sesamoid cartilage. In this superficial position piriform recess (recessus piriformis). When food
the interarytenoid cartilage is embedded in or fluid is forced past the closed laryngeal open
connective tissue which attaches the arytenoid ing in deglutition it largely occupies these
cartilages and the cricoesophageal tendon to recesses, which constitute the ventrolateral por
the cricoid lamina. tions of the laryngeal pharynx.
The laryngeal vestibule (vestibulum laryngis)
extends from the laryngeal opening to the ven
Cavity of Larynx and Laryngeal Mucosa tricular folds. It is a funnel-shaped cavity which
opens freely dorsocranially. It is bounded ven
The cavity of the larynx (cavum laryngis) is trally by the mucosa covering the large, dorsally
divided into three transverse segments: the ves concave epiglottis. The cranial part of the wall
tibule, or antechamber, which opens in the on either side is also composed of the epiglottis.
pharynx by the aditus laryngis; a middle, nar Dorsocranial to the ventricular folds the flat
row portion, called the glottis; and the infra- tened cuneiform processes form its wall. The
glottic cavity, which lies caudal to the glottis. rim of the vestibule (rima vestibuli) is the cau
The laryngeal opening (aditus laryngis) lies dal opening of the vestibule. It is bounded
directly caudal to the pharyngeal isthmus and bilaterally by the vestibular folds and the mu
faces dorsocranially. Air entering or leaving cosa covering the cuneiform processes. The
the larynx can travel either by way of the nasal ventral boundary is formed by the mucosa cov
part, or by way of the oral part of the pharynx. ering the thyroid cartilage directly caudal to
In lolling (rapid breathing with the tongue the thyroepiglottic ligament.
hanging out), most of the air passes through the The vestibular (ventricular) fold (plica ves
mouth and oral pharynx; in slow, shallow tibularis) (Fig. 14-12) is a short, wide plica of
breathing it passes through the nasal fossae and mucosa, containing a few elastic fibers, which
nasal pharynx. The margin of the laryngeal runs from the expanded ventral margin of the
opening forms an imperfect triangle, with the cuneiform cartilage to the cranial dorsal surface
base located caudally. The margin of the epi of the thyroid cartilage. It is less than one-half
glottis forms its lateral boundaries and apex. the dorsoventral diameter of the larynx, and its
The caudal boundary is formed by the right and width approximates that of the vocal fold which
left aryepiglottic folds. lies caudal to it.
Each aryepiglottic fold (plica aryepiglottica) The glottis consists of the paired arytenoid
(Fig. 14-9) runs from the dorsal portion of the cartilages dorsally and the paired vocal folds
arytenoid cartilage and the closely associated ventrally which form a narrow passageway into
comiculate process to the caudolateral angle of the larynx, which is called the rima glottidis. The
the epiglottic cartilage. Two prominent tuber portion of the rima glottidis between the vocal
cles which are separated by a deep notch are folds is the intermembranous part (pars inter-
present in the fold. The more dorsocaudal tu membranacea); that between the medial surfaces
bercle is formed by the underlying comiculate of the arytenoid cartilages is the intercartilagi-
process and is called the comiculate tubercle nous part (pars intercartilaginea). The rima
(tuberculum corniculatum). It is a rounded glottidis is the most important part of the larynx,
process approximately 1 cm. long by 0.5 cm. from the standpoint of veterinary medicine, be
wide. It and the opposite tubercle form the cause it is the narrowest part of the laryngeal
most dorsal and the least expandable part of the passageway, and it contains the vocal folds which
laryngeal opening. Ventral to these tubercles are necessary for vocalization—barking, baying,
the dorsal parts of the arytenoid cartilages lie whining, and growling.
L arynx 725
Epiglo
Vocal f o l d -
Vestibular fo ld
L a t v entr i cl e -
— Laryngeal inlet
^Corniculate p r oc es s of arytenoid c a r t i l a g e
M. v e n t r i c u l a r i s
' tM. arytenoideus transversus
Cuneiform p r o c e s s of oryten oid c a r t i l a g e
i i i _' M. cricoarytenoi deus dorsalis
id c ar ti la ge
Epiglottis
Ar ti cul at io n with
t hyroid c a r t i l a g e
Laryngeal s ac cu l e
~M. cric o a r yt en o i de us lat.
M. t h y r o a r y t e n o i d e u s -M. vocal is
~Cri cothyroi d lig.
Thyroid carti
othy roideus
(reflected)
F ig . 14-10. Laryngeal muscles, lateral aspect. (The thyroid cartilage is cut to the left of the mid line and reflected.)
M. ventricularis
Corniculate process of arytenoid c a r t i l a g e
M. thy r o a r y t e n o j
Vo c al lig. ( Cricothyroid lig.
'M. vocal is
F ig . 14-11. Laryngeal muscles, lateral aspect. (The thyroid cartilage is cut to the left of the mid line and removed. The mm.
thyroarytenoideus, arytenoideus transversus, and cricoarytenoideus dorsalis have been removed.)
726 C h ap ter 14. T he R e s p ir a t o r y S ystem
The vocal fold (plica vocalis) (Fig. 14-12), on ing phonation. All three levels may be involved
either side, extends from the vocal process of in normal closure, or only one level.
the arytenoid cartilage to the dorsocaudal part
of the trough of the thyroid cartilage. It is ap
proximately 13 mm. long and 6 mm. wide in a Innervation of the Larynx
medium-sized dog. It is separated from the ves
tibular fold by the slitlike opening of the lateral The nerves of the larynx in the dog (Fig. 10-
ventricle. The vocal ligament (lig. vocale) is a 17) have been reported upon by Franzmann
strap of elastic fibers which is enclosed in the (1907), Lemere (1932a and b, and 1933), Vogel
vocal fold. It forms the supporting framework (1952), and Bowden and Scheuer (1961).
for the cranial border of the vocal fold and is The cranial laryngeal nerve leaves the vagus
covered by mucous membrane. It measures 1 at the level of the nodose ganglion. It divides
to 2 mm. in maximum thickness and has a thin into an external branch, which supplies the m.
cranial border. Caudally, it is continuous with cricothyroideus, and an internal branch, which
the m. vocalis, which is a portion of the thyro- receives fibers from the mucosa of the larynx
arytenoideus muscle mass. cranial to the vocal folds. The internal branch
The ventricle of the larynx (ventriculus laryn usually anastomoses (ramus anastomoticus) with
gis) includes the slight dorsoventral depression the caudal laryngeal nerve. A nerve from the
between the vestibular and vocal folds, as well pharyngeal plexus (pharyngeal ramus of the
as the laryngeal saccule (sacculus laryngis), vagus or medial laryngeal nerve) may also sup
which lies medial to the thyroid cartilage and ply the m. cricothyroideus. The distribution of
lateral to the vocal and vestibular folds. The sensory fibers in the dog and man is similar, ac
mucosa of the laryngeal saccule is continuous cording to Pressman and Kelemen (1955).
with that of the rima glottidis and rima vestibuli The caudal laryngeal nerve is the motor sup
through an opening which maintains a constant ply to all of the intrinsic muscles of the larynx
width of about 1.5 mm. and a length equal to except the m. cricothyroideus. It is the terminal
that of the cranial border of the vocal fold which segment of the recurrent laryngeal nerve, which
forms the caudal border of the opening. In the in turn originates in the thorax by leaving the
production of sound the vocal and vestibular vagus. Rethi (1951) reports that cutting the in
folds can vibrate into the cavity of the laryngeal tracranial portion of the accessory nerve re
saccule as well as into the cavity of the glottis. sults in degeneration of the vast majority of the
Leonard (1957) describes eversion of the lateral fibers in the recurrent nerve. Cutting the intra
ventricles (laryngeal saccules) as one of many cranial portion of the vagus leaves the motor
possible causes of dyspnea in brachycephalic fibers of the recurrent nerve intact, whereas the
breeds. motor component of the cranial laryngeal nerve
The infraglottic cavity (cavum infraglotti- shows complete degeneration. Section of the re
cum) of the larynx extends from the rima glottidis current trunk or of the cranial laryngeal trunk
to the cavity of the trachea. It constitutes a third is followed by degeneration similar to that
subdivision of the larynx. The infraglottic cav which follows intracranial nerve severance.
ity is wide dorsally, corresponding to the lamina Several investigators, including Lemere (1932a)
of the cricoid cartilage, and narrow ventrally. and Pierard (1963), describe an inconstant para-
Because of this disparity in the lengths of the recurrent nerve which leaves the recurrent near
dorsal and the ventral wall of the infraglottic its origin in the thoracic cavity, parallels the re
cavity, the cavity of the larynx is in nearly a current nerve along the trachea, and has var
frontal plane, whereas that of the trachea is in iable anastomoses along its course. It is con
an oblique caudoventral plane. tinuous cranially with the ramus anastomoticus
Pressman and Kelemen (1955), in their excel of the cranial laryngeal nerve.
lent review of laryngeal physiology, consider
the comparative functional anatomy responsible
for the sphincter action of the larynx. Closure TRACHEA
or constriction of the larynx may be accom
plished at three levels: at the inlet (aryepiglottic The trachea runs from a transverse plane
folds) by muscular action during deglutition; at through the middle of the body of the axis to a
the ventricular folds by passive action for cre similar plane through the fibrocartilage between
ating intrathoracic or intra-abdominal pressure; the fourth and fifth thoracic vertebrae. It is a
and at the vocal folds by muscular action dur relatively non-collapsible tube which extends
T ra ch ea 727
P i r i f o r m r e c es s
Se sa mo i d c a r t i l a g e
Aryepiglottic fold
Interarytenoid c a r t i lage
Epiglottis cartilage
Locat i on of l aryngeal s ac c u l
V e s t i b u l a r fold
Basihyoid-
Thy r o hy o i d ligament ■Tracheo
Thyroid c a r t i l a g e '
L a t ventri cl e
Vocal f o l d
E pi gl ot t i s - - Cricoarytenoid lig.
L a r yn g e a l s a c c u l e - Cricoid cart i l age
Cuneiform p r o c e s s of~
aryt enoi d cartilage Vocal process af
arytenoid cart i l age
V e s tib u la r ligament ~
- - Trac he a
Thyroid cartilagei
Lat. ventricle'
Vocal ligj 'Cricothyroid lig.
M. vocalis '/W. cricoarytenoideus lat.
F ig . 1 4 -1 3 . Mid-sagittal section of the larynx. (The mucosa has b een rem oved to expose muscles and ligaments.)
728 Chapter 14. T he R e s p ir a t o r y Sy stem
from the cricoid cartilage of the larynx to its (1960). This process of branching continues un
bifurcation (bifurcatio tracheae) dorsal to the til the respiratory bronchioles are formed. The
cranial part of the base of the heart. The crest bronchi are cylindrical tubes which are kept
of the partition at the site where the trachea patent by flattened, overlapping, curved carti
divides into the two principal bronchi is called lages. The cartilaginous elements end when the
the tracheal carina (carina tracheae). Approxi diameter of the terminal bronchiole is reduced
mately 35 C-shaped hyaline tracheal cartilages to 1 mm. or less. In addition to the cartilages,
(cartilagines tracheales) form the skeleton of the the bronchioles have spiral bands of smooth
trachea. The space left by failure of these car muscle in their walls which continue peripher
tilages to meet dorsally is bridged by fibers of ally on the respiratory (alveolar) bronchioles.
the smooth, transversely running tracheal mus The respiratory bronchioles (bronchioli re-
cle (musculus trachealis) and connective tissue. spiratorii) give rise to alveolar ducts (ductuli al-
The rings so formed are united in a longitudinal veolares), alveolar sacs (sacculi alveolares), and
direction by bands of fibroelastic tissue, called pulmonary alveoli (alveoli pulmonis). The re
the annular ligaments of the trachea (ligg. an- spiratory portion of the bronchial tree in the
nularia trachealia). They are about 1 mm. wide, dog, including the arteries, veins, and lymphat
compared with the 4 mm. width of each tra ics, was modeled and described by Miller (1900,
cheal cartilage. The annular ligaments allow 1937). Boyden and Tompsett (1961), in their
considerable intrinsic movement of the trachea excellent paper on the postnatal growth of the
without breakage or collapse of the tube. Mack- lung in the dog, point out the substantial reduc
lin (1922) reconsidered the network of longi tion postnatally in the number of non-respira-
tudinal elastic fibers in the tunica propria of the tory branches of both the axial bronchus and the
trachea and bronchial tree. This elastic mem peripheral bronchioles. At birth, many non-
brane beneath the epithelium is thick in the respiratory peripheral bronchioles lined by
trachea and larger bronchi but thin in the ter cuboidal epithelium are converted into respira
minal respiratory passageways. It provides a tory units. Concomitantly, new alveoli and al
recoil mechanism for the lung. veolar sacs arise along the terminal bronchioles.
Boyden and Tompsett conclude that the num
ber of non-respiratory branches on the axial
BRONCHIAL TREE
stem (dorsocaudal bronchus of the intermediate
lobe) is reduced from 38 rami before birth to 32
The bronchial tree (Fig. 14-14) begins at the
at maturity.
bifurcation of the trachea by the formation of
Loosli (1937) studied the microscopic struc
the right and left principal bronchi (bronchi
ture of adult alveoli in several species of mam
principales). Each principal bronchus divides
mals, including the dog. He described and illus
into lobar bronchi (bronchi lobares), which are trated interalveolar communications in both
also known as secondary bronchi. These supply
normal and pathologic lungs, and pointed out
the various lobes of the lung, and are named
the significance of these pores in the normal
according to the lobe supplied. Within the lobe
mechanism of respiration and in the spread of
of the lung the lobar bronchi divide into seg
infection.
mental bronchi (bronchi segmentales), which
are sometimes referred to as tertiary bronchi.
THORACIC CAVITY AND PLEURAE
The segmental bronchi and the lung tissue
which they ventilate are known as bronchopul
To obtain a clear understanding of the lungs
monary segments (segmenta bronchopulmo- and how they function passively in the mechan
nalia). Adjacent bronchopulmonary segments ical act of breathing, it is necessary first to un
normally communicate with each other in the derstand the morphology of the thoracic cavity
dog. Various injection and reconstruction tech and its lining membrane, the pleurae.
niques have been employed to delineate these
segments in the dog (Angulo et al. 1958, Tucker
Thoracic Cavity
and Krementz 1957, and Boyden and Tomp-
sett 1961). The thoracic cavity (cavum thoracis) (Fig.
The segmental bronchi usually branch di- 14-15), in a narrow sense, is bounded by the
chotomously (Miller 1937) into small bronchi, subserous endothoracic fascia. In a wider sense,
which have been referred to as subsegmental its walls are formed by the muscles, bones, and
bronchi by Nickel, Schummer, and Seiferle ligaments of the thoracic wall.
T h o r a c ic C a v it y a n d P leu ra e 729
L. c ommon c a r o t i d a
Se c o n d r i b
Es o p h a g u s -
R. a p i c a l l o b e
L . a p i c a l lobe - -
Precava
L su be I a v i o n a
- Trachea
A o r t i c arch
Azygos v
Li gament um art eri osum- -
R. p u l m o n a r y a.
L p u l m o n a r y a.
R c a rd ia c lobe
L b r o n c h us
~-R. bronchus to
apical v cardiac lobes
L. a t r i u m - '
~R. bronchus to
L . p u l m o n a r y V. diaphragmatic lobe
~ R.pulmonary v.
L diaphragmatic
lobe
Postcava
Aortic groove-
I n t e r m e d i a t e lobe
E s o ph a g u s -
R. diaphragmatic
l obe
C e n t r a l tendon
L i ne o f p l e u r a l
re fle c tio n s
Diaphragm
A p ica l lobe
Diaphragm atic lobe
C a r d ia c lobe
Central tendon
Fig. 14-15. Thoracic cage and lungs (lungs hardened in situ). A. Left side. B. Right side.
T h o r a c ic C a v it y a n d Pleurae 731
The endothoracic fascia (fascia endothorac- cess. The recess is formed by the diaphragm
ica) is the areolar tissue which attaches the centrally, and the ribs and intercostal structures
costal and diaphragmatic pleurae to the under peripherally. In normal respiration the dia
lying muscles, ligaments, and bone. The endo phragm undergoes a craniocaudal excursion of
thoracic fascia is scanty where it closely attaches about IV2 vertebral segments, yet even in forced
the costal pleura to the ribs. Dorsally and ven inspiration the margin of the lungs never com
trally it dips into the mediastinal space and be pletely invades the recesses so created. In a
comes the connective tissue that invests the beagle-type dog the diaphragm protrudes for
organs and other structures which lie in the ward from its costal attachment for a maxi
mediastinum. Cranially it passes through the mum distance of 6 cm. For the details of the
thoracic inlet and is continued into the neck. intrinsic structure of the diaphragm see Figures
It blends with the deep cervical fascia, particu 3-33 and 3-34. The thoracic cavity contains the
larly with the prevertebral portion of this fascia. following organs: pleurae, lungs, fibrous and
The thoracic wall is formed bilaterally by the serous pericardium, heart, thymus gland, and
ribs and the intercostal muscles, and dorsally by lymph nodes. The structures which partly or
the bodies of the thoracic vertebrae and the completely traverse the thoracic cavity are the
intervening fibrocartilages. Cranial to the sixth aorta, precava and postcava, azygos and hemi
thoracic vertebra the right and left longus colli azygos veins, thoracic duct and smaller lymph
muscles cover the thoracic vertebral bodies and vessels, esophagus, and vagal, phrenic, and
lie directly under the pleura and endothoracic sympathetic nerves.
fascia. Ventrally, the narrow sternum and the The thoracic inlet (apertura thoracis [crani
paired flat transversus thoracis muscles con alis]) is the roughly oval opening into the
tribute to the thoracic wall. Caudally, the base cranial part of the thoracic cavity. It is bounded
of the thoracic cavity is formed by the dome bilaterally by the first pair of ribs and their
shaped, obliquely placed, musculotendinous cranially extending costal cartilages. In a bea
diaphragm. gle-type dog its dorsoventral dimension is about
The shape of the thoracic cavity of the dog 4 cm. Its greatest width is about one-fourth less
differs greatly from that in man. Its walls are than its dorsoventral dimension. The aperture
laterally compressed, with the result that in is wider dorsally than ventrally. The first tho
dogs of usual proportions its average dorsoven- racic vertebra and the paired longus colli mus
tral dimension is greater than either the average cles bound the thoracic inlet dorsally; the
lateral or craniocaudal measurement. Cranially, manubrium of the sternum bounds it ventrally.
the thoracic cavity opens to the exterior at the Traversing the aperture are the trachea, esopha
thoracic inlet. The external contour of the gus, vagosympathetic nerve trunks, recurrent
thorax differs from the internal limits of the laryngeal nerve, phrenic nerve, first two tho
thoracic cavity. In cross section the thorax is racic nerves, and several vessels. The apices of
roughly oval in shape, wider above than below, the pleural cavities lie in the thoracic inlet.
and long dorsoventrally. A cross section of the
thoracic cavity cranial to the diaphragm is Mediastinum
heart-shaped, with the apex located ventrally.
The base, located dorsally, is widened to accom The mediastinum is the space between the
modate the epaxial muscles, thoracic vertebral right and left pleural sacs which encloses the
bodies, aorta, and smaller associated struc thymus, heart, aorta, trachea, esophagus, the
tures. The thorax is a laterally compressed cone vagus nerves, and other nerves and vessels. It is
with a base (diaphragm) which is convex cra divided by the heart into three transverse divi
nially. The greatest cranial encroachment of sions. The portion of the mediastinum lying in
the diaphragm is to a transverse plane through front of the heart is known as the cranial medi
the sixth intercostal spaces and about 5 cm. astinal cavity (cavum mediastinale craniale).
dorsal to the sternum. In addition to being con The portion containing the heart is called the
vex, the diaphragm is oblique in position; the middle m ediastinal cavity (cavum mediastinale
most cranial dorsal attachment is approximately medium). The caudal mediastinal cavity
eight vertebral segments caudal to its most cra (cavum mediastinale caudale) is that part of the
nial ventral attachment. From an origin on the mediastinum lying caudal to the heart. In man
inner surfaces of the ribs and costal cartilages the mediastinum contains a considerable
the diaphragm bilaterally extends almost di amount of collagenous tissue and is sufficiently
rectly cranially, forming a slitlike space or re strong so that one lung can be collapsed inde
7 32 Chapter 14. The R e s p ira to ry S y s te m
pendently of the other. In the dog the tissue in The costal pleura (pleura costalis) is that por
the mediastinum is extremely scanty, but the tion of the pleura which attaches to the inner
pleura which covers it is not fenestrated. In all surfaces of the lateral walls of the thoracic cav
mammals the mediastinum contains the esopha ity. The true costal part is firmly adherent to the
gus, trachea, heart, and the nerves and vessels inner surfaces of the ribs and is thin. The costal
which enter or leave the heart. Only the post pleura, which covers the inner surfaces of the
cava, certain lymph vessels, and the right intercostal muscles, the aorta, and related struc
phrenic nerve have separate folds as they partly tures, is thicker and less firmly attached to them.
or completely traverse the thoracic cavity. The The pleura and the underlying endothoracic
thymus, trachea, and thoracic duct are located fascia possess considerable elasticity.
primarily in the cranial portion. The mediastinal pleura (pleura mediastinalis)
For purposes of description the mediastinum may be divided into four parts to facilitate
may also be divided into dorsal and ventral por description. These are the ventral, cranial (pre
tions by a frontal plane passing through the cardial), middle (pericardial), and caudal (post
roots of the lungs. In the dog the ventral surface cardial) mediastinal pleurae.
of the heart is attached to the sternum by the The ventral mediastinal pleura (pleura medi
folds of pleura which extend from the thoracic astinalis ventralis) is composed of the ventral
inlet to the diaphragm. The minute space lo portions of the three other parts of the medias
cated between these portions of the pleural tinal pleura. It is much more delicate and
sacs may be called the ventral mediastinal cav expansive than the comparable part in man,
ity (cavum mediastinale ventrale). The cranial which is restricted to the area ventral to the
portion of the ventral mediastinum is occupied heart. In old dogs it extends as a loose mid-ven
by the thymus in growing dogs. It also contains tral fold from the thoracic inlet to the dia
the paired internal thoracic arteries and veins. phragm. Ventrally, it attaches to the sternum.
The pleural reflections which form the ventral
mediastinal pleura continue from the dorsum
Pleurae of the sternum on the transversus thoracis mus
cles to form the right and the left costomedias-
The pleurae are the serous membranes which tinal recess (recessus costomediastinalis). The
cover the lungs, line the walls of the thoracic ventral borders of the various lobes of lungs
cavity, and cover the structures in the medi cranial to the diaphragm lie in these recesses.
astinum, or in places form the mediastinum. The The cranial part of the ventral mediastinal
pleurae form two complete sacs, one on either pleura continues dorsally as the precardial
side, which are known as the pleural cavities. mediastinal pleura. In young dogs it covers the
Each pleural cavity (cavum pleurae) in life is ventral portion of the thymus gland, and in all
essentially only a potential cavity, because it dogs an elevation of the pleura is produced bi
contains only a capillary film of fluid which laterally by the paired internal thoracic vessels
moistens the flat mesothelial cells paving its sur before these vessels disappear ventral to the
face. Except for this capillary fluid, the visceral transversus thoracis muscles. The cranial por
pleura of the lungs, or pulmonary pleura, lies in tion of the ventral mediastinal pleura averages
contact with the wall or parietal pleura. Only slightly over 1 cm. in width in beagle-sized
when gas (air) or fluid collects between the vis dogs. Because of its loose attachments, it may
ceral and parietal pleurae and prevents a lung be increased 2 or 3 times this size without rup
from expanding does it exist as a real cavity. turing. It extends from the third to the sixth
The right pleural cavity is larger than the left sternebra ventrally. Dorsally, the mediastinal
because of displacement of the postcardial pleural sheets separate and enclose the fibrous
mediastinal wall to the left side. The pleural pericardium as the middle mediastinal pleura.
cavities do not communicate with each other, The ventral mediastinal pleura widens caudal
although their medial walls and the tissue be to the heart as it forms a sagittally triangular
tween them are poorly developed. sheet. Ventrally, it attaches to the dorsum of the
For purposes of description the pleura is sternum caudal to the sixth sternal segment.
designated as the parietal and the pulmonary Dorsally, it attaches to the diaphragm.
pleura. The cranial mediastinal pleura (pleura medi
The parietal pleura (pleura parietalis) forms astinalis cranialis) covers most of the thymus
the walls of the pleural cavities. It is further gland, which is large in young dogs. In old dogs
designated as costal, mediastinal, and dia fatty remnants of the gland and vessels persist.
phragmatic. Coursing between the pleural sheets in a caudo-
T h o r a c ic C a v it y and Pleu ra e 733
ventral direction are the right and left internal acutely forward to cover the diaphragm, and
thoracic vessels. Dorsally the pleural leaves sep the caudal leaf continues on the lateral thoracic
arate to cover the precava, brachiocephalic wall as the costal pleura. The diaphragmatic
artery, thoracic duct, and the phrenic, vagal, attachment of the oblique portion irregularly
recurrent laryngeal, cardiosympathetic, and attaches to the diaphragm at an increasing
cardiovagal nerves. The sternal and the cranial dorsocranial level. The line of attachment of
mediastinal lymph nodes also lie in the cranial this portion of the mediastinal pleura follows the
mediastinal space. Above the trachea and left convex portion of the diaphragm parallel to
esophagus the pleural leaves clothe the sides of the muscle bundles which compose its muscular
the longus colli muscles and reflect on the tho periphery.
racic wall as the costal pleurae. The diaphragmatic pleura (pleura diaphrag-
The middle mediastinal pleura (pleura medi- matica) is the pleural covering of the dia
astinalis medialis) leaves the sternum as a deli phragm. It is heavier at the muscular periphery
cate membrane which is a continuation of the of this dome-shaped partition, and more loosely
costal pleura. This portion of the ventral attached, than it is on the tendinous center.
mediastinal pleura may be 4 cm. wide cranially, Where the diaphragm is attached to the lateral
1.5 cm. wide where the heart lies closest to the thoracic wall the diaphragmatic pleura reflects
sternum, and approximately 2 cm. wide cau acutely forward to form the costal pleura. In
dally. The right and left middle mediastinal life a capillary space is present on each side
pleural sheets, upon reaching the fibrous peri between the diaphragm and the caudal lateral
cardium, diverge from each other to enclose the wall of the thorax. These spaces are called the
heart. The middle mediastinal pleura may en right and left costodiaphragmatic recesses (re-
close a portion of the thymus gland when this cessus costodiaphragmatici). After death the
organ is well developed. The right leaf of the diaphragm and its pleura lie in contact with the
middle mediastinal pleura, upon leaving the costal pleura throughout a circular zone which
fibrous pericardium, covers the right phrenic is about 4 cm. in width. The right and left
nerve, precava, trachea, and longus colli mus costodiaphragmatic recesses extend cranioven-
cle before bending ventrally as the costal pleura trally on each side of the mediastinal pleura as
of the right thoracic wall. The left leaf of the the right and left costomediastinal recesses (re-
middle mediastinal pleura, upon leaving the fi cessus costomediastinales). These pleural spaces
brous pericardium dorsally, covers the left are formed between the ventral medias
phrenic and vagal nerves, the left subclavian tinum and the lateral thoracic walls. Although
artery, and the left longus colli muscle before it in cross section they form acute angles, they
reflects on the lateral thoracic wall as the costal are large enough to receive the ventral borders
pleura. The large central portion of the middle of the lobes of the lungs during inspiration.
pleura, after passing over the phrenic nerves, The apical portion of each pleural sac ex
continues dorsally on the pericardium and at tends through the thoracic inlet into the base of
the hilus of the lung of either side the right and the neck, forming a pleural pocket known as the
left pleural sheets reflect on the roots of the re pleural cupula (cupula pleurae). The left cupula
spective lungs and become the pulmonary is the larger and extends further cranial to the
pleurae. first rib than does the right. The right pleural
The caudal mediastinal pleura (pleura medi- cupula is wide dorsoventrally but extends only
astinalis caudalis) lies caudal to a transverse about half as far forward as does the left.
plane passing through the apex of the heart. For The pulmonary pleura (pleura pulmonalis)
descriptive purposes it will be divided into tightly adheres to the surfaces of the lungs and
sagittal and oblique parts. The sagittal part is a follows all of their irregularities. It is the visceral
continuation of the middle mediastinal pleura. portion of the pleura. Its greatest intrapulmo-
It is a triangular fold which bridges the space nary extensions correspond to the adjacent free
between the heart cranially, the sternum ven surfaces of the lobes of the lungs. In all inter
trally, and the diaphragm caudally. It contains lobar fissures, except that between the left ap
the stemopericardiac ligament. The left pleural ical and the left cardiac lobe, the pleura extends
layer of the fold for the postcava is elaborated to the bronchus of the lungs. Between the left
from its right pleural leaf. As a fold from the left apical and the left cardiac lobe the lung paren
pleural leaf the oblique portion runs to the ven chyma does not extend to the bronchus, and
tral part of the left costodiaphragmatic recess. therefore the pleura does not extend as deeply
The cranial pleural sheet of this fold reflects in this fissure as it does in others. In some speci
734 Chapter 14. The R e s p ira to ry S y s te m
mens the borders of the lungs are notched or pericardium to the diaphragm.
fissured by clefts of varying depth. These clefts Histologically, the pleura is more delicate in
occur more often on the apical lobes. the dog than it is in other domestic animals. It
The pulmonary ligament (lig. pulmonale) is a contains smooth muscle fibers, and under the
triangular fold of pleura on each side, which epithelium is a dense network of elastic fibers
leaves the respective lung caudal to the hilus. It which separate the true serosa from the collage
is continuous with the pleura covering the root nous subserosa. The subserosa also contains
or hilus of the lung, and contains no visible elastic fibers, mainly on its deep surface, where
structures. On the left side it extends about 3 they communicate with those of the lobules of
cm. caudodorsally from the large pulmonary the lung (Trautmann and Fiebiger 1957). The
vein leaving the left diaphragmatic lobe, and surface of the pleura is covered by a pavement
ends in a falciform border which stretches from layer of flat, mesothelial cells. In life the pleura
the medial surface of the lung to the left sheet is covered by a capillary film of fluid so that
of mediastinal pleura directly ventral to the the friction between the pulmonary pleura and
aorta. The right pulmonary ligament is about the parietal pleura or between adjacent layers
the same size and shape as the left. It leaves the of the pulmonary pleura is reduced to a mini
acute dorsal border of the right diaphragmatic mum. Elastic fibers are also present in the parie
lobe, extends over the right portion of the inter tal pleura, so that both pulmonary and parietal
mediate or azygos lobe, and becomes the right pleura are capable of stretching. The pulmonary
sheet of mediastinal pleura which covers the pleura is more tightly adherent to the lung
esophagus and aorta. It ends in a falciform bor parenchyma than is the parietal pleura to the
der about 1 cm. cranial to the diaphragm. The thoracic wall.
pulmonary ligaments are relatively avascular.
The pleurae which form them are continuous LUNGS
with the caudal portions of the middle medias
tinal pleurae which reflect on the roots of the The lung (pulmo) is the organ in which oxy
lungs. Small connecting pleural plicae occasion gen from the atmosphere and carbon dioxide
ally extend between adjacent lobes of a lung at from the blood are exchanged. The lungs serve
its hilus. a passive function in the mechanical act of res
The plica venae cavae is a thin, loose fold of piration. The diaphragm and several segmental
pleura which surrounds the postcava. It occu muscles which are attached to the ribs, when
pies the triangular space bounded by the post they contract, bring about an increase in the
cava dorsally, the pericardium cranially, and size of the thoracic cavity, and thus air is drawn
the diaphragm caudally. Because of its delicate into the lungs because of the negative pressure
nature the right portion of the intermediate which is produced. Aiding in expulsion of the
lobe of the right lung is visible through it. The air from the lungs are the abdominal muscles,
right phrenic nerve runs from the base of the which contract and force the abdominal viscera
heart to the diaphragm in a separate plica, only against the caudal surface of the diaphragm.
a few millimeters wide, which leaves the right The two lungs (pulmo sinister et dexter) pos
pleural leaf of the main plica immediately ven sess many features in common. Each has a
tral to the postcava. The plica venae cavae slightly concave base (basis pulmonis), which
leaves the ventral third of the diaphragm about lies adjacent to the diaphragm, and an apex
1 cm. peripheral to its tendinous center. Ven (apex pulmonis), which lies in the thoracic inlet.
trally, it blends with the sagittal portion of the The apex of the left lung is more pointed and
caudal mediastinal pleura. The plica venae extends further forward than the apex of the
cavae on the right and the oblique portion of the right lung (Bourdelle and Bressou 1927). The
caudal mediastinal pleura on the left form a curved lateral surface of each lung is called the
pocket between the heart and diaphragm in costal surface (facies costalis), and the flattened
which is located the intermediate lobe of the surface which faces the opposite lung is called
right lung. The sagittal portion of the postcar- the medial surface (facies medialis). Because
dial mediastinal pleura is located ventral to this the vertebral bodies protrude ventrally from
space where it is formed by the left pleural the dorsal wall of the thorax and intervene be
layer of the plica venae cavae on the right and tween the two lungs this portion of the medial
right pleural layer of the oblique portion of the wall of each lung is known as the vertebral part
caudal mediastinum on the left. Where these (pars vertebralis). The remaining ventral portion
layers come together the pericardiacodiaphrag- of each medial wall faces the mediastinum and
matic ligament runs from near the apex of the is known as the mediastinal part (pars medias-
L ungs 735
tinalis). The medial surface of each lung is (pulmo sinister, lobus apicalis) (Fig. 14-16) is
deeply indented by the heart over an area be transversely compressed between the heart and
tween the third and sixth ribs. This is called the the lateral thoracic wall. Its caudal margin may
cardiac impression (impressio cardiaca). On he medial or lateral to the caudally lying cardiac
each side the pleura covering the pericardial sac lobe. In a 22-pound dog it is 10 cm. long, 3 cm.
is displaced sufficiently by the underlying heart wide, and 1 cm. thick. It extends from the dor
so that its ventral portion lies in contact with sal part of the fifth rib to and through the tho
the costal pleura. The cardiac notch of the right racic inlet, where its apex lies not only cranial
lung (incisura cardiaca pulmonis dextri) is V- to a transverse plane through the first ribs but
shaped, with the apex located dorsally. The also largely to the right of the median plane. It
right cardiac notch is formed by the ventrally is the only lobe in the lung of the dog which
diverging borders of the apical and cardiac regularly fuses with an adjacent lobe. In the left
lobes. Its dorsal apex lies opposite the begin lung the parenchyma of the apical and cardiac
ning of the distal fourth of the fourth rib. On lobes are fused over a transverse distance of 2.5
the left side usually no obvious cardiac notch is cm. from the vertebral border to the fissure
formed. between the apical and cardiac lobes.
Each lung has a diaphragmatic surface (facies The cardiac lobe of the left lung (pulmo sinis
diaphragmatica), which is concave, because it ter, lobus cardiacus) presents a thin dorsocra-
lies against the convex surface of the diaphragm. nially convex border which overlies the caudal
The diaphragmatic surface of the right lung is thickened portion of the apical lobe, or these
about one-third larger than that of the left lung, features and positions of the adjacent lobes are
this larger area being caused by the intermedi reversed. The ventral margin of the cardiac lobe
ate lobe of the right lung extending ventrally of the left lung lies nearly in a frontal plane 1
and to the left, ending in a process at the apex cm. from the mid-ventral line. The left lung
of the heart. does not possess a cardiac notch.
The costal surface of each lung is continuous The diaphragmatic lobe of the left lung
with the medial surface at an acute angle ven (pulmo sinister, lobus diaphragmaticus) is py
trally, lying in the costomediastinal recess. This ramidal in shape and is completely separated
margin, extending from the apex to the base of from the cranially lying cardiac lobe by the
the lung, is called the ventral margin of the lung oblique fissure, which extends from the costal
(margo ventralis pulmonis). Caudally, the ven surface to the root of the lung. When the lungs
tral margin of the lung is continuous with the are moderately distended the fissure begins at
peripheral margin of the base of the lung, or the vertebral end of the sixth rib and ends near
the caudal margin (margo caudalis). the costochondral junction of the seventh rib.
The area of each lung which receives the Right lung. The right lung (Fig. 14-17) is sim
bronchi and furnishes passages for the pul ilar to the left lung in that its parenchyma is
monary and bronchial vessels and nerves is completely divided into cranial and caudal por
known as the hilus of the lung (hilus pulmonis). tions by an oblique fissure, comparable to that
The root of the lung (radix pulmonis) consists of the left lung, but lying about 1 cm. further
of the aggregate of those structures which enter caudad. The right oblique fissure separates the
the organ at the hilus. Each lung is divided cranially lying apical and cardiac lobes from the
down to its root by the oblique fissure (fissura caudally lying intermediate and diaphragmatic
obliqua). The left lung is divided into two lobes.
nearly equal masses by this fissure. The oblique The apical lobe of the right lung (pulmo dex
fissure of the right lung is located about 1 cm. ter, lobus apicalis) extends from the dorsal part
caudal to the oblique fissure of the left lung, but of the oblique fissure cranially and ventrally to
at a comparable angle. Each lung is further the right of the median plane. It does not end
divided by deep fissures into three or four lobes. in a definite apex, as does the left lung, but
The surfaces of adjacent lobes which lie in con rather its most cranial and ventral part has a
tact with each other are called the interlobar gently curved convex border which goes from
surfaces (facies interlobares). an acute, cranial, and dorsal margin to a slightly
convex surface cranial to the heart. This portion
Shape of Lobes and Position of Interlobar of the apical lobe extends across the median
Fissures plane to the left side, whereas its most cranio
ventral portion lies in contact with the caudal
Left lung. The apical lobe o f the left lung portion of the apex of the left lung which ex-
736
Ch.aP te r
H
Tre
sp'f<AT,
°K Y Sy<
'S*E M
T-0,
' a P t> ra n
!Obe 9 ^ Q t,
L ungs 737
Diaphragmatic lobe-
Apical lobe
C ardiac lobe -
P ulm onary a. I n t e r m e d i a te
P u l m o n a r y kk •> lobe
D i ap hr ag ma t i c lobe
A pi c al lobe
C ardiac lobe
B
F ig . 14-17. Right lung. A. Lateral aspect. B. Medial aspect.
738 Chapter 14. T he R e s p ir a t o r y System
tends across the mid line to the right side. The Relations of Lungs to Other Organs
caudoventral margin of the apical lobe of the
The heart produces large impressions on the
right lung is separated from the craniodorsal
medial surfaces of each lung. The cardiac im
portion of the cardiac lobe by the curved hori
zontal fissure (fissura horizontalis). pression of the right lung (impressio cardiaca
pulmonis dextri) is a deep excavation of the
The cardiac lobe of the right lung (pulmo dex
medial or mediastinal surfaces of the right ap
ter, lobus cardiacus) begins at the horizontal
ical lobe cranially, the cardiac lobe laterally,
fissure where its costal surface is broad and
and the intermediate lobe caudally. Ventrally
tapers to a narrow, pyramid-shaped ventral
and on the right the apical and cardiac lobes
extremity which lies caudal or caudosinistral to
fail to cover the heart, so that the pericardial
the apex of the heart. Its medial surface is
pleura lies in contact with the costal pleura
deeply excavated by the heart, resulting in the
over an area which is V-shaped, with the apex
cardiac impression. The cranioventral one-
fourth of its border diverges sharply from the located dorsally. This notch in the right lung is
rounded transversely located caudal portion of called the cardiac notch of the right lung (in
the apical lobe, thus exposing a portion of the cisura cardiaca pulmonis dextri). The cardiac
sternocostal surface of the heart to the thoracic notch exposes about 5 square cm. of the sterno
wall. This notch in the right lung is the cardiac costal surface of the heart to the thoracic wall.
The ventral margin of the left cardiac lobe is
notch (incisura cardiaca).
arciform; it is located, approximately in a
The intermediate lobe of the right lung (pulmo
dexter, lobus intermedius) is the most irregular frontal plane, about 1 cm. from the median
of all of the lobes of the lungs. Caudally it is plane, ventrally. The notch for the postcava (in
molded against the diaphragm; cranially it lies cisura venae cavae caudalis) (see Fig. 4-2) is
in contact with the diaphragmatic surface of the located between the dorsal and the right lateral
heart and the adjacent portions of both dia process of the intermediate lobe. Passing
through this notch in close association with the
phragmatic lobes. The intermediate lobe pos
sesses a thickened middle portion and three postcava is the right phrenic nerve as it runs
processes—a dorsal, a ventral, and a right lat to the diaphragm. Both structures are sur
eral. The dorsal process is a sharp-pointed rounded by the plica venae cavae, which at
taches to the diaphragm. The medial surfaces
pyramid-shaped eminence which extends cau
dally in contact with the caudoventral face of of the apical lobes of the lungs lie in contact
the dorsomedial portion of the diaphragmatic with the thoracic portion of the thymus gland
when the thymus is present. When it is fully de
lobe of the right lung. Its free caudal apex does
not reach as far caudally as do the diaphrag veloped, the thymus gland ends caudally
opposite the left fifth costal cartilage. It there
matic lobes. The ventral process of the inter
fore enters into the formation of the cardiac
mediate lobe runs almost directly ventrally to
impression of the left cardiac lobe.
the dorsal surface of the sixth stemebra. It is
wedged in the space between the diaphragm
Pulmonary Vessels
and the diaphragmatic surface of the heart.
Separating the dorsal and the right lateral lobe The pulmonary arteries carry non-aerated
is the notch through which the postcava and blood from the right ventricle of the heart to the
the right phrenic nerve pass. lungs for gaseous exchange. The pulmonary
The diaphragmatic lobe of the right lung veins return aerated blood from the lungs to the
(pulmo dexter, lobus diaphragmaticus) is similar left atrium of the heart. McLaughlin, Tyler,
in shape and comparable in location to the left and Canada (1961) have shown that the spatial
diaphragmatic lobe, except that it lies to the and functional arrangements of the pulmonary
right of the median plane. It is smaller than the vessels differ greatly between various species.
left diaphragmatic lobe, and-does not extend as In the dog, the pulmonary artery, in addition
far ventrally as does the left diaphragmatic lobe. to supplying the distal portion of the respiratory
Furthermore, its diaphragmatic surface is irreg bronchiole, alveolar duct, and alveoli, continues
ularly excavated in its central part by the inter on to supply the thin pleura.
mediate lobe. Around its periphery it is concave The pulmonary trunk (truncus pulmonalis) is
in all directions, in conformity with the convex the stem artery arising from the fibrous pulmo
surface of the diaphragm against which it lies. nary ring which extends into the media of the
L ungs 739
pulmonary trunk peripherally and centrally vein which immediately receives the vein from
serves for the attachment of muscle fibers from the right cardiac lobe so that blood from all of
the conus arteriosus. The pulmonary trunk bi these areas is returned to the right lateral part
furcates into thte' left and right pulmonary arter of the left atrium by a single large vein 1 cm.
ies, which ramify in the left and right lungs. The in diameter. The blood from the right diaphrag
left pulmonary artery (a. pulmonalis sinistra) matic and intermediate lobes is drained by a
curves dorsally cranial to the vein from the api single trunk which lies to the right of a similar
cal lobe which crosses the main bronchus to this trunk from the left diaphragmatic lobe. The
lobe. Just prior to this crossing a pulmonary pulmonary lobar veins from the left lung
arterial trunk arises and bifurcates. The larger usually open individually into the dorsum of the
terminal branch runs cranially as the main ves left atrium. The pulmonary veins lie most ven
sel to the left apical lobe. It lies dorsal to the trally. The pulmonary arteries circle dorsocau-
apical bronchus, and caudal to the apical vein. dally above the veins, and the lobar bronchi are
The branch to the left cardiac lobe lies cranial insinuated between the arteries and the veins.
to the cardiac bronchus and caudal to the large
vein which drains the cranial part of the left Bronchial Vessels
cardiac lobe.
The small bronchial arteries are variable in
The portion of the pulmonary artery which
origin, although in the majority of dogs the
ramifies in the left diaphragmatic lobe is larger
parent trunk is the bronchoesophageal artery
than all the other branches of the left pulmonary
(see Fig. 4-50), which arises from the right fifth
artery combined. Before entering the lobe it
intercostal artery close to its origin from the
passes dorsal to the bronchus which bifurcates
aorta. The bronchoesophageal artery crosses
to ventilate the left apical and left cardiac lobes.
the left face of the esophagus and contributes
Upon entering the lobe it branches irregularly
an esophageal branch before entering the root
to vascularize the whole lobe. The veins from
of the lung as the bronchial artery. In its course,
all of the lobes compose the most ventral part
the bronchial artery supplies the tracheobron
of the root of the lung.
chial lymph nodes, peribronchial connective
The right pulmonary artery (a. pulmonalis
tissue, and the bronchial mucous membrane. At
dextra) is shorter than the left. It runs caudo-
the level of the respiratory bronchiole the bron
laterally ventral to the left lobar bronchi and
chial artery terminates in a capillary bed which
dorsal to the large left lobar veins. The artery
is continuous with that of the pulmonary artery.
divides unequally into a small branch which
Miller (1937) and, more recently, McLaughlin,
runs to the right apical lobe and a large branch
Tyler, and Canada (1961) were unable to dem
which courses caudally into the right diaphrag
onstrate normally occurring bronchial artery-
matic lobe. Near the origin of the large artery
pulmonary artery anastomoses in the dog.
to the diaphragmatic lobe the relatively small
Notkovitch (1957) found single bronchial
right cardiac lobar artery runs laterally and
arteries on each side in 75 per cent of his speci
enters the dorsal third of the lobe. It is related
mens, and double bronchial arteries on each
to the dorsal surface of its satellite vein and lies
side in 10 per cent. In all cases they arose from
dorsocranial to the right cardiac lobar bronchus.
the first to the fourth right intercostal artery.
It may arise from the right apical lobar artery.
Berry, Brailsford, and Daly (1931) found the
The pulmonary lobar artery to the intermediate
right and left bronchial arteries arose from the
lobe of the right lung enters the thickened mid
right sixth intercostal artery by a common trunk
dle portion of the lobe and trifurcates—a
which also supplied twigs to the esophagus.
branch supplying each of the three processes of
They also described small bronchial vessels
the lobe. This lobar artery lies ventral to the
which supplied the hilus of the lung and which
bronchus to this lobe and dorsal to its satellite
arose from the pericardiacophrenic or internal
vein.
thoracic arteries.
The pulmonary veins (w. pulmonales) (see
True bronchial veins are found only at the
Fig. 4-3) are variable in number. Except for
hilus of the lung. They empty into the azygos
that from a limited area around the hilus, all of
vein or intercostal vein at the level of the sev
the blood distributed to the bronchial tree is re
enth thoracic vertebra.
turned by the pulmonary veins. There is one
main vein from each lobe, although there may
Pulmonary Lymphatics
be two veins which drain the right apical lobe.
The latter veins anastomose to form a larger The afferent lymph vessels from the lobes of
740 Chapter 14. T he R e s p ir a t o r y Sy stem
each lung run to the tracheobronchial lymph anastomoticus and ganglion cells of the superior laryn
nodes of the respective side and to the middle geal nerve. Anat. Rec. 54: 389-407.
--------------- . 1933. Innervation of the larynx; III. Experimen
tracheobronchial lymph node. From these lo tal paralysis of the laryngeal nerves. Arch. Otolaryng.
cations lymph is drained via a chain of cranial 18: 413-424.
mediastinal lymph nodes. Correll and Langston Leonard, H. C. 1956. Surgical relief for stenotic nares in a
(1958) state that a crossing of the lymphatic dog. J. Amer. vet. med. Ass. 128: 530.
vessels draining the lower lobes of the lung oc --------------- . 1957. Eversion of the lateral ventricles of the
larynx in dogs; five cases J. Amer. vet. med. Ass. 131:
curs but is uncommon. Reference is made to 83-84.
Chapter 6 and to Figure 6-20, for further in Loosli, C. G. 1937. Interalveolar communications in normal
formation on the pulmonary lymphatics. and in pathologic mammalian lungs. Arch. Path. 24:
743-776.
BIBLIOGRAPHY Macklin, C. C. 1922. A note on the elastic membrane of the
bronchial tree of mammals with an interpretation of its
functional significance. Anat. Rec. 24: 119-135.
Allison, A. C. 1953. The morphology of the olfactory system McLaughlin, R. F., W. S. Tyler, and R. O. Canada. 1961. A
in the vertebrates. Biol. Rev. 28: 195-244. study of the subgross pulmonary anatomy in various
Angulo, A. W., V. P. Kownacki, and E. C. Hessert, Jr. 1958. mammals. Amer. J. Anat. 108: 149-165.
Additional evidence of collateral ventilation between Miller, W. S. 1900. Das Lungenlappchen, seine Blut- und
adjacent bronchopulmonary segments. Anat. Rec. 130: Lymphgefasse. Arch. Anat. Physiol., Anat. Abtheilung,
207-211. Pp. 197-228.
Berry, J. L., J. F. Brailsford, and I. B. Daly. 1931. The bron --------------- . 1937. The Lung. Springfield, 111., Charles C
chial vascular system in the dog. Proc. roy. Soc. B 109: Thomas.
214-228. Negus, V. E. 1949. The Comparative Anatomy and Physiol
Bourdelle, E., and C. Bressou. 1927. Le cul de sac anterieur ogy of the Larynx. London, W. Hineman Ltd.
de la cavete pleurale chez les carnivores en particulier --------------- . 1958. The Comparative Anatomy and Physiol
chez le chien et chez le chat. Rec. Med. v^t. 103: 457- ogy of the Nose and Paranasal Sinuses. Edinburgh, Liv
466. ingstone.
Bowden, R. E. M., and J. L. Scheuer. 1961. Comparative Nickel, R., A. Schummer, and E. Seiferle. 1960. Lehrbuch
studies of the nerve supply of the larynx in eutherian der Anatomie der Haustiere. Band II: Eingeweide. Ber
mammals. Proc. zool. Soc. London 136: 325-330. lin, Paul Parey.
Boyden, E. A., and D. H. Tompsett. 1961. The postnatal Notkovitch, H. 1957. Anatomy of the bronchial arteries of
growth of the lung in the dog. Acta anat. (Basel) 47: the dog. J. thorac. Surg. 33: 242-253.
185-215. Pierard, J. 1963. Comparative Anatomy of the Carnivore
Correll, N. O., Jr., and H. T. Langston. 1958. Pulmonary Larynx. Thesis, Cornell University.
lymphatic drainage in the dog. Surg. Gynec. Obstet. Pressman, J. L., and G. Keleman. 1955. Physiology of the
107: 284-286. larynx. Physiol. Rev. 35: 506-554.
Duckworth, W. L. H. 1912. On some points in the anatomy Rethi, A. 1951. Histological analysis of the experimentally
of the plica vocalis. J. Anat. Physiol. 47: 80-115. degenerated vagus nerve. Acta morph. Acad. Sci. hung.
Franzmann, A. F. 1907. Beitrage zur vergleichenden Anato Tome I, Fasc. 2: 221-230.
mie und Histologie des Kehlkopfes der Saugetiere mit Trautmann, A., and J. Fiebiger. 1957. Fundamentals of the
besonderer Berucksichtigung der Haussaugetiere. Bonn, Histology of Domestic Animals (translated and revised
C. Georgi. from the 8th and 9th German editions, 1949, by R. E.
Horning, J. G., A. J. McKee, H. E. Keller, and K. K. Smith. Habel and E. Biberstein). Ithaca, N.Y., Comstock Pub
1926. Nose printing your cat and dog patients. Vet. Med. lishing Assoc.
21: 432-435. Tucker, J. L., Jr., and E. T. Krementz. 1957. Anatomical cor
Lemere, F. 1932. Innervation of the larynx. I. Innervation of rosion specimens; I. Heart-lung models prepared from
laryngeal muscles. Amer. J. Anat. 51: 417-437. dogs. Anat. Rec. 127: 655-665; II. Bronchopulmonary
--------------- . 1932a. Innervation of the larynx. I. Innervation anatomy in the dog. ibid. pp. 667-676.
of laryngeal muscles. Amer. J. Anat. 51: 417-437. Vogel, P. H. 1952. The innervation of the larynx of man and
--------------- • 1932b. Innervation of the larynx; II. Ramus the dog. Amer. J. Anat. 90: 427-447.
CHAPTER 15
TH E U R O G E N IT A L SYSTEM
A N D M A M M A RY GLANDS
B y G E O R G E C. C H R IS T E N S E N
The urogenital apparatus or system (appa Because of the close physiological relationship
ratus urogenitalis or systema urogenitale) is of the mammary glands, in the female, with the
made up of the urinary organs and the repro reproductive organs, they too are described in
ductive organs. this chapter.
T H E U R IN A R Y O R G A N S
The urinary organs (organa uropoetica) in an oblique direction, tilted cranioventrally. The
clude the kidneys (renes), the ureters, the blad right kidney is more firmly attached to the dor
der (vesica urinaria), and the urethra (urethra sal wall than is the left, and has a correspond
masculina or urethra feminina). ingly larger retroperitoneal area.
Both kidneys are invested with a fibrous cap
THE KIDNEYS sule, are embedded in adipose tissue (perirenal
fat), and are fixed in position by renal fascia
The kidneys (renes) (Figs. 15-1, 15-2) are (modified subperitoneal connective tissue). The
reddish brown, paired, compound tubular kidneys are not rigidly fixed; they may move
glands that secrete the urine. They are charac during respiration or be displaced by a full
teristically bean-shaped, and each presents two stomach. In some lean animals, it is possible to
extremities, two borders, and two surfaces. The examine the kidneys by deep abdominal palpa
cranial and caudal extremities are joined by a tion. The kidneys (especially the left one) are
convex lateral border and a straight medial bor less firmly attached in the cat than in the dog.
der. The medial border is indented by an oval Upon abdominal palpation, the “floating” kid
opening, the hilus, which opens into a space, neys of the cat may be mistaken for foreign
the renal sinus. The hilus transmits the ureter, bodies in the digestive tract or for fetuses in a
renal artery and vein, lymph vessels, and gravid uterus.
nerves. Of these structures, the renal artery is The kidney in the dog averages 6 to 9 cm. in
the most dorsal, and the renal vein the most length, 4 to 5 cm. in width, and 3 to 4 cm. in
ventral. The nerves and lymphatics lie in close thickness. The weight of the freshly excised
relationship to the vein. kidney averages 25 to 35 gm.
Both kidneys are retroperitoneal, located in
the sublumbar region, one on either side of the Investment and Fixation
aorta and postcava. The dorsal, or parietal, sur
face of each kidney is less convex than the A strong, thin fibrous capsule (capsula fibro
ventral, or visceral, surface. The dorsal surface sa) covers the surface of the kidney. The capsule
is in partial contact with the areolar tissue un dips inward at the hilus to line the walls of the
derlying the sublumbar muscles, whereas the sinus and to form the adventitia of the renal
entire ventral surface is covered by peritoneum. pelvis. It also invests the renal vessels and
The cranial pole of each kidney is covered nerves before they pass into the hilus. The
with peritoneum on both the dorsal and the fibrous capsule of normal kidneys is easily re
ventral surface; only the ventral surface of the movable, except in the renal sinus, where it is
caudal pole is covered. The kidneys thus lie in adherent to blood vessels and to the diverticula
741
742 Chapter 15. T he U r o g e n it a l S y s t e m and M am m ary G la n d s
R, a d r e n a l g l a n d L.adrenal gland
/L.crus of d i a p hr a g m
R. k i d n e y
/ ' P h r e n i c o - a b d o mi n a l a.v v.
/ . ' ' R e n a l a. v v.
Oviductk
. - Ovarian a.rv.
Opening of ova r i a n bu r s a
B r o a d lig. ( m e s o v a r i u m ) —
P r o p e r lig. o f ovary.
L-U terine a . v v.
R. u t e r i n e horn
Aorta
R. u r e t e r
R o u n d lig. of u t e r u s
Deep ci r c u m f l e x i l i a c
Colon U t e r i n e a. ? v.
R. u r e t e r E x t e r n a l i l i a c a.
Re n a l artery
Renal vein
U reteral arterg
U reter
of the renal pelvis. Fat of the adipose capsule passes on into the ureter. Since the kidney of
(capsula adiposa), in which the kidney is par the dog is unipyramidal, there are no calyces
tially embedded, extends through the hilus into connected to the renal pelvis. However, the
the sinus. pelvis of the kidney is elongated in a craniocau-
dal direction, and is curved to conform with
Position and Relations the lateral border of the kidney. It extends into
the renal parenchyma both dorsally and ven
The craniolateral surface of the left kidney is trally by means of curved diverticula. There are
in contact with the dorsal end of the medial generally five or six diverticula curving outward
surface of the spleen, the greater omentum, and and toward the median plane from each border
the greater curvature of the stomach. Cranially, of the pelvis. The cranial and caudal extremities
it is in contact with the pancreas and left adre of the renal pelvis are designated terminal re
nal gland. Dorsally, the kidney, with its adipose cesses, and the funnel-shaped portion of the
capsule, is related to the quadratus lumborum, pelvis is termed the middle recess.
transversus abdominis, and psoas muscles, as
well as to the deep layer of the lumbodorsal fas Structure
cia underlying the retroperitoneal or pararenal
The parenchyma of the kidney is made up of
fat. Caudally, the left kidney of the female is in
an internal medulla (medulla renis) and an ex
contact with the descending colon and the mes-
ternal cortex (cortex renis) (Fig. 15-3). When
ovarium. The peritoneum on the ventral surface
the kidney is cut longitudinally, the medulla ap
of the kidney blends with the peritoneum sus
pears striated. The portion of the medulla clos
pending the ovary. In the male, the renal peri
est to the renal sinus is comparatively light in
toneum is reflected onto the dorsal body wall
color and projects into the renal pelvis. This
as parietal peritoneum. Medially, the left kid
portion, the common renal papilla, is thickest
ney of the male is related to the descending
longitudinally. Transverse, curved secondary
colon, mesocolon, and ascending duodenum.
ridges project from each side of the common
The descending colon is also related to the ven
papilla. These are in contact with the divertic
tral surface of the kidney. The medial edge of
ula of the renal pelvis. The pyramid, from which
the left kidney is located approximately 1 cm.
the common renal papilla projects, is smooth in
from the mid-dorsal line in an average-sized
section and is marked with fine striae that con
dog; the cranial pole, or extremity, lies about
verge from base to apex. These striations are
5 cm. caudal to the upper third of the last rib.
formed by the renal tubules. The common
The right kidney has its cranial pole embed
renal papilla is marked on its intrapelvic sur
ded in the fossa of the caudate lobe of the liver.
face by a variable number of foramina, which
This extremity is located at the level of the 13th
are the openings of the papillary ducts. These
rib. It may be a few centimeters craniad or cau
ducts carry urine into the renal pelvis. The
dad, depending on the degree of gastric or, in
branches of the renal artery and vein diverge
the female, uterine distention. It may be in between the diverticula of the renal pelvis to
contact with the diaphragm and retractor cos reach the cortex.
tae muscle. The right adrenal gland is also re The sectioned peripheral portion of the renal
lated to its cranial pole. Medially, the right parenchyma, the cortex, is granular in appear
kidney is in close proximity to the postcava and, ance owing to the presence of renal corpuscles
ventrally, it is in contact with the right limb of and convoluted tubules. When the kidney is cut
the pancreas and the ascending colon. in a frontal plane, numerous cut ends of arcuate
The renal sinus (sinus renalis) is the cavity of arteries and veins are apparent at the cortico
the kidney. Its opening on the medial border of medullary junction. The thickness of the renal
the kidney is called the renal hilus (hilus re cortex is approximately the same as the trans
nalis). The sinus contains the renal pelvis, a verse diameter of the renal medulla. The
variable amount of adipose tissue, and branches peripheral surface of the cortex is covered by
of the renal artery, vein, lymphatics, and nerves. the fibrous capsule.
After they pass through the sinus, the vessels
and nerves enter the parenchyma of the kidney. Renal Tubules (Tubuli Renales)
The renal pelvis (pelvis renalis) (Fig. 15-3)
is a funnel-shaped, saclike structure that col The nephron (Fig. 15-4), composed of a re
lects urine from the collecting ducts of the kid nal corpuscle (corpusculum renis) and a portion
ney. Urine is mixed in the pelvis before it of straight tubule (tubulus renalis rectus) is the
744 Chapter 15. T he U r o g e n it a l Sy st e m and M am m ary G la n d s
Cor t ex
Medulla
I n t e r l o b a r a.*v.—
Renal capsule
A r c u a t e a.rv.'
Renal pyramid
Medul
Cortex—
------ Re n a l p e l v i s
Di vert i cul um o f p e l vi s -
Pe I vi s
Ureter
E f f e r e n t a r t e r i o l e ------ -
Afferent arteriole—
D istal
convol ut ed tubul e
— C o l l e c t i n g tubul e
I n t e r l o b u l a r a.vv.— H -
Descend!nj l im b
fLoop of H e n l e
unit of urine production. The double-layered of cuboidal cells, but the larger collecting
glomerular capsule (Bowman’s capsule) begins tubules and papillary ducts possess columnar
the renal tubule. It is invaginated by a spherical epithelium.
confluence of blood capillaries, the glomerulus. Glomerular filtration, tubular reabsorption,
The glomerulus and capsule together form the and tubular secretion all probably occur to
renal (malpighian) corpuscle. Renal corpuscles some degree in glomerular vertebrate kidneys
are present in the renal cortex, but not in the (Marshall, 1934). Physiologically, Marshall be
medulla. From each glomerular capsule (cap lieves that the relative importance of the filtra-
sula glomeruli), the proximal convoluted tubule tion-reabsorption mechanism and of the tubular
(coiled and twisted in appearance) descends secretion of any given substance, in a particular
toward the medullary portion of the kidney animal, depends on the substance in question
through the pars convoluta (peripheral region and on the conditions prevailing at the time of
of the cortex) to the pars radiata, where it observation. Shannon (1939) has made an excel
straightens out as the descending limb of Hen- lent survey on the mechanism of renal tubular
le’s loop. This narrow tube extends into the excretion in many mammals, including the dog.
medullary pyramid. After progressing down Detailed studies of renal morphology have also
ward for a variable distance, it makes a U-tum been made by Vimtrup (1928), Oliver (1939),
and again runs peripherally, increases in diam and Smith (1943, 1951).
eter, and becomes the ascending limb of Henle’s
loop. It now reaches the pars convoluta as the
distal convoluted tubule. Vessels and Nerves
After further twisting and coiling, it extends
into the pars radiata, becomes smaller in diam The kidney is a highly vascular organ.
eter, and opens into a straight collecting tubule Briefly, blood enters the renal artery from the
along with other distal convoluted tubules. The aorta, goes through end arteries, interlobar ves
collecting tubule runs through the medulla, sels, arcuate vessels, interlobular arteries, and
united with other collecting tubules, and opens finally to glomeruli via afferent arterioles. Effer
onto the crest of the common renal papilla as a ent arterioles (arteriolae rectae spuriae) leave
papillary duct (ductus papillaris, or duct of Bel the glomeruli and course directly into the outer
lini). layer of the medulla, giving rise to long capil
The epithelium of the renal tubule varies lary nets which extend to the apical end of the
throughout its course. In the adult, the visceral pyramid, or they branch directly into inter-
layer of Bowman’s capsule, the part that covers tubular capillary networks.
the glomerulus, is composed of a single layer of The renal artery (arteria renis) bifurcates
flat cells which dips in between capillary loops. into dorsal and ventral branches. The site of
According to Bensley and Bensley (1930), the bifurcation is extremely variable (Christensen
glomerular epithelium is a continuous layer 1952). Variations in the renal artery are com
over the entire surface of the glomerulus. These mon, ranging from a single vessel to one with
workers describe the glomerular epithelium as numerous branches or to completely doubled
being made up of separate cells and as resting renal arteries. The two primary branches of the
upon an optically structureless basement mem renal artery, end branches, divide into two to
brane which forms the supporting wall of the four interlobar arteries (aa. interlobares renis).
glomerular capillary. Some authors consider These branch into arcuate arteries at the corti-
the continuity of the visceral layer to be imper comedullary junction. The arcuate arteries (aa.
fect. The outer layer of Bowman’s capsule is arcuatae) radiate toward the periphery of the
also made up of flat epithelial cells. The convo cortex, where they redivide into numerous
luted tubules (tubuli renales contorti) are com interlobular arteries (aa. interlobulares). Affer
posed of irregularly cuboidal cells containing ent arterioles (vasa afferentes) leave these to
spherical nuclei. The cells of the convoluted supply the glomeruli and thence the efferent
tubules bear a brush border on their inner sur arterioles. The mean glomerular diameter in a
faces. According to Trautmann and Fiebiger 35-pound dog is 170 /i. According to Rytand
(1952), the functional status of the cells deter (1938), there are 408,100 glomeruli in one
mines their structure. The epithelium flattens canine kidney, with a total glomerular volume
again in the descending limb of Henle’s loop of 1,247 cu. mm. Pease and Baker (1950),
and changes to cuboidal in the ascending limb. working on the rat kidney, found that the extra-
The smaller collecting tubules are also made up glomerular capillary system of the cortex and
T he U reters 747
the venous capillaries of the medulla lack a con (1951) found that the innervation of the renal
tinuous endothelium, blood being contained by blood vessels in the cat is derived chiefly
only the basement membrane. through the renal plexus, that the efferent
Venous drainage of the kidney stems from nerves are solely sympathetic, and that the af
the numerous stellate veins (venulae stellatae) ferent nerves include only spinal nerve com
in the fibrous capsule. These connect with veins ponents. Those authors believe that the wide
of the adipose capsule and empty into inter distribution of afferent nerve fibers associated
lobular (venae interlobulares), arcuate (vv. ar- with the renal vessels indicates that impulses
cuatae), and interlobar (vv. interlobares) and, coming from the kidneys may play an impor
finally, into the main trunk of the renal vein tant part in the reflex regulation of the organs.
(vena renis) which joins the postcava. Venous
arcuate vessels, unlike their arterial counter
Anomalies
parts, unite to form elaborate arches. Arcuate
veins span the medulla to join the dorsal and
Because of development of the kidney from
ventral parts of the kidney.
the nephrogenic blastema and the mesonephric
The reported existence of two independent
duct, malformations are fairly common. Con
circulatory pathways in the kidney, emanating
genital renal cysts result from improper em
from the renal artery (Trueta et al. 1947), was
bryonic union of these structures. Polycystic
not confirmed by Moyer et al. (1950), Schlegel
kidneys may occur in the dog, but are most fre
and Moses (1950), and Christensen (1952).
quently found in cats and hogs. Isolated renal
Daniel et al. (1952) stress the view that blood cysts are more commonly found. Uremia may
passing through either the cortex or the med
result from bilaterally cystic kidneys.
ulla in normal animals passes through glomer
Other congenital anomalies of the kidney in
uli. White (1940) found that, under conditions clude hypoplasia (very small kidney), aplasia
normally encountered in the dog, glomeruli are (failure of one kidney to develop), and hypo
open all of the time. However, the possibility genesia (bilateral rudimentary development).
that severe hemorrhage or sudden increments Fetal lobations may persist in the adult dog
of circulating epinephrine may bring on glo as the result of embryonic retardation. Varia
merular intermittence is not excluded. Moses tions in size, shape, and position are not infre
and Schlegel (1952) found that collateral renal quent. Variations in the arrangement of the
circulation in the rabbit, by virtue of anasto renal arterial supply are frequently observed
moses between arcuate and interlobular arter (Christensen 1952).
ies, is sufficient to maintain renal nutrition under
conditions of stress. Renal vascularization has
also been studied in detail by Huber (1907), Clinical Considerations
MacCallum (1926, 1939), Morison (1926), and
Fitzgerald (1940). When the need for unilateral nephrectomy
Capsular and parenchymal lymphatics are is indicated in the dog, a high paracostal inci
connected to interlobular plexuses which pass sion and a retroperitoneal approach to the
into trunks that leave the kidney at the hilus kidney is one method of choice (Markowitz et
(Trautmann and Fiebiger 1952). They termi al. 1954). Others prefer a mid-ventral abdomi
nate in the lumbar lymph nodes. According to nal approach (Blakely 1952b). Diagnosis of
Peirce (1944), lymphatics in the kidney accom renal anomalies in the dog is aided by the use
pany the interlobular, arcuate, and interlobar of pyelography and urinalysis. Focal and sys
vessels, surrounding them in an irregular net temic infections affecting the kidney are treated
work. The periarterial rete is thicker than the according to the nature of the causative agent.
perivenous network. Cortical and perirenal
lymphatics anastomose. Peirce is of the opinion THE URETERS
that medullary rays and medullary substance
lack lymphatics. The ureters are fibromuscular, slightly flat
Renal nerves, consisting of myelinated and tened tubes that carry urine from the kidneys
unmyelinated fibers derived from sympathetic to the bladder. The diameter of a single ureter
and parasympathetic (vagus) nerves, form dense measures 0.6 to 0.9 cm. when it is distended.
plexuses around the renal blood vessels. They The length of the ureter depends on the size of
also innervate the renal tubules and the mus the animal, averaging between 12 and 16 cm.
culature of the renal pelvis. Christensen et al. in a 35-pound dog. The right ureter is slightly
748 Chapter 15. T he U r o g e n it a l Sy st em and M am m ary G la n d s
longer than the left, because of the more cra structed ureter. Obstruction may be caused by
nial position of the right kidney; it is also longer calculi, tumors, scars, ligatures, or developmen
in the male than in the female. tal anomalies.
The ureter begins at the renal pelvis, which
receives urine from the common renal papilla. Clinical Considerations
Running caudoventrally and mesially toward
the urinary bladder, it is bounded dorsally by When treatment of ureteral obstructions is
the psoas muscles and ventrally by the perito feasible, surgical intervention is usually indi
neum (Fig. 15-1). The ureters lie dorsal to the cated.
internal spermatic vessels in the male and to
the utero-ovarian artery and vein in the female. THE URINARY BLADDER
The right ureter lies in close association with
the postcava and is 1 to 2 cm. lateral to the The urinary bladder (vesica urinaria, urocyst)
aorta. The ureters pass ventral to the deep cir (Fig. 15-5) is a hollow, musculomembranous
cumflex iliac and external iliac arteries and organ that varies in form, size, and position,
veins. In the male, the ureter crosses dorsal to depending on the amount of urine it contains.
the ductus deferens, 2 cm. from the junction of It receives urine from the kidneys, through the
the ductus deferens with the neck of the blad ureters, and stores it until it is disposed of
der. It enters between the two layers of peri through the urethra. The bladder in a 25-pound
toneum forming the lateral ligament of the dog is capable of holding 100 to 120 ml. of urine
bladder and reaches the dorsolateral surface of without being overly distended. When relaxed,
the bladder just caudal to its neck. In the fe the bladder in a 25-pound dog measures 17.5
male, it reaches the lateral ligament of the blad cm. in diameter and 18 cm. in length. When
der after being associated with the broad liga contracted, it measures 2 cm. in diameter and
ment of the uterus. The ureters enter the 3.2 cm. in length. The bladder may arbitrarily
bladder obliquely and open by means of two be divided into a neck (cervix vesicae) connect
slitlike orifices. ing with the urethra, a blunt cranial end (fundus
vesicae), and a body (corpus vesicae).
Structure The ventral surface of the bladder is sepa
rated from the abdominal wall, just cranial to
The muscular wall (tunica muscularis) of the the pubis, by visceral and parietal layers of peri
ureter is divided into three thin layers: an outer toneum. The greater omentum frequently oc
longitudinal, a middle circular, and an inner cupies the space between peritoneal layers.
longitudinal. Only longitudinal fibers are pres Dorsally, the bladder is in contact with the small
ent at the junctions of the ureters with the blad intestine (jejunum and, frequently, ileum) and
der. The ureteral mucosa (tunica mucosa) is with the descending colon (in the female), cra
made up of transitional epithelium. nial to the divergence of the uterine horns from
the body of the uterus. In the male the deferent
Vessels and Nerves ducts lie dorsal to the neck of the bladder,
whereas in the female the cervix and body of
The blood supply to the ureter is derived the uterus are in contact with the dorsal surface
from the renal artery (cranial ureteral artery) of the bladder (Fig. 15-25). When empty, the
and the urogenital artery (caudal ureteral ar bladder lies entirely, or almost entirely, within
tery). Cranial and caudal ureteral arteries anas the pelvic cavity. The space on each side of the
tomose on the ureter. The ureteral arteries have bladder is occupied by the ureters and small
venous counterparts. The autonomic nerves to intestine.
the ureter come from the celiac and pelvic
plexuses. Structure
In the male, the sphincter lies deep to the pros tend cranially to the umbilicus, and each con
tate gland. The mucosa of the urinary bladder, tains an umbilical artery (round ligament of the
like that of the ureter and renal pelvis, is made bladder) and a ureter. Before birth, the bilateral
up of transitional epithelium. It is irregularly umbilical arteries (branches of the internal iliac
folded when the bladder is empty. The mucosal arteries) carry blood from the fetus to the pla
folds disappear during vesical distention. A centa and are components of the umbilical cord.
loose tela submucosa lies between the mucosa When the cord is severed at birth, the arteries
and the muscular layer. retract and become fibrous cords between the
Internally, a triangular area near the neck of bladder and the umbilicus. The narrowed lumen
the bladder is termed the vesical trigone (trigo of each vessel remains patent between the in
num vesicae). The apex of the trigone is at the ternal iliac artery and the bladder, where the
urethral orifice, and the base is indicated by a relatively minute cranial vesical artery leaves
line connecting the ureteral openings. This area the umbilical artery to vascularize the vertex
is free from the characteristic mucosal folds, but and body of the bladder. The lateral umbilical
poorly developed ridges, converging toward the ligaments, in the female, blend laterally with
urethral crest, denote the boundaries of the the broad ligament of the uterus (mesometrium)
trigone. as well as with the lateral pelvic wall. The ureter
and round ligament of the bladder cross at
Fixation nearly right angles to each other at the junction
of the broad and lateral ligaments. The ureter
The reflections of the peritoneum from the is the more mesial of the two structures. In the
lateral and ventral surfaces of the urinary blad male, the ureter and ductus deferens cross each
der to the lateral walls of the pelvis and to the other a few centimeters from the entrance of
ventral abdominal wall are known as ligaments the ureters into the bladder (Fig. 15-5). The
of the bladder. These are made up of double ductus deferens is located in the deferential or
layers of peritoneum separated by intercalated genital fold of peritoneum, which arises from
blood vessels, nerves, lymphatics, and adipose the peritoneal fold containing the internal sper
tissue, as well as by the ureters, deferent ducts, matic vessels and nerves. The peritoneal pocket
and vestiges of embryonic structures. The larg between the rectum and the bladder is termed
est peritoneal fold, the middle umbilical liga the rectovesical pouch. It is partially subdivided
ment (lig. umbilicale medianum), or middle by the genital fold into the rectogenital and
ligament of the bladder, is reflected from the vesicogenital pouches. In the female, the recto-
ventral surface of the bladder to the symphysis genital and the vesicogenital pouch are com
pelvis and the mid-ventral line of the abdominal pletely separate. A small vesicopubic pouch,
wall as far cranially as the umbilicus. It is me between the bladder and pubis, is present in
dian in position and triangular in shape. The the male. In the female, the peritoneum is re
middle umbilical ligament in the fetus contains flected directly from the bladder to the ventral
the urachus (stalk of the embryonic allantois). abdominal wall at the level of the pubic brim,
This normally disappears shortly after birth, so that a vesicopubic pouch normally does not
leaving only the peritoneal fold. Even in the exist.
adult, a vestigial fibrous urachus may sometimes
be found in the free edge of the ligament. In an Vessels and Nerves
average-sized dog the umbilical ligament is wid
est caudally (6 cm.) and narrows down to form The urinary bladder receives its blood sup
an acute angle with the abdominal wall at the ply through the cranial vesical artery, a branch
umbilicus. Caudally, the ligament ends approxi of the umbilical artery, and through the caudal
mately at the level of the vaginovestibular junc vesical artery, a branch of the urogenital artery.
tion in the female, and at the level of a trans The latter is the termination of the visceral
verse plane through the middle of the prostate branch of the internal iliac artery.
gland in the male. The plexus of veins on the urinary bladder
The lateral umbilical ligaments (ligg. umbili- drains primarily into the internal pudendal
cales laterales, chordae arteriae umbilicales), or veins.
lateral ligaments of the bladder, connect the The lymphatics of the bladder drain into the
lateral surfaces of the bladder to the lateral pel internal iliac and lumbar lymph nodes.
vic walls. They are also triangular in shape. The The urinary bladder receives its innervation
lateral ligaments of the bladder in the fetus ex from three sources: somatic, sympathetic, and
Chapter 15. T he U r o g e n it a l S y s t e m and M am m ary G lands
--U reter
r-D u c tu s deferens
-Bladder
U reteral o rific e
( O r if ic e s of
- -Prostate ( du c t u s d e f e r e n t e s
C o l l i c u l u s semi nali s
■ - U r e t h r a I m.
- - U r e t h r a l m.
--U rethra
- Urethra
parasympathetic. The pudendal nerve, derived An enlarged prostate may be the indirect cause
from sacral nerves 1, 2, and 3, supplies its so of dilatation of the bladder. Also, the bladder
matic innervation. It is distributed to the exter may be displaced into the pelvic part of the
nal sphincter of the bladder and to the striated peritoneal cavity, causing a unilateral or bilat
musculature of the urethra. Volitional impulses eral external swelling near the anus (perineal
of urination pass through this nerve. Sympa hernia).
thetic innervation to the bladder is by means of
the hypogastric nerve; parasympathetic im
Clinical Considerations
pulses are carried by the pelvic nerve. Stimula
tion of the pelvic nerves results in contraction
Since the bladder may extend as far cranially
of the bladder and relaxation of the sphincter,
as the umbilicus when it is distended, it is essen
with subsequent urination. McCrea and Kim-
tial that the bladder be catheterized before a
mel (1954) found additional nerves to the blad
dog is subjected to abdominal surgery. Other
der in the human. These arise from the sacral
wise, it is difficult to incise the abdominal wall
nerves and pass to the bladder along the blood
and parietal peritoneum without puncturing
vessels in the endopelvic fascia. Some of the
the bladder.
nerves terminate independently in the pelvic
In case of vesico-umbilical fistula, ligation or
plexus, whereas others join the plexus. It is
cauterization of the patent urachus is indicated,
thought that these nerves maintain normal vesi
provided the urethra is normal and functional.
cal function following radical pelvic surgery.
Torsion of the bladder, although rare, may be
According to Jacobson (1945), section of the
corrected by laparotomy and manipulation.
hypogastric nerves and excision of the lumbar
Calculi in the bladder (Stephenson 1939) may
sympathetic ganglia in the dog result in a tem
possibly be removed with the aid of forceps in
porary decrease in the capacity of the bladder
and a slight increase in its tone. This emphasizes the female, but laparotomy and incision of the
the fact that the principal innervation of the bladder must be resorted to when moderate- to
bladder is by way of the sacral or pelvic nerves large-sized calculi are present. Resection of a
portion of the bladder wall is advocated in the
(parasympathetic).
case of a localized tumor.
Anomalies
THE URETHRA
Anomalies of the urinary bladder include di
verticula, which are also quite common in the Both the male and the female urethra are dis
human (Lower 1914), strictures of the neck of cussed in connection with the genital organs,
the bladder, urachal cysts, and patent urachus. with which they are inseparably associated.
Int. i l i a c a. ( p a r i e t a l b r ) t
I nt. p u d e n d a l a.l Femor al a. ? v.
I n t p u d e n d a l v.,
Lat. c o c c y g e a l a. v v.
A. o f p e n i s ,
C au d al r e c t a l a. r v. x
U r e t h r a I a.vv.
P e r i n e a l a. r v.
Comm. tr. of v of u r e t h r a l
b ul b vdeep v. of p e n i s
A r t e r y of b ul b ■
R i g h t cr us of p e n i s -
P e lvic u reth r a -
Deep a. of p e n i s ■
R. dorsal a. * v. o f p e n i s
B o d y of p e n i s /
S u p e r f i c i a l br. of d o r s a l
\ / ' I
Deep br. of d o r s a l aJ 1
' i
P r e p u t i a l tyn o f d o r s a l a.1 t
R e g i o n of b u l b u s y l a n d i s 1
Reyi on of p a r s l onya y l a n d l s 1
S u p e r f i c i a l vein of yl an s'
Prepuce]
Pre p u c e - p t r ie ta l layer
--------- G l a r s p e n i s
E x t a b d . o b i iqu/e m .—
F e m o r a I a. v v . _____
Ext. p u d e n d a l a. v v.
Pectineus m . --------
A d d u c t o r m . ----
Spermatic cord—
G r a c i I i s m. ~
M e d i a n septurn of s c r o t u m
R. t e s t i s
Epididymis
ered with fine scattered hairs. Sebaceous and which secretes serous fluid into the cavity sepa
tubular (sudoriparous) glands are well devel rating it from the serous membrane covering
oped. Deep to the outer integument of the the testis (tunica vaginalis propria, tunica vagi
scrotum is the dartos, a poorly developed layer nalis visceralis). This potential space, the vagi
of smooth muscle mixed with collagenous and nal cavity (cavum vaginale), is merely the capil
elastic fibers. The dartos forms a common cov lary space between the two serous membranes,
ering for both halves of the scrotum and also filled only with enough fluid to allow free move
helps to form the scrotal septum (Fig. 15-8, C), ment of the testes within the scrotum. A true
but is frequently incomplete in the septum. The cavity does not exist in the unopened scrotum.
separated strands of dartos muscle run obliquely The proper vaginal tunic is also an extension of
craniodorsally from the ventral aspect of the the parietal peritoneum into the scrotum, but
scrotum. Dorsally, the tissue forming the sep this layer closely invests the testis, epididymis,
tum blends with the abdominal fascia. and constituents of the spermatic cord. The
Deep to the dartos tunic and intimately proper and common vaginal tunics are continu
blended with it is the external spermatic fascia. ous with each other along the dorsocaudal
This is an inseparable combination of super border of the cord and testis.
ficial and deep abdominal fascia. The fibrous The above terminology, applied to the vagi
layer of the common vaginal tunic (internal nal tunic, differs slightly from that generally
spermatic fascia) is an extension of the fascia used in human anatomy. However, this nomen
covering the transversus abdominis muscle. This clature follows the precedent set in veterinary
is intimately fused with the external spermatic anatomy by Ellenberger and Baum (1943), Sis
fascia to form a morphologically common fascia son and Grossman (1953), and Miller (1952).
(spermatic fascia). The cremaster muscle of The scrotum, because of its abundant glands
either side inserts upon the fibrous layer of the and thin skin, lack of subcutaneous fat, and its
common vaginal tunic (tunica vaginalis com ability to contract toward or relax away from
munis) dorsomedial to the ipsilateral testis. Each the body, functions as a temperature regulator
muscle arises from the caudal free border of the for the testes (Moore 1942).
internal abdominal oblique muscle and runs
ventrally between internal and external sper Vessels and Nerves
matic fasciae. The cremaster muscles pull the
scrotum and its contents close to the abdominal The principal blood vessel to the scrotum is
wall. Contraction of the dartos muscle causes the external pudendal artery. The cremasteric
the integumentary layer of the scrotum to in artery arises from the femoral or deep femoral
vest the testes more intimately, and thus helps artery. The scrotal arteries run along the cranio-
to bring them closer to the body. In obese dogs, ventral surface of the testis, superficial to the
abdominal fat may totally or partially envelop common vaginal tunic. The perineal branches
the spermatic cord down to the bottom of the of the external pudendal artery supply the scro
scrotum. The fat does not envelop the testis, tum in part. The draining veins follow the same
but continues as fatty cords on the ventrolateral course in reverse.
and dorsolateral surfaces of the fibrous layer of The genital nerve (external spermatic), a
the common vaginal tunic. branch of the genitofemoral nerve (from the
The peritoneal layer of the common vaginal third and fourth lumbar nerves), innervates the
tunic is deep to the fibrous layer. This is an ex skin of the scrotum, the inguinal region, and
tension of the parietal peritoneum into the scro the prepuce. The perineal nerve, a branch of
tum. The common vaginal tunic (tunica vagi the pudendal (from sacral nerves 1, 2, and 3),
nalis communis, tunica vaginalis parietalis) is helps supply this region. The smooth muscula
differentiated from parietal peritoneum at the ture of the scrotum is supplied by the internal
point where the latter dips into the inguinal spermatic plexus of nerves from the pelvic
canal. The peritoneal ring marking this division plexus.
is termed the vaginal ring (Figs. 15-9, 15-10,
15-11). The cavity of the vaginal process is Clinical Considerations
continuous with the peritoneal cavity in domes
tic animals, unlike in the human, where the sac Scrotal and inguinal hernias occur in male
is partially obliterated in development and the dogs, scrotal hernia being the more common of
connection with the peritoneal cavity is lost. the two. A fold of omentum or a loop of intes
The common tunic is an epithelial membrane tine protrudes into the vaginal process and lies
T he T estes 755
in the vaginal cavity (between the proper and buginea joins the mediastinum testis by means
the common vaginal tunic), forcing the testis of connective tissue lamellae (septula testis),
and scrotal wall apart. The hernia may be bi which converge centrally. The mediastinum
lateral or unilateral. Scrotal hernia is character testis is a cord of connective tissue running
ized by an elongated enlargement of the scro lengthwise through the middle of the testis. The
tum and a dilatation of the inguinal canal. lobuli testis (wedge-shaped portions of testicu
Inguinal hernia (intestines or omentum pushing lar parenchyma) are bounded by the septula.
into the inguinal canal peripheral to the vagi The lobuli contain the seminiferous tubules
nal process) is recognizable as a soft, fluctuating (tubuli seminiferi), a large collection of twisted
enlargement to one side of the penis. When re canals. Spermatozoa are formed from the epi
placement by palpation or by holding the ani thelial lining of the tubules, which contains
mal up by the hind legs is unsuccessful, surgical spermatogenic cells and Sertoli cells (supportive
intervention is indicated. cells) that nourish the sperm cells. Straight tu
Wounds and injuries of the scrotum are not bules (tubuli recti) are formed by the union of
infrequent. These are treated locally. If the tes the seminiferous tubules of a lobule. These con
tes are exposed, castration (orchiectomy) is gregate in the rete testis. The mediastinum tes
usually indicated. tis contains a network of confluent spaces and
Tumors of the scrotum, usually fibromas in ducts called the rete testis. Testicular blood
aged dogs, are also treated by castration, in or vessels and lymphatics enter and leave through
der to relieve pressure irritation. the mediastinum. The lobuli testis also contain
A malformed, undeveloped scrotum (infantile interstitial cells (of Leydig), which are thought
scrotum) is the result of undescended testes. to produce testosterone, an internal secretion
The condition may be unilateral or bilateral, de (Pollock 1942, Hooker 1944).
pending on whether one or both testes are re
tained or absent. Attachments
.Tail of e p i d i d y m i s
jSi nus o f e p i d i d y m i s Common v a g i n a l t u n i c
( Pampi ni f o r m Lig. o f ) _
\ p l e x u s o f veins scrotumJ
C r e m a s t e r m. Ski n
Ep i d i d y m i Dartos t u n ic
anastomoses with the testicular artery. The tes as atrophy of the opposite testis, metaplasia of
ticular vein follows the arterial pattern but the epithelium of the prostatic urethra, enlarge
forms an extensive pampiniform plexus in the ment and alopecia of the prepuce, and enlarge
spermatic cord, surrounding the internal sper ment of the prostate gland. Male pseudoher
matic artery, lymphatics, and nerves. The right maphroditism and true hermaphroditism have
testicular vein empties into the postcava at the been reported in the dog (Lee and Allam 1952,
level of the origin of its arterial counterpart. Brodey et al. 1954).
The left drains into the left renal vein. Harrison
(1949) has made a detailed comparative study Clinical Considerations
of the vascularization of the mammalian testis.
The testicular and epididymal lymphatics Injuries, wounds, and tumors of the testes are
anastomose into a variable number of trunks usually treated by surgical castration. Occasion
which drain into the lumbar lymph nodes. ally, in the human, the descent of undescended
The nerve supply to the testis is derived from testes has been hastened by the use of chorionic
the sympathetic division of the autonomic ner gonadotropic hormones. Comparatively little
vous system. According to Kuntz (1919a), the success has been reported with their use in the
testicular (external spermatic) nerves accom dog. They do not restore fertility.
pany the spermatic arteries distally and enter
the gland with either the blood vessels or the THE EPIDIDYM IDES
efferent ducts. Indirectly, they are derived from
the fourth, fifth, and sixth lumbar ganglia of the The epididymides (Fig. 15-8) are the organs
sympathetic trunk. The hypogastric nerve is where spermatozoa are stored before initiation
thought to supply only the proximal end of the of ejaculation. The epididymis is comparatively
ductus deferens. The blood vessels and smooth large in the dog and consists of an elongated
muscle fibers in the testis receive a sympathetic structure composed of a long convoluted tube
nerve supply, but the seminal epithelium and the coils of which are joined by collagenous con
the interstitial secretory tissue do not. Elimina nective tissue. It lies along the dorsolateral
tion of the sympathetic nerve supply to the tes border of the testis. The head (caput) of the
tis is followed by degeneration of the seminal epididymis begins on the medial surface of the
epithelium and hypertrophy of the interstitial testis but immediately twists around the cranial
secretory tissue (Kuntz 1919b). The degenera extremity to attain the lateral side. It is slightly
tive changes are considered to be the result of larger than the remainder of the epididymis. It
paralysis of the blood vessels in the spermatic continues as the body (corpus), which runs
cord and testis. Hypertrophy of the interstitial along the dorsolateral surface of the testis, and
secretory tissue accompanies degeneration of then as the tail (cauda epididymidis), which is
the seminal epithelium. attached to the caudal extremity of the testis by
the ligament of the epididymis. It is continued
Anomalies craniodorsally up the spermatic cord as the duc
tus deferens. The curved epididymis has its con
Cryptorchidism is the most important con cave surface in juxtaposition with the testis. Its
genital anomaly of the testis. This condition is medial edge is attached to the testis by the
comparatively frequent, and is of particular proper vaginal tunic. This dips in between the
economic importance in horses and swine, be lateral edge of the epididymis and the testis,
ing hereditary in swine. In a cryptorchid animal under the body of the epididymis, forming a
one or both testes are retained either in the ab potential space, the sinus epididymis (bursa
dominal cavity (in the region of the inguinal testicularis), which is continuous with the vagi
canal) or between the external (subcutaneous) nal cavity. The sinus is limited cranially and
inguinal ring and the scrotum. Sterile, crypt caudally by the epididymal head and tail, which
orchid dogs usually possess normal sexual adhere tightly to the testis.
desire. More rare than cryptorchidism is anor-
chidism, bilateral absence of the testes, and mon- Structure
orchidism, the presence of only one testis.
Either condition is ascertained only by autopsy In addition to being a storage place for sper
or extensive laparotomy. According to Run- matozoa, the epididymis slowly transports
nells (1954), testicular tumors of dogs have been spermatozoa to the ductus deferens. Circular
reported to cause anatomical alterations, such smooth muscle fibers aid seminiferous tubule
758 Chapter 15. T he U r o g e n it a l S y s t e m and M am m ary G la nds
Peritoneum
'ransversalis fa sc ia
Transversus abdom. m.
Superf. <fdeep a b d o m i n a l
■Sk i n
Vaginal process
Epididymis
-----S c r o t a l l i g
S p e r m a t ic fascia
F ig . 15-10. Diagram of sagittal section of inguinal canal and vaginal process of the male.
76 0 Chapter 15. T he U r o g e n it a l S y s t e m and M am m ary G la n d s
M. s p h i n c t e r ani ext.
\
M. u r e t h r a I i s N
\
\
S ym physis pelvis
M. r e t r a c t o r p e n i
M.bulbocavernosus — ^
Urethra-
Corpus c a v e r n o s u m peni s
E pid idyrnis —
S crotal I ig.
Parietal
peritoneum
Co mmon v a g i n a l t u n i c ^ /
B u i b u s g-l and i s ^ ’ Os peni s
Proper va yin a l tunic/
V i s c e r a l l a y e r of p r e p u c e 1 Pars l o n g a g l a n d i s
P a rie ta l l a y e r of p r e p u c e '
enters the prostate gland before opening into Along the path of the spermatic cord from
the prostatic urethra; the small artery of the the abdominal inguinal ring to the testis, the
ductus deferens, which arises from the urogeni relationships of the vaginal tunics and the en
tal artery; and the vein of the ductus deferens, closed structures remain constant. The tunics
which drains into the internal iliac vein. The also reflect over the testis, joining along the
testicular artery, originating from the ventral dorsomedial border of the organ. A small cir
surface of the aorta, is also in the cord. The left cumscribed area on the tail of the epididymis is
testicular artery arises approximately 4 cm. free of tunic, allowing the scrotal ligament (em
cranial to the origin of the caudal mesenteric bryonic gubernaculum testis) to attach the
artery, and the right arises 4.6 cm. cranial to epididymis to the spermatic fascia. The vaginal
the same artery. The testicular artery runs lat tunics in domestic animals are continuous from
erally and caudally, crossing the ventral surface the abdominal cavity to the scrotum, unlike the
of the ureter, at which point it is joined by the tunics in man, where the connecting portion of
testicular vein and nerve. The peritoneal fold both visceral and parietal layers degenerate at
enclosing the testicular structures is attached to an early age.
the abdominal wall in a line slightly lateral to The spermatic cord passes through the in
the junction of the transversus abdominis and guinal canal in its course through the abdomi
psoas muscles. If the fold of peritoneum is nal wall. The canal is a sagittal slit between the
stretched out, its edge is 4 cm. from the body at abdominal muscles, connected by internal
tachment at the widest point. The left testicular (abdominal) and external (subcutaneous) ingui
vein empties into the left renal vein and the right nal rings (slitlike in appearance). The abdom inal
into the postcava. The plexus of the testicular inguinal ring is located approximately 1 cm.
nerves arises from the area of the sympathetic craniomedial to the femoral ring. The femoral
trunk between the third and the sixth ganglion. ring affords passage for the femoral vessels. The
The testicular lymph vessels pass to the lumbar abdominal ring is bounded medially by the rec
lymph nodes. tus abdominis muscle, cranially by the internal
The components of the spermatic cord (vagi oblique muscle, and both laterally and caudally
nal process) are joined together by loose con by the aponeurosis of the external abdominal
nective tissue and peritoneum, which is desig oblique muscle. The spermatic cord dips in be
nated as the proper vaginal tunic. At the ab tween medial and lateral branches of the caudal
dominal inguinal ring, the parietal peritoneum deep epigastric arteries which lie along the bor
dips into the inguinal canal and scrotum as the ders of the internal ring. The external (subcu
common vaginal tunic (tunica vaginalis parie- taneous) ring, located 2 to 4 cm. lateral to the
talis). The peritoneal ring formed by the dipping linea alba, is merely a slit in the aponeurosis of
of the common tunic into the abdominal inguinal the external abdominal oblique muscle. The
ring is termed the vaginal ring. It encircles the cranial wall of the canal is made up of the
space between the peritoneal cavity of the ab transversus abdominis and internal abdominal
domen and the vaginal cavity within the vaginal oblique muscles, as well as the aponeurosis of
process (spermatic cord). The vaginal ring aver the external abdominal oblique muscle. Only
ages 0.5 to 1 cm. in diameter in the male. There the latter forms the caudal wall of the canal.
is usually an irregular mass of fat at the vaginal As the spermatic cord and testis pass through
ring, covered by peritoneum. It overlaps the the inguinal canal, transversalis fascia (under
cranial border of the ring and probably acts like lying parietal peritoneum) is reflected onto the
a valve to decrease the possibility of intestinal or tunica vaginalis communis, and is here known
omental herniation (scrotal hernia). The fat mass as internal spermatic fascia. The combined
may be in two separate parts. superficial and deep abdominal fascia, from the
The ductus deferens, with its vessels, is en outer surface of the external abdominal oblique
veloped by one fold of peritoneum at the vagi muscle, is reflected onto the spermatic cord as
nal ring, and the testicular vessels and nerves it emerges from the inguinal canal. It then lies
are covered by another. The double layer of superficial to the internal spermatic fascia and
peritoneum uniting these two folds to each is known as the external spermatic fascia. The
other and to the edge of the vaginal ring is cremaster muscle, a caudal fasciculus of the
termed the mesorchium. It may be compared internal abdominal oblique muscle, passes
to the mesentery, which attaches the intestines down between the internal and the external
to the abdominal wall. spermatic fascia.
Scrotal hernia is relatively rare in the dog.
762 Chapter 15. T he U r o g e n it a l S y s t e m and M am m ary G la n d s
Abdominal viscera slip into the vaginal ring, ered on its dorsal surface by muscle fibers from
pass down the vaginal cavity between the parie the wall of the urinary bladder. Capsular tra
tal (common) and visceral (proper) vaginal beculae subdivide the prostate into lobules.
tunics, and reach the scrotum. The condition The columnar glandular epithelium changes to
may be corrected by surgical intervention if it is transitional in the excretory ducts opening into
of fairly recent occurrence. the urethra. It is thought by some that prostatic
secretion renders the spermatozoa actively mo
THE PRO STA TE GLAND bile and neutralizes any acidity in the urethra
resulting from the passage of urine. Huggins
The prostate gland (prostata) (Fig. 15-5) is a (1945) believes that the prostate is essential for
musculoglandular body that completely encom fertilization, because the quantity of spermatic
passes the proximal portion of the male urethra fluid in the ductus deferens is too small to be
and the neck of the bladder. In a mature 25- delivered from the urethra at ejaculation. Ac
pound dog, the prostate is ovoid in shape, but it cording to Huggins, the prostate functions only,
may vary from 1.7 cm. in length, 2.6 cm. in by its secretion, to thin and to increase the vol
transverse diameter, and 0.8 cm. in dorsoven- ume of semen.
tral diameter, to an almost perfect spheroid, 2
cm. in diameter. The average weight of the Vessels and Nerves
prostate for 25-pound dogs, ranging from 2 to
5 years of age, is 6.8 gm. The normal size and The arterial supply of the prostate gland is
weight of the prostate vary, depending on age, derived from the prostatic branch of the uro
breed, and body weight (Zuckerman and Mc- genital artery. The latter arises from the visceral
Keoun 1938). As it encircles the urethra, the branch of the internal iliac. The prostatic artery
prostate is thin dorsally, slightly thicker later ramifies over the surface of the gland, sending
ally, and thickest ventrally. A mid-ventral longi small branches into the parenchyma and, in
tudinal cleft sometimes reaches the urethra. some cases, supplying the prostatic urethra.
The dorsal surface of the prostate is sepa The venous network of the gland drains, ulti
rated from the ventral surface of the rectum by mately, into the internal iliac vein. The pros
two layers of peritoneum which bound the po tatic lymph vessels empty into the iliac lymph
tential rectogenital space. Ventrally, the pros nodes.
tate normally lies on the symphysis pelvis, par The gland receives autonomic innervation
tially separated from it by a double layer of from the pelvic plexus. The hypogastric nerve
peritoneum (Fig. 15-11). Depending on the de (sympathetic) carries impulses which induce
gree of distention of the urinary bladder, the prostatic secretion. According to Huggins
relationships of the prostate vary. If the bladder (1945), the prostate of the dog secretes small
is full, the gland may be in the abdominal amounts of fluid at frequent intervals (resting
region cranial to the pubis, or, if the bladder is secretion) and the secretion is greatly aug
contracted, it may lie 2 to 3 cm. caudal to the mented by parasympathetic stimulants (stimu
brim of the pelvis. lated secretion). During the “resting secretion”
The two deferent ducts enter the craniodor- phase, 0.1 to 2 ml. of prostatic fluid is secreted
sal surface of the prostate. They lie adjacent to per hour.
each other, one on either side of the median
plane. They run caudoventrally through the Anomalies and Variations
dorsal part of the gland to open into the urethra
by two slits, one on each side of the colliculus Prostatic hypertrophy occurs frequently in
seminalis. The prostatic part of the pelvic ure old dogs. Prostate glands enlarged to as much
thra runs slightly off-center through the length as 3 inches in diameter are not rare. The hyper
of the gland. trophy may be glandular, fibrous, or a combina
tion of the two types. Cysts are often present.
Structure The exact pathogenesis is not known, but im
proper hormone balance is generally considered
The prostate consists of numerous compound to be the causative factor. Since the condition
tubular glands enclosed in interstitial connec occurs primarily in dogs that are trained to re
tive tissue containing smooth muscle fibers. The tain urine for long periods of time, Schlotthauer
comparatively thick, fibromuscular capsule of (1937) believes that environment is also a sig
the prostate (capsula prostatae) is partly cov nificant factor.
T he P e n is 763
Clinical Considerations covering it. The body of the penis begins at the
blending of the crura. The corpus cavernosum
A hypertrophied prostate gland may inter penis arises, one on either side, from the ischial
fere with urination or with defecation. Huggins tuberosity, and each runs distally in the dorso
(1945) considers the condition is neoplastic in lateral part of the body of the penis as far as the
nature (cystic hyperplasia). A large number of os penis. In contrast, the corpora cavernosa
aged dogs have this disturbance. Castration penis of the tomcat extend halfway into the
usually causes involution of the gland (Moore glans penis. A fibrous median septum com
1944), and endocrine therapy is often beneficial pletely separates the right and the left caver
(Herman 1940). If the preservation of sexual nous body in the dog. Superficially, each corpus
potency is desired, prostatectomy is performed. cavernosum penis is enveloped by a thick layer
A surgical approach is usually made through of collagenous and elastic fibers, the tunica al
the abdominal wall, and the gland is either buginea. The role of the corpora cavernosa
completely dissected away from the neck of the penis in erection is comparatively limited.
bladder, urethra, and deferent ducts, or only The extremes and averages of the dorsoven
the capsule of the gland is removed or incised. tral diameter of the crus and corpus caverno
Removal or incision of the capsule usually re sum penis of one side (from 150 specimens) are
sults in prostatic atrophy, according to Gadd as follows: of the crus—2.8 to 7.4 mm., average
(1944). 5.2 mm.; of the middle of the corpus caverno
sum penis—0.7 to 2.4 mm., average 1.5 mm.;
THE PENIS of the apex—0.7 to 2.5 mm., average 1.6 mm.
Similar figures for the side-to-side measure
The male copulatory organ, the penis, is ments are: crus—2.8 to 5.7 mm., average 4.2
composed of three principal divisions: the root mm.; of the middle of the corpus cavernosum
(radix penis), the body (corpus penis), and the — 1.8 to 5.4 mm., average 3.4 mm.; apex— 1.5
distal free part (glans penis). The latter is sub to 4.9 mm., average 3.2 mm.
divided into a bulbus glandis and a pars longa The corpus cavernosum urethrae (Figs. 15-
glandis. Unlike the penis of the human, which 12, 15-13) is located in a groove on the urethral
is completely covered by integument and side of the penis and surrounds the penile ure
hangs free from the body except at its proximal thra throughout its course. The bulb of the
end, the root and body of the penis of the dog urethra (bulbus urethrae) is a bilobed expansion
are firmly attached to the ventral body wall and of this erectile body, located between the
are covered by integument only on their ventral crura at the ischial arch. The corpus caverno
and lateral surfaces. Most of the glans penis is sum urethrae narrows in diameter from its ori
enveloped by epithelium and, in non-erection, gin just within the pelvic cavity until it dips
the glans is entirely withdrawn into the prepuce. into the glans penis. There it gives off numerous
The prepuce is attached to the ventral abdomi shunts that supply the bulbus glandis with
nal wall except for its distal open end, which is venous blood. It then continues in the pars
free. The penis has two primary surfaces, a longa glandis to the external urethral orifice.
dorsal (dorsum penis) and a ventral or urethral The cavernous spaces are generally much
surface (facies urethralis). larger than those of the corpus cavernosum
Measurements of the non-erect penis in over penis. From one to four valves are located in
150 mature dogs of assorted breeds show that each of the venous connections between the
its length ranges from 6.5 to 24 cm., with an bulbus glandis and the corpus cavernosum ure
average of 17.9 cm. thrae, preventing blood from leaving the bulbus
The root and body, which are directly con glandis by this route. At the distal quarter of
tinuous, are made up of the corpora cavernosa the glans penis, the corpus cavernosum urethrae
penis, the ventrally located corpus cavernosum diverges ventrally from its groove in the os penis.
urethrae, containing the penile urethra, and The corpus cavernosum urethrae in the non-
the enlarged proximal end of the os penis (bacu- erect penis varies greatly in diameter, depend
lum). The corpora contain enlarged venous ing on the size of the dog. Average dorsoventral
spaces. The root is attached to the tuber ischii, diameters are as follows: urethral bulb— 11.7
and the parts of the root to the left and the right mm.; middle of corpus cavernosum urethrae—
are termed the crura. Each crus is made up of 4.1 mm.; corpus cavernosum urethrae just
the proximal part of the ipsilateral corpus caver proximal to the bulbus glandis—4.6 mm. Aver
nosum penis and the ischiocavernosus muscle age side-to-side diameters are: bulb of urethra
76 4 Chapter 15. T he U r o g e n it a l S y s t e m and M am m ary G la n ds
—unilateral, 8.5 mm., bilateral, 18 mm.; middle horse have no rigidity in the non-erect state. In
of corpus cavernosum urethrae—5.1 mm.; cau the bull and the boar the penis is heavily fibrous,
dal to the bulbus glandis—3.4 mm. with comparatively little erectile tissue. Didier
In contrast to the glans penis of man and the (1946) has made a comprehensive study of the
stallion (Markee 1953, Sisson and Grossman os penis.
1953), the glans of the dog is not a simple cap There are four paired extrinsic penile mus
shaped expansion of the corpus cavernosum cles in the dog (Figs. 15-14, 15-15, 15-16).
urethrae. The glans penis of the dog is a bipar The retractor penis muscles, composed princi
tite structure, consisting of a bulbus glandis and pally of smooth muscle fibers (Fisher 1917),
a pars longa glandis. The distal three-fourths of arise indirectly from the first and second coc
the glans penis is made up primarily of the pars cygeal vertebrae, by fascial slips, and blend
longa glandis. The distal half of the bulbus glan with the anal sphincters and levator ani muscles.
dis is overlapped by the caudal third of the pars Each runs ventrally along the peripheral border
longa glandis. of the anal sphincter to the urethral surface of
The bulbus glandis, a cavernous expansion the penis, and inserts in the penis at the fornix
of the corpus cavernosum urethrae, surrounds of the prepuce. Bands of muscle fibers leave the
the proximal part of the os penis. Its thickest retractor penis at the level of the caudal edge
part and area of greatest potential expansion is of the scrotum and disperse in the septum
located on the dorsal surface of the penile bone. scroti. The short, broad, paired ischiocaverno-
The bulbus glandis contains large venous sinuses sus muscles cover the crura of the penis. Each
bounded by trabeculae rich in elastic tissue. originates from the ischial tuberosity and has a
The pars longa glandis and bulbus glandis are broad insertion upon the corpus cavernosum
separated from each other by a thick layer of penis. The bulbocavemosus muscles, consisting
dense connective tissue. This layer continues mainly of transverse fibers, cover the superficial
distally as a sheath for the corpus cavernosum surface of the urethral bulb. Each arises from
urethrae and the os penis. Although similar to the external anal sphincter and fuses with the
the bulbus glandis in structure, the pars longa retractor penis muscle at the proximal third of
glandis is not capable of comparable expansion. the body of the penis. Each ischiourethralis
The bulbus glandis averages 2.4 cm. in muscle (m. compressor venae dorsalis penis of
length, compared with 5.3 cm. for the pars Houston 1830) originates from the dorsal sur
longa glandis. In diameter, the bulbus glandis face of the ipsilateral ischial tuberosity and in
averages 2.3 cm. dorsoventrally and 2 cm. from serts into a fibrous ring which encircles the
side to side. Comparable diameters for the pars common trunk of the right and left dorsal veins
longa glandis are: dorsoventrally— 1.4 cm. dis of the penis (Fig. 15-14). A strong, short liga
tal to the bulbus and 1.7 cm. in middle; side to ment attaches the fibrous ring to the area adja
side— 1.2 cm. distal to the bulbus and 1.5 cm. cent to the caudal edge of the symphysis pelvis.
in middle. There is also a ligament which attaches the
The os penis (baculum) (Fig. 15-12) is long, fibrous ring to the concave surface of the penis
compared with that of the tomcat. The other at the level of the ischial arch. The two liga
domestic animals do not possess penile bones. ments represent the superficial layer of the uro
Generally considered to be the ossified distal genital diaphragm of man.
portion of the corpus cavernosum penis, the os
penis of the dog tapers in a proximodistal direc Structure
tion. The proximal end, located in the body of
the penis just caudal to the bulbus glandis, is The corpus cavernosum urethrae contains
comparatively broad. It is thicker dorsoventrally numerous sinuses, separated by connective tis
than from side to side. The distal end of the sue trabeculae, in which pass the blood vessels
bone is small in diameter and is extended by a and nerves. Smooth muscle, fibrous connective
slightly curved fibrocartilaginous projection. tissue, and adipose tissue are found in the inter-
The proximal two-thirds of the bone is indented sinusoidal lamellae of the corpus cavernosum
ventrally by a distinct groove. The corpus penis. The glans penis is covered by stratified
cavernosum urethrae and the penile urethra squamous epithelium. The bulbus glandis
occupy the groove until they diverge from the morphologically resembles and is continuous
bone toward the external urethral orifice. The with the corpus cavernosum urethrae. The pars
os penis holds the organ relatively stiff when it longa glandis, also similar to the corpus caverno
is not in erection. The penis of man and of the sum urethrae structurally, has a dense network
T he P e n is 765
of arteries and veins in its trabeculae. Profuse dal artery in the parietal wall of the prepuce.
vascular anastomoses are located under the epi The superficial branch runs ventrodistally deep
thelial surface of the entire glans penis. to the epithelium of the glans penis, extending
almost to the tip of the glans. The deep branch
Vessels and Nerves of the dorsal artery dips into the tunica albu
ginea and reaches the dorsolateral surface of the
The principal source of blood to the penis is os penis, deep to the bulbus glandis (Fig. 15-
the internal pudendal artery, a ramification of 18). It passes into the pars longa glandis, termi
the visceral branch of the internal iliac. This is nating near the penile tip. Distal to the bulbus
augmented by the external pudendal, which glandis, a large anastomotic branch is given off
anastomoses with the preputial branch of the to the corpus cavernosum urethrae. The three
dorsal artery of the penis (Fig. 15-6). After branches of the dorsal artery of the penis and
giving off the urogenital, urethral, and caudal the external pudendal artery supply blood to the
rectal (hemorrhoidal) arteries, the internal pu pars longa glandis.
dendal gives rise to the perineal artery, which The internal and external pudendal veins
supplies the superficial part of the penile root drain blood from the penis. The internal puden
and the perineum (Fig. 15-17). The artery of dal joins the internal iliac vein, and the external
the penis is that portion of the internal puden pudendal drains into the external iliac. The iliac
dal between the perineal artery and the three veins on each side unite to form the common
principal vessels of the penis: artery of the ure iliac vein, the two common iliac veins then
thral bulb, deep artery of the penis, dorsal converging into the postcava.
artery of the penis. Typically, the artery of the The intrinsic penile veins partially parallel
urethral bulb arises from that of the penis proxi the arteries at the root of the penis (Fig. 15-17).
mal to the deep artery of the penis. At this The dorsal veins of the penis are united at the
point, the artery of the penis is continued by the ischial arch for a short distance before they di
dorsal artery (Fig. 15-13). verge into the right and left internal pudendal
The paired arteries of the urethral bulb di veins. Unlike the corresponding arteries, the
verge into two or three branches which divide deep vein of the penis and the vein of the ure
again before entering the corpus cavernosum thral bulb unite in a common trunk before en
urethrae. These supply the spaces and tissue of tering the internal pudendal vein. The perineal
the corpus cavernosum urethrae, the penile vein empties into this common trunk.
urethra, and the pars longa glandis (Christen The dorsal veins of the penis arise from either
sen 1954). The principal trunk is partially coiled side of the bulbus glandis and run along the
in the non-erect state. Its branches anastomose dorsolateral surface of the penile body as far as
with the end branches of the dorsal artery of the ischial arch. The superficial vein of the
the penis as well as with branches of the deep glans runs from the dorsal surface of the pars
artery. longa glandis to the external pudendal vein
The deep artery of the penis gives off two to (Fig. 15-19).
five branches and passes through the tunica al The dorsal vein of the penis (deep dorsal
buginea to enter the corpus cavernosum penis. vein of Deysach, 1939), in its course between
In this cavernous body, the artery again divides the bulbus glandis and the common venous
into clumps of spiral or looped vessels, the heli- trunk, has distinct semilunar valves regularly
cine arteries, which open directly into the spaced along its entire length (Fig. 15-20). The
cavernous spaces. According to Vaerst (1938), vein of the urethral bulb drains the cavernous
Kiss (1921), and von Ebner (1900), helicine spaces of the proximal half of the corpus
arteries retain their spiral shape in the non-erect cavernosum urethrae, arising at the junction of
penis, owing to contracted myoepithelium. its proximal and middle thirds. Typically, two
The dorsal artery of the penis runs diagonally valves are located in the vein of the urethral
distally to the bulbus glandis, anastomosing with bulb between its emergence from the cavernous
the deep artery and artery of the bulb. Proximal body and its junction with the deep vein of the
to the glans penis, the dorsal artery trifurcates penis. The common trunk of the deep vein and
into a preputial branch, a deep branch, and a the vein of the urethral bulb receives the peri
superficial branch. The preputial branch runs neal vein. One to five valves are present be
dorsodistally over the bulbus glandis, supplying tween the origin of the common trunk and its
the dorsal surface of the pars longa glandis as junction with the perineal vein.
well as anastomosing with the external puden The superficial vein of the glans arises from
(Text continued on page 773.)
766 Chapter 15. T he U r o g e n it a l S y s t e m and M am m ary G la n d s
✓ Bul bocavernosus m.
Bulb of urethro
A / Urethra
, Ret ra ct or penis m.
*
/ , Corpus cavernosum urethroe
•
/ i Corpus covernosum penis
i Bul bus g l a n d i s
i
i Visceral l ayer of prepuce
i
i iPars longa glandis
I 1
i Os penis
i i F i broc ort i l ogi nous end
i of os penis
/Septum penis
/
- Ur et hr a ' / Corpus cavern, penis
------- Corpus cavern, urethroe Tunica olbuginea
— Bui b of urethra Urethra
- - Bui bocovernos u s m
V — Corpus cavern urethrae
- Re t r oc t or penis m. ' Ret ract or penis m.
B ------- Os penis
,
■ Crus of corpus cavern penis (Anast of bulbus glandis
* corpus cavern urethrae
Ischiocavernosus m
____ - -Ur e thr o
~~ Corpus cavern urethrae Squamous epithelium
'Bulbocavernosus m
Retractor penis m.
Fic. 15-12. Internal morphology of the penis. Upper drawing, a parasagittal section. A to E, cross sections at five levels indi
cated by letters on upper drawing. (From Christensen 1954.)
T he P e n is 76 7
_ - Deep v. * a. af penis
^ V ir a. of urethral bulb
„ Circumflex br of darsal a.
yDorsal a r v. of penis
, Preputial br of dorsal a
/
I /Superficial v of glans
I Deep v. of glans
i Os penis
C o rp cavern urethrae1 ■
i
Co rp cavern, penis'
Deep br of dorsal a.
Superf i ci al br. af dorsal a
Bulbus glandis
Corpus cavernosum urethrae
Deep br af darsal a
Fig. 15-13. SemidiaRraminatic view of penis. The pars longa glandis and the muscles of the root are illustrated as if transparent.
The vessels of only one side are shown. (From Christensen 1954.)
768 Chapter 15. T he U r o g e n it a l S y s t e m and M am m ary G la n d s
M. b u l b o c a v e r n o s u s M. r e t r a c t o r penis
Cut bul b of ur et hra
M. I s c h i oc a v e r n o s u s
Ur e t h r a
M. i schiourethral i s - _
Fibrous ri ng
T u be r i s c h i i - _
||^J Common trunk of
Sacrotuber ous lig. - - ' dor s al vv. of penis
M. o b t u r a t o r int. In t. pudendal a t v .
M. c o c c y g e u s '
M. l e v a t o r a n i - - . Prostate
F ig . 15-14. Dorsal view of ischiourethral muscles. (The pelvic urethra cut off at the urethral bulb and removed.)
M. rectococcyyeus- -
M. r e t r a c t o r pen is-
M. b u I b o c a v e r n o s u s - -
'•M. o b t u r a t o r int.
M. i schi ocavernosus-
sTuber i schi i
M. i schiourethral is
— M. rectococcycjeus
- M. coccyqeus
- M. l e v a t o r a n i
M . s p h i n c t e r ani
M. o b t u r a t o r int.
"R
M. i sc h i o u r e t hr a l is
M. r e t r a c t o r p e n i s
M. i schi ocavernosus
M. b u l b o c a v e r no s u s
Int. pudendal a. # v.
A. of pen i s - -
Peri neal a v
— Urethra
Anastomosis of deep a
of penis <r a of u bulb. Dorsal a <r v of penis
Deep a of penis
Fic. 15-17. Corrosion preparation of proximal half of the penis. (From Christensen 1954.)
T he P e n is 771
Circumflex branch
R dorsal a *v. of penis
Preputial br of dorsal a
[Deep branch
j ;Superficial branch Corpus cavern, urethrae
| 'Bulbus glandis
i Anastomosis of deep br.
i it a of urethral bulb
Deep br of dorsal a
S u p e r f i c i a l vv of gl an S i ,Fornix of prepuce
I I
Preputi al br
i D e e p vv of glans
Ft. dorsal v r a of penis.
Pars langa glandis ( c u t )
the deep surface of the pars longa glandis, three sacral nerves, enters the sacral plexus, and
which it helps to drain, and runs dorsoproxi- then gives off perineal and rectal (hemorrhoidal)
mally to the fornix of the prepuce, where it branches before continuing as the dorsal nerve
bends acutely and drains into the external pu of the penis. Perineal branches supply the skin
dendal vein by two or more connections (Figs. around the anus and scrotum. Rectal branches
15-6, 15-19). One or more valves are present go to the external anal sphincter, ischiocaverno-
in each branch of the vein. sus, bulbocavernosus, ischiourethralis, ure-
The deep vein of the glans drains blood from thralis, and the retractor penis muscles. The
the pars longa glandis into the bulbus glandis. smooth muscle fibers of the retractor penis also
It arises, on each side of the midline, from the receive sympathetic and parasympathetic im
middle of the deep surface of the pars longa pulses (Oppenheimer 1938). The dorsal nerves
glandis and runs proximally along the dorsolat of the penis pass cranially, on the dorsolateral
eral surface of the os penis to enter the bulbus surface of the organ, to the glans penis (sensory
glandis. A semilunar valve prevents blood in nerves of the glans).
the bulbus glandis from going to the pars longa Lymph vessels from the penis drain into the
glandis. superficial inguinal lymph nodes.
There is a double connection, on each side
of the os penis, between the deep vein of the Mechanism of Erection
glans and the dorsal vein of the penis (Fig. 15-
21). A ventral shunt, through irregularly located Erection in the dog results from the filling of
openings, receives blood from the venous spaces the spaces of the cavernous bodies with an
of the bulbus glandis and the corpus caverno abundance of blood. Stimulation of the nerves
sum urethrae. The openings are directed proxi of erection causes increased penile blood pres
mally and are bordered distally by a lip of endo sure, partial inhibition of venous drainage (Fig.
thelium which diverts blood toward the dorsal 15-20), and dilatation of the arteries in the
vein of the penis. There is also a dorsal shunt penis. The phenomenon of delayed erection in
through the bulbus glandis which is narrower the dog is due to slow engorgement of the bul
in diameter and less clearly defined than the bus glandis and the pars longa glandis.
ventral shunt. Numerous branches go from the Initially, the muscles in the helicine branches
dorsal shunt into the spaces of the bulbus. Blood of the deep arteries and the arteries of the ure
going through the dorsal shunt disseminates into thral bulb relax, allowing more blood to enter
cavernous spaces of the bulbus, whereas blood the cavernous bodies. Arterial blood pressure in
in the ventral shunt is directed into the dorsal the penis rises, but venous outlets are still suffi
vein of the penis without detouring through the cient to accommodate the increased inflow of
venous sinuses of the bulbar erectile tissue. blood. The corpus cavernosum urethrae re
Nerves emanating from the pelvic and sacral ceives the greater share of the blood through the
plexuses supply the penis. These are the paired artery of the urethral bulb. Venous blood con
pudendal nerve and the paired pelvic nerve tinues to flow into the bulbus glandis and into
(n. erigens of Eckhard, 1863). The pelvic plexus the vein of the urethral bulb. Arterial blood is
lies on the pelvic wall dorsal to the prostate shunted from the artery of the urethral bulb
gland, lateral to the rectum. It receives sympa into the corpus cavernosum penis via anasto
thetic fibers through the hypogastric nerve, motic branches. The deep vein of the penis is
which runs caudally from the caudal mesenteric not of sufficient diameter to drain the increased
plexus. The pelvic plexus receives parasympa amount of arterial blood emptying into the cav
thetic fibers through the ventral branches of the ernous spaces from the helicine arteries. Inter
first, second, and sometimes the third sacral nal pressure against the tunica albugiftea causes
nerves. Infrequently only fibers from the second a stiffening of the corpus cavernosum penis.
sacral nerve go to the plexus. The intrinsic veins tend to be compressed.
Fibers leave the pelvic plexus and go to the In the second stage, after intromission has
bladder, prostate, pelvic urethra, rectum, and occurred, partial venous occlusion becomes a
penis. Sensory (afferent) fibers are present in factor in erection of the glans penis. The in
the pelvic nerve, as well as efferent parasympa creased arterial blood supply is largely directed
thetic fibers, according to Gruber (1933). The into the dorsal artery of the penis. Blood in the
hypogastric nerves (sympathetic) are responsi artery of the urethral bulb is shunted to the
ble for ejaculation and prostatic secretion. dorsal artery through circumflex branches. Im
The pudendal nerve (mixed) arises from all pulses in the pudendal nerve stimulate contrac-
77 4 Chapter 15. T he U r o g e n it a l S y s t e m and M am m ary G la n ds
F ig . 15-21. Diagram of venous pathways in the bulbus glandis connecting the deep vein of the glans and the dorsal vein of the
penis. The ventral shunt (B to A) is the principal route whenthe penis is relaxed; the dorsal shunt (C to D) is utilized
during erection. (From Christensen 1954.)
77 6 Chapter 15. T he U r o g e n it a l S y s t e m and M am m ary G la n d s
tion of the extrinsic penile muscles: the ischio- deep veins of the glans and in the venous con
cavemosus, ischiourethralis, and bulbocaverno nections with the corpus cavernosum urethrae
sus. The ischiourethralis contracts enough to prevent blood from leaving the bulbus by any
lessen the free flow of venous blood through route except through the now inefficient dorsal
the common trunk of the dorsal veins. When the penile veins.
paired ischiourethralis muscles contract, the The engorgement of the bulbus glandis and
fibrous ring (encircling the common venous pars longa glandis is facilitated by relaxation of
trunk) is pulled caudally and laterally, narrow the smooth muscles of the intersinusoidal trabec
ing the lumen of the ring and partially squeez ulae and by stretching of the elastic fibers in
ing the vein. The fibrous ring is anchored to the the trabeculae. The elastic tissue covering of the
symphysis pelvis by a strong, short ligament. glans, unlike the thick tunica albuginea of the
Muscular contraction does not affect the arter corpus cavernosum penis, permits maximum
ies, which lie outside the encircling fibrous ring. expansion.
The ischiocavernosus and bulbocavernosus mus The branches of the dorsal artery of the penis
cles, upon contracting, slow down egress of lose their helicine-like appearance as the pars
blood through the deep vein, vein of the ure longa glandis is distended. The deep branch of
thral bulb, and the common trunk of these the dorsal artery uncoils and straightens out
veins. Before intromission, the superficial veins during the height of erection. As the glans penis
of the glans, as well as the dorsal veins of the becomes engorged and firm, less arterial blood
penis, allow free flow of venous blood away can be accommodated, and it is shunted from
from the glans. With constriction of the veins the dorsal artery to the corpus cavernosum
leaving the two cavernous bodies, venous blood penis, increasing its stiffness (Fig. 15-20, B).
is further directed into the bulbus glandis via After ejaculation, the extrinsic penile muscles
shunts from the corpus cavernosum urethrae. relax and arterial blood pressure drops to nor
More arterial blood enters the capillaries of the mal. Venous pressure declines as the erectile
pars longa glandis through the branches of the bodies shrink. The bulbus glandis decreases in
dorsal artery of the penis and the artery of diameter sooner than the pars longa glandis,
the urethral bulb. Constriction of the sphincter owing to its greater venous drainage. Elastic
muscle of the female vestibule causes increased recoil of the intersinusoidal trabeculae helps
occlusion of the dorsal veins of the penis and force blood out of the glans.
also prevents blood from leaving the pars longa For further physiological discussion of the
glandis through the superficial vein of the glans mechanism of erection, reference is made to
(Fig. 15-20, B). Because of their relatively thick articles by the following authors: Eckhard
muscular walls and greater intrinsic pressure, (1863), Francois-Franck (1895), Langley (1896),
the arteries of the penis are not occluded by ex Henderson and Roepke (1933), and Christen
trinsic muscular contraction. sen (1954). Semans and Langworthy (1938)
Since the superficial veins of the glans no have made a notable addition to the knowledge
longer permit outflow of blood, all blood in the of the neurophysiology of sexual function in the
pars longa is directed toward the deep veins of cat. An excellent study of psychosexuality in
the glans. The spaces of the bulbus glandis re dogs has been made by Gantt (1938, 1949).
ceive venous blood from two sources: deep
veins of the glans and the corpus cavernosum
urethrae. When the dorsal veins of the penis Anomalies and Variations
permit free exit of blood from the bulbus, the
blood traverses the ventral shunt in the bulbus Congenitally, the penis may be short or bent.
without expanding the erectile tissue. With the Short retractor penis muscles may inhibit erec
dorsal veins partially occluded, and the inflow tion. Anomalies may be corrected by surgical
of blood greatly increased, blood entering the treatment, but if the condition is congenital it
bulbus through the deep vein of the glans is is not advisable to use the animal for breeding
forced into the dorsal bulbar shunt (Fig. 15-21). purposes.
Because of the abundance of openings in the The flaccid penis varies greatly in length and
dorsal shunt excess venous blood is permitted to diameter, depending on individual variations
enter and engorge the spaces of the bulbus as well as on breed and size of the animal.
glandis. The caliber of the dorsal veins of the Physiological changes caused by temperature,
penis is not sufficient for them to accommodate urination, and sexual excitement also contribute
this increased amount of blood. Valves in the to marked variations of size.
T he M ale U reth ra 777
urethra may infrequently open on the ventral preputii) is a divergent strip of cutaneus trunci
surface of the penis (hypospadias) or on the muscle which extends from the area of the
dorsal surface (epispadias). Hypospadias is con xiphoid cartilage to the dorsal wall of the pre
sidered to result from failure of the urethral puce. There it inserts in apposition with the
groove to close normally, whereas epispadias is preputial muscle of the opposite side. Two func
caused by an embryonic displacement of cells tions are attributed to the preputial muscles—
forming the cloacal membrane, resulting in a to prevent the cranial free end of the prepuce
reversal of the cloacal membrane location. from hanging loosely in non-erection, and to pull
the prepuce back over the glans penis after
Clinical Considerations erection. During erection, the preputial muscles
are relaxed.
Catheterization of the urinary bladder is per
formed through the urethra, and careful sterile Vessels and Nerves
technique is necessary to prevent inflammation
or infection of the bladder and urethra. Hypo The parietal layer of the prepuce is supplied
spadias and epispadias are usually treated by by an intricate, anastomosing network of ar
surgical removal of all of the penis distal to the teries located between the outer and the middle
actual external urethral opening. Occasionally layer of the prepuce. The arteries are branches
the functional urethral opening is undisturbed of the external pudendal artery and the prepu
and the open part of the penis is closed. tial branch of the dorsal artery of the penis (Fig.
15-6). The inner, visceral layer of the prepuce,
THE PREPUCE which invests the glans penis, is supplied by the
three anastomosing branches of the dorsal ar
The prepuce (preputium) (Figs. 15-6, 15-11) tery of the penis, the external pudendal artery,
is a complete tubular sheath of integument and to a lesser degree by the artery of the ure
which, when the penis is not erect, contains and thral bulb.
covers the pars longa glandis and part of the The veins from the parietal layer drain into
bulbus glandis. It is firmly attached to and con the external pudendal vein. The visceral layer
tinuous with the skin of the ventral abdominal is drained by the superficial vein of the glans
wall. into the external pudendal vein and, deeply, by
the dorsal vein of the penis via the deep vein
Structure of the glans and the bulbus glandis.
Preputial lymph vessels go to the superficial
The prepuce is composed of two layers of in inguinal lymph nodes.
tegument dorsally, except for the cranial 1 to Sensory (afferent) innervation of the visceral
3 cm., where it is free of the abdominal wall and layer of the prepuce, covering the glans penis,
there are three layers. Ventrally and laterally is through the dorsal nerve of the penis to the
there are three layers of integument. The outer pudendal. Superficial branches of the ilioingui
layer is skin. The inner layers, parietal and vis nal and iliohypogastric nerves innervate the
ceral, are made up of stratified squamous epi parietal layer.
thelium which is smooth and thin, and stippled
with lymph nodules and nodes. These are more Anomalies and Variations
numerous on the parietal layer and along the
fornix of the preputial cavity than on the vis The two most common anomalies of the pre
ceral layer. The parietal, or middle, layer is a puce are phimosis and paraphimosis. Phimosis
continuation of the outer skin layer onto the is the existence of an abnormally small preputial
wall of the preputial cavity. It extends to the opening, which prevents protrusion of the
fornix, which is located in a transverse plane penis. Paraphimosis, the inversion of the pre
through the middle of the bulbus glandis. In putial opening after penile protrusion, prevents
erection, the parietal layer is reflected from the the return of the glans penis into the preputial
preputial orifice directly onto the proximal half cavity.
of the bulbus glandis and the body of the penis.
The visceral, or inner, layer extends from the Clinical Considerations
preputial fornix to the external urethral orifice,
where it is continuous internally with the caver Phimosis is surgically corrected by incising
nous urethra. Each preputial muscle (protractor the dorsal, free surface of the prepuce suffi
T he B road L ig a m e n t s 779
ciently to allow protrusion of the penis. The 23). The peritoneal extension of the broad liga
outer (skin) and middle (parietal) layers of the ment through the inguinal canal into the sub
prepuce are sutured together. If local manipu cutaneous region of the vulva, known as the
lation, anesthesia, and vasoconstrictor drugs vaginal process, corresponds to the peritoneal
fail to correct paraphimosis, the prepuce is slit outpocketing (vaginal tunics) into the scrotum
on the abdominal side enough to allow retrac of the male. The vaginal process of the female
tion of the penis. Balanitis (inflammation of the varies from none at all, when the peritoneum
visceral and parietal layers) is treated locally, does not even dip into the internal inguinal ring
and venereal granuloma (a tumor of viral origin) (no vaginal ring), to a long process of perito
is corrected by excision and cauterization of the neum containing the round ligament of the
site, followed by six months’ rest from breeding. uterus and fat. It extends into the subcutaneous
tissue of the labia. The round ligament of the
uterus and the ovarian ligaments contained in
FEM A LE GEN ITAL ORGANS the broad ligament are described more fully in
the discussions of the utems and ovaries.
The female genital organs consist of ovaries, Morphologically, the broad ligament is di
oviducts, uterus, vagina, and vulva. The ovaries vided into three regions: mesovarium, mesosal
produce the female sex cells (ova), which are pinx, and mesometrium. The mesovarium is that
transported by the oviducts (fallopian tubes) to part of the broad ligament which attaches the
the uterus. There implantation takes place, if ovary to the dorsolateral region of the abdomi
fertilization has occurred, and the embryo is de nal wall. The cranial boundary of the broad
veloped. The vagina is a canal leading from the ligament, where the suspensory ligament of the
uterus to the external genitalia, the vulva. ovary attaches to or near to the thirteenth rib,
marks its beginning, and it ends at a transverse
THE BROAD LIGAMENTS plane just caudal to the ovary. It contains the
utero-ovarian vessels (Figs. 4-72, 15-22).
The ovaries, oviducts, and uterus are attached The mesosalpinx, another double fold of peri
to the dorsolateral walls of the abdominal cavity toneum, extends laterally from the dorsal peri
and to the lateral walls of the pelvic cavity by toneal layer of the mesovarium. It curves around
paired double folds of peritoneum called the the dorsal and ventrolateral borders of the ovary
right and left broad ligaments (plicae latae uteri) to attach to the medial surface of the broad lig
(Figs. 15-1, 15-22). Each broad ligament con ament just dorsal to the ovary. It encloses the
tains an ovary, oviduct, and uterine hom. It also ovary within a small peritoneal cavity, the ova
contains vessels and nerves to the genitalia and rian bursa. The ovarian bursa is variable in size,
finger-like streaks of fat. It does not support or depending on the age and size of the animal. It
suspend the genitalia in the body cavities, but averages 2 cm. in length in a 25-pound dog. The
rather unites its components. amount of fat between the peritoneal layers of
The broad ligament is attached dorsally along the mesosalpinx depends on the condition of the
or near the junction of the psoas and transver animal. The bursa is completely closed except
sus abdominis muscles. Cranially, it is attached for a narrow slit in its ventromedial surface, con
by means of the suspensory ligament of the necting it with the peritoneal cavity (Figs. 15-
ovary to the junction of the middle and distal 22, 15-24). In a 25-pound dog, the opening
thirds of the last rib. The ligament is reflected averages 0.8 cm. in length. Topographically, it
off the vagina onto the rectum dorsally, ven is located caudoventral to the kidney. Several
trally onto the urethra and bladder, and in a structures come close together at the opening of
curved line laterally onto the wall of the pelvic the bursa. These are the proper and suspensory
cavity as far as the internal inguinal ring. The ligaments of the ovary, and the fimbriae arising
ligament is broadest at the level of the ovary, from the ovarian end of the oviduct. The entire
tapering from there to its cranial and caudal ex oviduct lies between the peritoneal layers of the
tremities. In a 25-pound dog, the distance mesosalpinx.
spanned by the broad ligament at the ovarian The mesometrium begins at the cranial edge
level varies from 6 to 9 cm. A peritoneal fold of the uterine horn, where it is continuous with
arises from the lateral surface of the broad liga the mesovarium, and extends caudally to a point
ment and extends from the ovary to or through where the peritoneum of the broad ligament re
the inguinal canal. It contains the round liga flects onto the bladder and the colon. It leaves
ment of the uterus in its free border (Fig. 15- the uterine horn, the body of the utems, the
780 Chapter 15. T he U r o g e n it a l S y s t e m and M a m m a r y G la n d s
cervix, and the cranial part of the vagina to at nial portion of the free border of the broad liga
tach along the abdominal and pelvic walls. The ment. It is continued caudally by the proper
cranial and caudal uterine arteries and veins ligament of the ovary (ovarian ligament, chorda
run between the peritoneal layers of the meso- utero-ovarica). This in turn attaches to the cra
metrium. nial end of the uterine horn. There it is continu
ous with the round ligament of the uterus,
THE OVARIES which extends caudally toward the vaginal
process. Both the proper and the suspensory
The ovaries (ovaria), or female gonads, are ovarian ligament are composed of connective
paired oval organs, located in the abdominal tissue mixed with smooth muscle fibers.
cavity caudal to the kidneys. They are the sites
of ova formation and development, as well as
the source of certain hormones. In a 25-pound Structure
dog, an ovary averages 1.5 cm. in length, 0.7
cm. in width, 0.5 cm. in thickness, and 0.3 gm. The ovary is divided into a medulla and a cor
in weight. In its normal position, an ovary may tex. The medulla (zona vasculosa) contains
be described as having cranial and caudal poles, blood vessels, nerves, lymphatics, smooth mus
medial and lateral borders, and dorsal and ven cle fibers, and connective tissue fibers. The cor
tral surfaces. The ovary is smooth in appear tex consists of a connective tissue stroma which
ance before estrus, which occurs for the first contains a large number of germ cells, follicle
time between 6 and 9 months of age. In multip- cells, and graafian follicles (folliculi oophorive-
arous bitches its surface is rough and nodular. siculosi). In the absence of germ cells, the de
In a sexually mature, 25-pound dog, the left velopment of interstitial cells appears to be
ovary is located approximately 12 cm. caudal correlated with the activity of the follicle or in
to the middle of the 13th rib and 1 to 3 cm. different cells (Kingsbury 1914). The interstitial
caudal to the corresponding kidney. Typically, cells are modified stroma cells of connective
it lies between the abdominal wall and the left tissue origin. The connective tissue condenses
colon. The right ovary is located approximately to form the tunica albuginea on the periphery
10 cm. caudal to the last rib of the right side. of the ovary. The tunica albuginea is covered
The ventral border and medial surface of the by germinal epithelium made up of a single
ovary are in contact with the mesovarium, layer of cuboidal or columnar cells which are
whereas the dorsal surface and lateral border, flattened in the adult (Trautmann and Fiebiger
free of an intimate peritoneal covering, face the 1952). The surface of the ovary facing the ovar
mesosalpinx across the ovarian bursa. In a ian bursa is covered by germinal epithelium
young animal it lies ventral to the adipose cap which is free of serosa. From the germinal epi
sule of the right kidney and dorsal to the de thelium, rows of germ cells grow inward during
scending duodenum. In animals that have un fetal life and produce follicles (vesicle-like struc
dergone numerous pregnancies, both right and tures) in which the ova are developed. Accord
left ovaries migrate caudally and ventrally. The ing to some workers, new ova are formed from
left ovary, however, always remains in a trans the germinal epithelium at each estrus. Huge
verse plane caudal to the right ovary, corre numbers of follicles degenerate and become
sponding to the relative positions of the kidneys. atretic. A graafian follicle is composed of a num
ber of cell layers. It envelops a cavity filled with
Ligaments follicular fluid. At one end of the cavity there is a
germ-hill (cumulus oophorus), which contains
In addition to the mesovarium (cranial por the maturing ovum. In intimate contact with
tion of the broad ligament), the ovary has two the ovum is a clear membrane, the zona pellu-
other ligamentous attachments. The suspensory cida. This is surrounded by a layer of radially
ligament of the ovary (plica suspensoria ovarii) arranged cells, the corona radiata. As the fol
is attached cranially to the middle and ventral licular fluid (liquor folliculi) increases, the fol
thirds of the last one or two ribs. Caudally, it licle migrates to the periphery of the ovary.
attaches to the ventral aspect of the ovary and When the follicle is under considerable tension
mesosalpinx, lying between the opening of the by pressure of the fluid, it ruptures, and the
ovarian bursa and the ascending oviduct (Fig. ovum is released into the ovarian bursa, which
15-24). The suspensory ligament lies between is actually an outpocketing of the peritoneal
the two layers of peritoneum and forms the cra cavity (Fig. 15-24). Strassmann (1941) de-
T he O v a r ie s 781
Wing o f s a c r u m^ |Ext. i l i a c a . r v .
V i s c e r a l b r o f int. i l i a c a.tv. IR o u n d l i g . of u t e r u s ( c u t )
P a r i e t a l b r of int. i l i a c a.tv., I U t e r i n e a. •? v
U rog en i f a I a. t v.i \R. u t e r i n e h o r n
Caudal vesi ca I a. t v. \Opemng of ovari an bursa
Cr an i a l g l u t e a l a. t v.
Superf. l a t co c c y g e a l a.tv.f |.Suspensory l/g.
Caudal g l u t e a l a.t
I n t. j pudejida b^Tty.
Caudal rectal a. t
M. c o n s tric to r
vu lv a e
1Postcava
R. u r e t e r
Colon O v a r i a n a. t v.
i
^ Aorta
M. co n stricto r Mesomef r i um
v e s t ib u l i i 1 j
Il 1 I 1 j , U t e r i n e a.rv.
P e r i n e a l a. t v .1 j j j / I
-Ovary
Oviduct
M. sphincter ani e x t -
‘ — —Me s o m e t r i um
C e rv i x -------------
- - U t e r i n e horn
M. constrictor vestibuli —
Round l ig. of u t e r us
Cl i t o r i s- " Bladder
S ym physis p e l vi H.H.
V a g in a l process> 1P a r i e t a l p e r i t o n e u m
scribes a “theCa cone” in the ovary of the dog, Vessels and Nerves
human, and other mammals. It is formed by ec
centric growth of the follicle and the one-sided The ovary is supplied with blood through the
proliferation of the theca interna toward the ovarian artery. Homologous to the testicular
ovarian surface. It has the special function of artery of the male, the ovarian artery arises from
bringing the follicle up to that part of the ovar the aorta approximately one-third to one-half
ian surface from which the ovum can escape the distance from the renal arteries to the deep
to the peritoneal cavity. circumflex iliac arteries. Usually the right ovar
After ovulation, relatively slight hemorrhage ian artery arises slightly cranial to the left (Fig.
occurs, filling the follicular cavity. As this is re 15-1). The degree of uterine development de
sorbed, the corpus luteum (yellow body) is termines the tortuosity and size as well as the
quickly formed. It contains yellow lipid ma position of the artery. In a nulliparous animal,
terial. If fertilization does not take place, the the artery extends laterally almost at right
corpus luteum gradually degenerates into a angles from the aorta, whereas in late preg
connective tissue scar, the corpus albicans. If nancy it is drawn cranioventrally, along with
the ovum is fertilized, the corpus luteum re the ovary, by the enlarged, heavy uterus. In ad
mains fully developed throughout pregnancy. dition to supplying the ovary, the ovarian artery
After parturition, it regresses. Involution of the supplies branches to the adipose and fibrous
corpus luteum again allows graafian follicles to capsules of the kidney. In addition, small tor
mature. Estrus in the bitch usually occurs twice tuous branches supply the oviduct and uterus.
a year, in the spring and again in the fall. The Caudally, the artery anastomoses with the uter
reproductive cycle averages 172 to 200 days ine artery (a branch of the urogenital artery).
(Leonard 1960). However, some bitches come Through this anastomotic connection, the uter
into heat three times a year. ine artery may be considered as a supplemen
The cyclic changes of the ovary are initiated tary source of arterial blood to the ovary. The
by two gonadotrophic hormones originating in arteries to the ovary supply the parenchyma of
the anterior lobe of the pituitary gland. The the medulla and cortex as well as the thecae of
follicle-stimulating hormone (FSH) causes ma the follicles. Capillary loops become extensive
turation of graafian follicles. The luteinizing during follicular enlargement but recede or dis
hormone (LH) converts follicular cells into appear during corpus luteum regression.
lutein cells of the corpus luteum. Follicular and The right and left ovarian veins have differ
lutein cells then secrete hormones which act on ent terminations. The right vein drains into the
the uterus and, reciprocally, on the pituitary postcava, whereas the left enters the left renal
gland. Neither FSH nor LH, acting alone, can vein. Similar to the corresponding arteries, the
cause ovulation (Hisaw 1947). Preovulatory en uterine and ovarian veins anastomose between
largement and rupture of the follicle is produced the peritoneal layers of the broad ligament. The
by the combined action of both hormones. Ovu ovarian vein receives a tributary which comes
lation is initiated by an increase in the secretion from the medial edge of the suspensory liga
of gonadotrophins by the pituitary gland and, ment of the ovary and the lateral surface of the
according to Hisaw, luteinizing hormone is the kidney. In some instances, the vein will also
principal one concerned. anastomose with the deep circumflex iliac vein.
Raps (1948) made a study of the develop The lymphatics drain into the lumbar lymph
mental changes in the dog ovary, from two days nodes. Polano (1903) has demonstrated the ovar
after birth to the sixth postnatal month. His ian lymphatics of the dog.
data show that primordial ovocytes surrounded The nerve supply to the ovaries is via the
by follicular epithelial cells are present at 4 days sympathetic division of the autonomic nervous
of age; that at 15 days true primary follicle for system. The nerves reach the ovaries by way of
mation with granulosa cell development occurs; the renal and aortic plexuses, which receive
and that antrum formation is not observable nerve fibers from the fourth, fifth, and sixth
until 6 months of age. Raps also found that ger lumbar ganglia of the sympathetic trunk (via
minal epithelial activity occurs in cycles and is the caudal mesenteric plexus). They accompany
not a continuous process; that cortical activity the ovarian artery to the ovary. The ovarian
is greatest at about 13 days of age and is in blood vessels receive an abundant sympathetic
versely correlated with medullary activity; and nerve supply, but, according to Kuntz (1919a),
that the tunica albuginea reaches its greatest the ovarian follicles and interstitial secretory tis
peak, developmentally, in early life. sue are devoid of sympathetic innervation.
784 C h ap ter 15. T he U r o g e n it a l System and M am m ary G lan ds
Descendi ng o v i d u c t s
\ I n f undi bul u m
M e s o s a I p i nx
Opening of ovari an bursa
M e s o v a r/um
Ovary Abd. osti um of o v i duc t
M e s o m e t r i um -----
layer (tunica mucosa) is made up of partially and uterus when sexual sterility is desired, or
ciliated simple columnar epithelium, the motion when primary infections of the uterus indicate
of the cilia being directed toward the uterine the necessity for this operation.
hom. The mucosa is folded slightly (plicae
tubariae) near the uterine ostium and greatly THE UTERUS
near the abdominal ostium. Structurally, the
fimbriae are highly vascular, but, unlike the The uterus is a hollow muscular organ which
tubal portion of the duct, they contain few serves as the habitation for the developing
muscle fibers. young. It gives attachment to the fertilized
ovum and functions as a source of fetal nour
Vessels and Nerves ishment. It also serves as the route by which the
sperm may reach the ovum in the oviduct.
The oviduct is supplied by the ovarian and The uterus consists of a neck or cervix (cer
uterine arteries. The two vessels anastomose vix uteri), a body (corpus uteri), and two homs
near the cranial extremity of the uterine hom. (cornua uteri). It is a tubular, Y-shaped organ
The veins are satellites of the arteries. The lym which communicates with the oviducts cranially
phatics follow the ovarian lymph ducts to the and the vagina caudally (Fig. 15-25). Its size
lumbar lymph nodes. Anderson (1927), Samp varies considerably, depending on age, size of
son (1937), and Ramsey (1946) have worked out animal, number of previous pregnancies, and
the detailed anatomy of the lymphatics of the whether the animal is currently pregnant. In
oviduct in various species of mammals. For the nulliparous mature female dogs of 25 pounds
most part, the numerous lymphatic channels in weight, the uterine horns average 10 to 14 cm.
the mucosal folds of the infundibulum and fim long and 0.5 to 1 cm. in diameter. They diverge
briae, as well as those of the tube, drain into the from the body of the uterus at a point 4 to 5 cm.
lymph vessels of the mesosalpinx. cranial to the symphysis pelvis. The uterine
Nerves to the oviduct are derived primarily body is 1.4 to 3 cm. long and 0.8 to 1 cm. in di
from the thoracolumbar (sympathetic) division ameter. The cervix averages 1.5 to 2 cm. long.
of the autonomic nervous system and pass A small caudal portion of the cervix may pro
through the aortic and renal plexuses. Fibers trude 0.5 to 1 cm. into the vagina. The diameter
from the pelvic plexus (parasympathetic) also of this intravaginal cervix is approximately 0.8
innervate the oviduct (Mitchell 1938). cm. The gravid uterus lies on the abdominal
floor during the last half of pregnancy (normal
Anomalies and Variations gestation period is 63 days). Unlike the divergent
horns in the non-pregnant state, the heavy gravid
Associated with the oviduct in the mesosal homs parallel and contact each other. During
pinx are the epoophoron and paroophoron. The uterine distention, the homs flex near the mid
epoophoron is homologous with the embryonic dle of their dorsal surfaces, bending the uterus
male appendix epididymis and efferent duct, cranially and ventrally upon itself. Both the
whereas the paroophoron is homologous with ovary, with its suspensory ligament, and the
the paradidymis (tubules) of the testis. In the vagina are drawn slightly cranioventrally.
female, these structures are rudimentary and The uterine horns (cornua uteri) are musculo-
represent the embryonic mesonephros and membranous tubes, slightly flattened dorsoven-
mesonephric (wolffian) duct. Hydatids of Mor- trally, which unite at the body of the uterus.
gagni (hydatides terminales) in the adult female The horns are located entirely within the ab
are pedunculated cysts, located on the fimbriae, domen. The cranial tip of each horn is con
which represent part of the epoophoron. The nected to the ovary by the proper ligament and
paroophoron may persist in the mesosalpinx and indirectly by the broad ligament. The oviduct
occasionally give rise to cysts. Congenital steno opens into the uterine horn at or near its tip.
sis of the oviduct is not infrequent. From a lateral view, the homs have an S-shaped
appearance, with the ovarian ends being most
Clinical Considerations dorsad. Typically, the right horn is slightly
longer than the left. Gravid homs contain a
Although rarely of clinical importance, pri series of dilatations (ampullae) which represent
mary conditions of the oviduct are generally the positions of the fetuses, separated by con
corrected by ovariohysterectomy. Secondarily, strictions.
the oviducts are removed along with the ovaries The body of the uterus (corpus uteri) is usu
T he U terus 787
ally located in both the pelvic and the abdomi mesovarium and mesosalpinx have been dis
nal cavity. Generally, the largest portion is in cussed in the descriptions of the ovary and of
the abdomen, and in multiparous bitches the the oviduct. The mesometrium begins on a
entire uterine body may be located cranial to the transverse plane through the cranial end of the
brim of the pelvis. The body extends from uterine horn and extends caudally as far as the
the point of convergence of the uterine horns to cranial end of the vagina. It is attached periph
the cervix (Fig. 15-25). There are three open erally to the lateral pelvic wall. The medial
ings into the uterine body: one from each uter surfaces of the uterine horns are connected to
ine hom and one from the internal orifice of the each other for approximately 1 cm. by a trian
cervix. An internal musculomembranous pro gular-shaped, double layer of peritoneum. The
jection extends 1 cm. into the body of the mesometrium and the lateral ligament of the
uterus, separating the horns. This partition is bladder fuse at their attachments to the pelvic
not discernible externally. The canal of the cer wall.
vix is directed caudoventrally from uterus to The round ligament of the uterus (ligamen-
vagina. The cervix lies diagonally across the tum teres uteri, chorda inguinalis) is attached
uterovaginal junction. Its ventral border at to the cranial tip of the ipsilateral uterine hom
taches to the uterine wall cranial to its dorsal and is, apparently, a caudal continuation of the
attachment. Consequently, the internal orifice suspensory and proper ligaments of the ovary.
of the cervical canal (ostium uteri internum) is All three ligaments consist largely of smooth
facing almost directly dorsally, whereas the ex muscle, allowing for stretching during preg
ternal orifice (ostium uteri externum) is directed nancy. The round ligament runs in the free edge
toward the vaginal floor. The external uterine of the peritoneal fold given off from the lateral
orifice opens on a rounded hillock projecting surface of the mesometrium. It extends cau
into the vagina (vaginal part of cervix). The dally, ventrally, and mesially, toward the inter
cervical canal averages 0.5 to 1 cm. in length nal inguinal ring. In some bitches, the round
and is closed during pregnancy by a mucous ligament with its peritoneal covering remains
plug. wholly within the abdominal cavity. In most,
however, the ligament with its peritoneal in
Relations vestments (processus vaginalis) passes through
the inguinal canal and terminates subcutane-
Dorsally, the uterus is in contact with the de ously in or near the vulva. Zietzschmann
scending colon, the psoas muscles, the trans (1928) found the vaginal ring to be absent on
verse abdominal muscle, and the ureters. both sides in 32 per cent of his specimens and
Ventrally, it contacts the urinary bladder, the absent on one side in 12 per cent. The vaginal
greater omentum, the jejunum, ileum, and de process is accompanied in its course through
scending duodenum (Fig. 15-22). The parietal the inguinal canal by the external spermatic
and visceral peritoneum investing these struc nerve and the external pudendal artery and
tures is, of course, interposed between the vein. The fascial layers enveloping the vaginal
above organs and structures as well as between process are the same as those described with
the neighboring blood vessels, nerves, and the spermatic cord and vaginal tunics of the
lymphatics. The rectouterine space, a potential male.
peritoneal cavity between the rectum and the
uterus, is continuous with the pararectal fossa. Structure
The vesicouterine space, between the urinary
bladder and the uterus (with its attached broad The uterus is made up of three tunics: se
ligament), is separated from the paravesical rosa, muscularis, and mucosa. The tunica
fossa by double peritoneal folds, the lateral serosa (perimetrium) is the layer of peritoneum
ligaments of the bladder. which covers the entire uterus. It is continuous
with the mesometrium of the broad ligament.
Ligaments The tunica muscularis (myometrium) con
sists of a thin, longitudinal outer layer and a
The broad ligaments, containing some fat thick, circular inner layer of involuntary mus
and unstriped muscle, attach the uterus and cle. Within the circular layer, close to its junc
ovaries to the body wall. The mesometrium is tion with the longitudinal layer, is a vascular
that part of the broad ligament which attaches layer (stratum vasculare) containing blood ves
the uterus to the dorsolateral body wall. The sels, nerves, and circular and oblique muscle
788 C h ap ter 15. T he U r o g e n it a l System and M am m ary G lan ds
fibers. The circular layer is especially thick in ceral branch of the internal iliac. The artery
the region of the cervix. enters the mesometrium at the level of the
In describing the process of labor in the cervix (Figs. 15-1, 15-22). Upon entering the
bitch, Rudolph and Ivy (1930) discuss the ac broad ligament, the artery lies relatively close
tion of uterine musculature in detail. Briefly, to the body of the uterus. It diverges from the
the fetus is advanced by a strong circular con uterine horn until it approaches the cranial ex
traction that progresses like a cylindrical band, tremity of the horn, where it anastomoses with
and by a longitudinal shortening. The “retreat” the ovarian artery. The uterine artery ramifies
of the fetus is prevented by a persistent longi in the wall of the uterus (stratum vasculare)
tudinal contraction. In the body of the uterus, and in the mesometrium. Branches supply
transverse circular contraction, with some both sides of the uterine horn.
longitudinal shortening, moves the fetus into The uterine and ovarian veins follow a
the vagina. Contraction of the abdominal mus course similar to that of the arteries, except at
cles, as well as of the vaginal musculature, their terminations. The right ovarian vein emp
causes the final expulsion of the fetus. Rey ties into the postcava at the level of the right
nolds (1937) describes uterine motility, during ovary, whereas the left enters the left renal
estrus, as being a series of simple myometrial vein. Both ovarian veins are very tortuous in
contraction waves, since intermediation of an their course between the peritoneal layers of
intrinsic innervation is not essential to it. the broad ligament.
The tunica mucosa (endometrium) is the Studying the physiological aspects of uterine
thickest of the three uterine tunics. Facing the circulation during pregnancy, Reynolds (1949)
lumen of the uterus is a layer of low columnar found two distinct phases in the adjustment of
epithelium whose cells are only temporarily the uterine vessels to the shape and size of the
ciliated (Trautmann and Fiebiger 1952). Simple conceptus. First, there is progressive stretch
branched tubular glands are present in the ing of the blood vessels and, secondly, the blood
lamina propria. Opening into the uterine cav vessels during the latter part of gestation sepa
ity, these glands are generally very long and rate from one another without increase of
are separated by shorter, inconstant glands or length. Burwell and his co-workers (1938)
crypts. The long glands in the bitch show rela found that blood pressure in the femoral and
tively little branching or coiling, in contrast uterine veins was elevated during pregnancy
with those of the mare or cow. They generally in the dog. However, pressure in the uterine
traverse the entire thickness of the endome vein is higher than that in the femoral vein. The
trium. Grossly, the mucosal surface of the uterine veins drain into the postcava. Two of
uterus is reddish in color and may either be the principal functions served by rhythmic
smooth or contain low longitudinal ridges uterine contractions during estrus, according
which obliterate the uterine cavity in the non to Fagin and Reynolds (1936), may be produc
pregnant state. The cervical canal does not tion of an increased volume flow of blood
contain the relatively high mucosal folds ob through enlarged, hyperemic vessels, and re
served in other domestic animals, but is closed moval of any edematous fluid.
in pregnancy by a mucous plug and muscular The lymphatics from the uterus pass to the
contraction. The cervix of the non-gravid uterus internal iliac and lumbar lymph nodes. In
is composed predominantly of fibrous connective studying the lymphatics of the uterus and
tissue, with an average of only 15 per cent of vagina of the Rhesus monkey, Wislocki and
smooth muscle (Danforth 1947). Dempsey (1939) found conspicuous lymphatic
networks in both the endometrium and the
Vessels and Nerves myometrium. In pregnancy, these networks
hypertrophy. The vagina possesses a dense
The uterus is supplied with arterial blood via plexus of lymphatics in its tunica propria.
the ovarian and uterine arteries. The origin of The uterus receives sympathetic and visceral
the ovarian arteries from the aorta has been afferent fibers through the hypogastric plexus
discussed with the description of the ovaries and parasympathetic and visceral afferent
and of the oviducts. The ovarian artery anas fibers via the pelvic nerves. Kollar (1953) found
tomoses with the uterine artery, one of the that bilateral nerve resection in the rat inter
principal branches of the urogenital artery, rupts the afferent portion of the “genital nerve-
arising from the urogenital 11 to 18 mm. distal pituitary pathway.” Thus copulatory stimuli,
to the origin of the latter vessel from the vis not reaching the anterior lobe of the pituitary,
T he U terus 789
fail to initiate the hormonal sequence of events straight, poorly developed uterine glands. If
which inhibit the next estrus and allow proges pregnancy does occur during estrus, a decidu-
tational changes to occur. The finer distribution ate, zonary placenta develops within the uterus.
of the nerves to the myometrium has been The uterine horns undergo marked enlarge
studied by Pallie et al. (1954) in the rabbit. ment, increase in weight, and become con
These authors found a plexus of finely myeli stricted around the growing fetuses. In late
nated and unmyelinated fibers in the myome pregnancy, the cornua lie on the ventral ab
trium and under the mesometrium. No ganglion dominal wall and pull the ovaries slightly
cells and no specific sensory endings were cranioventrally. Blood vessels supplying the
found. It is suggested that this “primitive” type uterus increase in size. The cervix softens or
of nerve supply is sufficient to control uterine relaxes. The gestation period in the bitch is ap
musculature, which is described as being proximately 63 days. Following parturition, the
“broadly coordinative, slow-acting, not sharply uterine musculature and mucosa involute.
localized; and . . . nonspecific in that the motor Congenital anomalies of the uterus include
response to nervous stimulation is conditioned uterus didelphys (two distinct uteri occurring
by the dominant ovarian hormone.” side by side), uterus unicornis (one hom), and
Rudolph and Ivy (1930) advance the theory atresia of the cervix. Pseudocyesis, or false preg
that the pelvic nerve exercises a tonic inhibi nancy, is of unknown origin (DeVita 1946).
tory action and the hypogastric a tonic motor It may occur in the nulliparous as well as the
action on the uterus of the dog. They state that multiparous bitch. The condition occurs dur
the muscular activity of the uterus of the dog ing metestrus and is evident at the time when
is controlled by its intrinsic nervous mechanism normal parturition should occur. In some
and that the extrinsic nerves play a relatively instances, it is accompanied by abundant lacta
minor role in regulating uterine motor activity. tion. Experimental pseudocyesis following in
duced ovulation persists 40 to 44 days (Foster
and Hisaw 1935). At the end of this period
Anomalies and Variations retrogressive changes are apparent in the uterus
and corpora lutea, and spontaneous uterine
The sexual cycle in the bitch, lasting from 4 motility, reduced during the false pregnancy,
to 6 months, is responsible for periodic uterine shows a definite increase.
changes. There are four phases to the uterine
cycle: proestrus, estrus, metestrus, and anestrus.
Proestrus lasts approximately 9 days (Evans and Clinical Considerations
Cole 1931). This period is followed by estrus
(heat), which is characterized by vulvar swell Ovariohysterectomy is frequently done to
ing. Estrus also lasts about 9 days, during sterilize the bitch or to correct pyometra, as
which time the endometrium becomes edem mentioned in the description of the ovaries.
atous and hyperemic, and is covered with a Dystocia (difficult birth) is a distinct clinical
sanguineous mucous secretion. The uterine problem, particularly in breeds of dogs that
glands become longer, the mucosal columnar normally have a small pelvis. Although dysto
epithelium becomes higher, and the external cia frequently may be corrected by the use of
genitalia become more swollen. Ovulation oc pituitary extract and manipulation, cesarean
curs during the first few days of estrus. Metes section is often the only solution. In breeds
trus, following estrus, is an indeterminate that are highly susceptible to dystocia, such a
period, during which the corpora lutea are de procedure is routine. It is usually performed
veloped. Evans and Cole (1931) describe through a mid-ventral abdominal incision. The
metestrus in the bitch as being similar to a uterine incision is made either through the
combination of metestrus and diestrus in poly- mid-ventral surface of the uterine body or
estrous animals. During diestrus, the uterine through the avascular surface of one uterine
glands become longer, more coiled, and more hom, peripheral to the broad ligament, the
active. If pregnancy does not occur, the uterine pups from both homs being manipulated
mucosa regresses and the corpora lutea degen through the one incision.
erate. The combined period of metestrus and Recurrent pseudocyesis is generally cor
diestrus usually takes three months. Anestrus, rected by ovariohysterectomy. Huggins and
which lasts two months, is characterized by Moulder (1944) recommend that the operation
low cuboidal epithelium in the uterus and for this purpose be done during anestrus.
790 Chapter 15. T he U r o g e n it a l System and M am m ary G lan ds
estrus. The everted vaginal mucosa may be in are soft and pliable, being composed of fibrous
flamed, edematous, or necrotic. Surgical re and elastic connective tissue, smooth muscle
moval of excess prolapsed tissue is usually fibers, and an abundance of fat. The vaginal
indicated. Benign neoplasms (leiomyoma, leio- processes, containing the round ligaments of
myofibroma, lipoma, and vaginal polyps) are the uterus, end in the subcutaneous connective
found. Venereal sarcoma is the most common tissue of the labia (Fig. 15-23). These are not
malignant tumor of the vagina. When neces present in all animals (see description of uter
sary, tumors are surgically removed. Venereal ine ligaments). The distance between the dorsal
sarcoma may be transmitted by copulation. commissure and the anus is 8 to 9 cm. The dorsal
Consequently, mating of afflicted bitches is commissure lies at or slightly below a frontal
generally prohibited (Allam 1952). According plane passing through the symphysis pelvis. The
to Runnells (1954), the condition may disap ventral portions of the labia, with their uniting
pear spontaneously and be followed by perma commissure, form a pointed projection extend
nent immunity. Prolonged dystocia may cause ing downward and backward from the body.
the development of fistulas into the rectum or The clitoris, the homologue of the male
bladder. Lacerations of the vaginal walls may penis, is composed of paired roots (crura cli-
occur during parturition. These traumatic con toridis), a body (corpus clitoridis), and a glans
ditions are corrected by local surgical repair. (glans clitoridis). The roots and body are hom-
ologues of the male corpora cavernosa penis,
THE VULVA AND FEMALE URETHRA and the glans clitoridis (possessing erectile tis
sue) is homologous with the glans penis, al
T h e V u lv a though it is not bipartite in structure. The body
of the clitoris in the dog is composed primarily
The vulva (pudendum femininum), or ex of fatty rather than erectile tissue. It is covered
ternal genitalia (partes genitales externae), con by a thick tunica albuginea. The clitoris of the
sists of three parts: vestibule, clitoris, and labia. dog does not contain any structures compara
The vestibule (vestibulum vaginae) is the ble to the os penis, the corpus cavernosum ure
space connecting the vagina with the external thrae, and the urethra of the male. Instead, in
genital opening (Fig. 15-25). It develops from the bitch, there are elongate masses of erectile
the embryonic urogenital sinus, the common tissue, the vestibular bulbs (bulbi vestibuli),
opening for genital and urinary tracts. The lying deep to the vestibular mucosa and united
space is variable in size, depending on the size to each other by an isthmus. They lie in close
of the animal and whether or not she is preg proximity to the corpus clitoridis, correspond
nant. In a non-pregnant, mature 25-pound dog, ing to the bulb of the urethra in the male. The
the external vulvar opening or cleft is approxi vestibular bulbs are each supplied by a termi
mately 3 cm. long. The distance from the ven nal branch of the internal pudendal artery,
tral commissure of the vulva to the urethral homologous to the artery of the urethral bulb
opening is 5 cm., and the diameter of the in the male. The glans clitoridis, erectile in
vaginovestibular junction is 1.5 to 2 cm. structure, is very small and projects into the
The urethral tubercle is a ridgelike projec fossa clitoridis. The fossa is partially folded
tion on the ventral floor of the vestibule, near over the glans clitoridis dorsally. This fold cor
the vaginovestibular junction. It contains the responds to the male prepuce. The free part of
external urethral orifice (ostium urethrae ex the clitoris (glans) is about 0.6 cm. long and 0.2
ternum). The tubercle, widest cranially, nar cm. in diameter in an average-sized dog; the
rows caudally to an apex located at a point ap distance from the ventral commissure of the
proximately half the distance from the urethral vulva to the glans clitoridis is 2 to 3 cm., and
opening to the clitoris. A shallow fossa or de that from the ventral commissure to the fundus
pression is present on each side of the tubercle. of the fossa of the clitoris is 3 to 4 cm. The open
The mucosa of the vestibule is not covered ing of the fossa is approximately 1 cm. in di
with distinct ridges, as is the mucosa of the ameter.
vagina, but is relatively smooth and red.
The labia (labia pudendi), or lips, form the Structure
external boundary of the vulva. Homologous
with the scrotum of the male, the labia fuse The mucosal surface of the vulva is covered
above and below the vulvar cleft to form dor by stratified squamous epithelium. A variable
sal and ventral vulvar commissures. The labia number of lymph nodules may cause promi-
79 2 Chapter 15. T he U r o g e n it a l Sy stem and M am m ary G la n ds
Suspensory l ig. o f ov ar y
—Oviduct
- ~ M e s o s a I pi nx
- - P r o p e r lig. of ov ar y
i--------- L. u t e r i n e horn
Body o f ut er us
Vagina- ^
U rethral opening S a g i t t a l s e c t i o n t h r ou g h c e r v i x
V e s t i b u l a r bulb-
V e stib u le ^
C onstrictor}
vestibuli j
Cl i t o r i s - - Labium
Fossa c l i t o r i d i s '
F ig . 15-25. Dorsal view of female genitalia, partially opened on mid line. Smaller view shows sagittal section through cervix.
T he V ulva and F em ale U reth ra 793
M. sacr ococcygeus v e n t r a l i s la
M. rect ococcygeus -
— M. coccygeus
F i b e r s f r o m c o c c y g e a l vertebrae I v l l — ®
M . s p h i n c t e r a n i ext. - M. l e v a t o r a n !
M. c o n s t r i ct o r v e s f i b u l
^ M . u r e t h r a l i s coveri ng urethra
Symphysi s p e l v i s
M. c o n s t r i c t o r v u l v a e
M. i s c h i o u r e t hr a l i s
C r u s c l i t or j i di s 1
lM. i schi acavernosus
nences to appear on the mucosa. Small minor ischii, on each side, and insert upon the poorly
vestibular glands, lobular in structure, open developed central tendon of the perineum.
ventrally on each side of the median ridge con The ischiocavernosus muscles are small in the
nected to the urethral tubercle. The glands are female. They arise bilaterally from the caudal
located deep to the constrictor vestibuli mus edge of the ischium and attach to the crura cli
cles. The body of the clitoris consists of fat, toridis. This is similar to the manner in which
elastic connective tissue, and a peripheral tunica they insert upon the corpora cavernosa penis in
albuginea. The glans clitoridis, made up of erec the male.
tile tissue, contains numerous sensory nerve
endings. The vestibular bulbs are also composed T he F em ale U rethra
of cavernous tissue. The labia, covered with
stratified squamous epithelium, are rich in se The urethra of the bitch (urethra feminina)
baceous and tubular glands and also contain fat, (Fig. 15-22) corresponds to that portion of the
elastic tissue, and smooth muscle fibers. male urethra which lies cranial to the prostatic
utricle. It is about 0.5 cm. in diameter and 7 to
Muscles 10 cm. long. It originates from the urinary blad
der at or near the cranial edge of the symphysis
In addition to the usual unstriped muscle pelvis. It extends caudodorsally to enter the
fibers, similar to those of the vagina, the vulva genital tract approximately 0.5 cm. caudal to
possesses two striated circular muscles (Figs. 15- the vaginovestibular junction. Its dorsal wall is
26, 15-27). The most cranial of the two is the in close apposition to the ventral wall of the
strong constrictor vestibuli muscle. It is incom vagina. Structurally, the female urethra resem
plete on the dorsal surface of the vestibule, but bles that of the male. It is lined by folded mu
fuses along its caudal border to the external cous membrane, allowing the urethral lumen to
sphincter of the anus. Its fibers run diagonally expand considerably when under pressure. The
in a cranioventral direction, encircling the ure mucosa is non-glandular, and the submucosa is
thra, vestibule, and caudal portion of the vagina highly vascular. Lymph nodules are also pres
before it joins its fellow of the opposite side. It ent. The musculature of the female urethra
constricts the vestibule and, during copulation, consists of outer and inner longitudinal and
tightens behind the bulbus glandis of the penis, middle circular layers of unstriped muscle. The
preventing the male from dismounting for about smooth muscles become less conspicuous near
15 minutes following initial ejaculation. It is the entrance of the urethra into the vestibule.
thought that some semen still flows during this At the external urethral orifice, voluntary mus
period. cle encircles all but the dorsal surface of the
Immediately caudal to the constrictor ves urethra, which is in close contact with the ves
tibuli muscle is the relatively weak, thin constric tibule. These circular fibers form a strong
tor vulvae muscle (the muscle of the labia). sphincter at the external orifice.
This muscle is continuous dorsally with the ex
ternal anal sphincter, arising from the coccygeal Vessels and Nerves
fascia ventral to the first and second coccygeal
vertebrae, and encircles the vulva and vestibule The external genitalia and urethra of the fe
about 1 cm. caudal to the point where the male are supplied with blood through the uro
urethra disappears into the genital tract. The genital and the external and internal pudendal
constrictor vulvae muscle blends with the vestib arteries. The external pudendal artery sends
ular constrictor to a slight degree. The vulvar branches to the labia (cranial labial artery), cor
constrictors fuse together below the vulva, cra responding to the scrotal branches in the male.
nial to the ventral commissure. They lift the The urogenital artery supplies the vulva by
labia dorsally prior to intromission of the penis, means of the cranial vestibular branches, from
allowing it to enter the vagina more easily. The the vaginal ramus, and the caudal vestibular
vestibular and vulvar constrictors together are branches from the termination of the urogeni
homologous with the bulbocavemosus muscles tal artery. The clitoris is supplied by branches
of the male. They lie superficial to the vestib of the internal pudendal artery, corresponding
ular bulbs. Their counterparts in the male are to the dorsal and deep penile arteries. The ves
peripheral to the bulb of the urethra. The tibular bulb is also supplied by the internal pu
ischiourethralis muscles (transversi perinei) dendal artery (homologous to the artery of the
arise from the caudomedial surface of the tuber urethral bulb in the male).
T he V u lva and F em ale U reth ra
M. rec t oc oc c ygeus
M. coccygeus
— M. l e v a t o r ani
- - M . s ph i nc t e r ani ext
M. o b t u r a t o r i nt . __
— Tuber i s c h i i
M. i schi ou r e t hr a l i s
^ - - M . ischiocavernosus
M. c o n s t r i c t o r v e s t i b u l i
- - M . c o n s t r i c t or vuivae
The bilateral veins from the clitoris (dorsal parturition and for correcting vaginal prolapse
veins of clitoris) join each other at the ischial that may occur during estrus. Permanent episi
arch and then separate again (after a distance of otomy is recommended by Blakely (1952a) in
1 or 2 cm.) into internal pudendal veins which dyspareunia. Episioplasty is frequently indicated
drain into the internal iliac veins. The vestibular in cases of perivulvar dermatitis.
bulb is drained by a separate tributary of the
internal pudendal. Valves are apparent in most
of the veins, including the common trunk of the E M B R Y O LO G Y O F T H E
dorsal veins of the clitoris. The dorsal arteries U RO G EN ITAL SY ST EM
and veins of the clitoris are not actually dorsal
to the clitoris, in the manner of the compara The development of the mammalian urogeni
ble penile vessels. They curve ventrally around tal system is briefly summarized here. For spe
the ischial arch and run caudoventrally along cific details, one should refer to textbooks of
that surface of the clitoris which corresponds to embryology and comparative anatomy, such as
the dorsum of the penis. Lymphatic drainage those by Arey (1954) and Romer (1950). The
compares with that of the external genitalia of homology of the structures in the male and fe
the male. male is shown in Table 1.
The sensory afferent nerves to the external The embryonic intermediate cell mass, or
genitalia are derived from the pudendal and the nephrotome, divides during early fetal life into
genital nerves. Both nerves innervate the labia. a urinary, or nephric, region and a genital re
The glans clitoridis receives its sensory nerves gion. Segmental tubules (the pronephros) de
from the pudendal (dorsal nerve of the clitoris). velop in the cranial portion of the nephric
Motor impulses to the urethral muscle and to region (mesomere). The tubules, one pair per
the vestibular and vulvar constrictors also pass segment, grow caudally and form a longitudinal
through the pudendal nerve. Autonomic inner duct (pronephric duct), which runs to the prim
vation to the external genitalia and urethra in itive cloaca. The funnel-shaped connection of
the female is through the hypogastric and pelvic each segment with the coelom (body cavity) is
nerves. It includes principally sympathetic called the nephrostome. Renal arteries push out
fibers which innervate the musculature of the from the aorta to form glomeruli, one per seg
blood vessels. ment. Caudal to the pronephros, other segmen
tal tubules unite with the pronephric duct to
Anomalies and Variations form the mesonephros, or wolffian body. As this
occurs, the pronephros degenerates, and the
Incomplete degeneration of the wolffian pronephric duct becomes what is called the
(mesonephric) ducts may be the cause of con mesonephric, or wolffian, duct. The mesoneph
genital anomalies. During pregnancy, the mu ric tubules are more complex than their prede
cosa of the vulva will exhibit changes similar to cessors. Although the pronephros is functionless
those occurring in the vagina. Normal physio in mammals, the mesonephros is thought to
logical changes during the estrus cycle (varia serve as a temporary organ of excretion. Unlike
tions of size and shape) are also apparent. the pronephros, the mesonephric glomerulus is
internal, and the nephrostome does not act as
Clinical Considerations a mouth for the tubule. Later, a bud develops
from the mesonephric duct near its entrance
Recognition of the location of the external into the cloaca. This bud gives rise to the renal
urethral orifice is important for the purpose of pelvis, ureter, and the collecting tubules of the
catheterization. Not uncommonly, the clitoris definitive, or metanephric, kidney. The secre
or the fossa clitoridis is mistaken for the urethral tory tubules and Bowman’s capsules of the
opening, and the catheterization attempt is un metanephros develop from the caudal portion
successful. The urethra opens on a tubercle 4 of the nephric ridge.
to 5 cm. cranial to the ventral commissure of The three types of kidneys in vertebrates ap
the vulva. pear successively in cranio-caudal sequence both
Episiotomy, or incision of the vulva, is fre ontogenetically and phylogenetically. The pro
quently performed on bitches. The operation, nephros, or more commonly the mesonephros, is
consisting of an upward incision in the dorsal the functional kidney of anamniotes (fish and
commissure of the vulva, is indicated for the amphibians), whereas the metanephros is the
purpose of extirpating certain tumors, for easing functional kidney of adult amniotes (reptiles,
birds, and mammals).
E m bryo lo g y of th e U r o g e n it a l System 797
Table 1. Homologies of Genital Organs in portion of the mesonephric tubules, which be
Male and Female Mammals come the epididymis. The middle tubules form
the ductus deferens, and the caudal ones persist
FEM A LE
M A LE as the vestigial paradidymis and ductuli aber-
Testis Ovary rantes. The mesonephric tubules do not join the
Mesorchium Mesovarium ovaries in the female, persisting as the vestigial
Appendix testis Abdominal ostium epoophoron and paroophoron. The mesoneph
of oviduct ric ducts also degenerate during the course of
Gubemaculum Round ligament of embryonic development. In rare instances,
uterus, proper lig Gartner’s ducts (remnants of the caudal parts of
ament of ovary the mesonephric ducts) open near the vagino-
Prostatic urethra Urethra vestibular junction in the adult.
(cranial to utricle) The ovaries and testes migrate caudally dur
Prostatic urethra (caudal Vestibule ing fetal development. In addition, the male
to utricle) and mem gonads leave the body cavity and descend into
branous urethra the scrotum, moving from their dorsolateral po
Penis Clitoris sitions in the abdomen toward the inguinal
Glans penis Glans clitoridis canals, and migrating between the abdominal
Corpus cavernosum Vestibular bulbs wall and the parietal peritoneum. During early
urethrae fetal development, when the testes are still high
Corpus cavernosum penis Corpus cavernosum in the abdomen, parietal peritoneum begins to
clitoridis push into the inguinal canals. These peritoneal
Scrotum Labia outpocketings are the primordia of the vaginal
Scrotal raphe Dorsal commissure processes. Each vaginal process evaginates
of labia through its inguinal canal into the scrotum.
Prepuce Fold of fossa During the latter part of embryonic life, each
clitoridis testis descends through the inguinal canal into
the scrotum, passing between the vaginal proc
ess and the scrotal wall. The testis always re
mains outside the peritoneal cavity, byt both
The growth of the kidneys and urinary blad the testis and gubemaculum are enveloped by
der in the fetal dog has been studied in de peritoneum before descent begins. The shorten
tail by Latimer (1951). Gersh (1937), in studying ing of the gubernaculum, both actual and rela
renal development in the rabbit, cat, and pig, tive, is thought to draw the testis into the
found that the mesonephros and metanephros scrotum. Some workers attribute the testicular
may function both simultaneously and con descent to a process of normal herniation
tinuously. Renal elimination is a continuous through a canal previously dilated by the guber
function without any interruption being caused naculum. The peritoneal layers within the scro
by degeneration of the mesonephros. tum form the tunics for the testis and spermatic
The gonads develop from the ventromedial cord. Unlike in the human, there is no disap
portion of the intermediate cell mass (genital pearance of tunics in the area between the ab
ridge). The sex of the embryo is genetically de dominal cavity and the testis. The vaginal
termined at fertilization, but is not distinguish process contains a cavity which is continuous,
able morphologically until much later in fetal throughout life, between the peritoneal cavity
life. of the abdomen and the vaginal cavity of the
According to Gruenwald (1942), the coelomic scrotum. The ductus deferens and the spermatic
wall forms a uniform gonad blastema during the vessels and nerves are pulled into the scrotum
initial phases of gonad development, by con by the testis and epididymis, causing the duc
tributions from mesothelium and mesenchyme. tus deferens to loop over the ureter.
The primary sex cords differentiate within the In the female, the ovaries remain in the ab
gonad blastema and other cords associated with dominal cavity, suspended by the broad liga
them form the rete and, in the testis, interstitial ments. The vaginal processes of the bitch, con
cells. Gruenwald states that the tunica albu taining the round ligaments of the uterus,
ginea forms in the testis, separating the sex migrate toward the labia in a manner similar to
cords from the superficial cell layer. the migration of the vaginal processes of the
The developing male testes join the cranial male toward the scrotum. The female counter-
798 Chapter 15. T he U r o g e n it a l Sy stem and M am m ary G la n ds
part of the male gubernaculum fails to unite in receiving the paired wolffian and miillerian
the same way because of the persistence of the ducts, develops into the urinary bladder, the
fused miillerian ducts. urethra, and the vestibule of the vagina in the
Mossman (1938) proposed the theory that the female. The wolffian duct in the male becomes
follicular epithelium and lutein cells of mam the ureter and ductus deferens. The ureter
malian ovaries are both ontogenetically and opens into the bladder, and the ductus deferens
phylogenetically specialized portions of the opens into that part of the urethra which con
coelomic epithelium and that the follicular cavi nects the bladder with the urogenital sinus. The
ties are homologous to the coelom. prostate gland, endodermal in origin, arises
Paired miillerian ducts also develop in both from the urethral epithelium. The prostatic tu
male and female (Fig. 15-28). In the bitch, the bules originate as solid epithelial projections
miillerian ducts open cranially into the perito from the prostatic urethra.
neal cavity as the abdominal openings of the During early fetal life the external genitalia
oviducts. Caudally, they unite, in part, to form are in an indifferent or indeterminate condition.
the oviducts and the bicornuate uterus and, Genital tubercles (rounded labioscrotal swell
completely, to form the vagina. In the male, the ings) bound the phallic eminence. The ventral
miillerian ducts degenerate, except for their surface of the phallus contains a urethral groove
cranial and caudal ends. Cranially, they remain and, on its floor, a urethral membrane. In the
as the vestigial appendix testis. Caudally, the male, the phallus develops into the penis (with
ducts unite and persist as the uterus masculinus its cavernous bodies). The urethral membrane
(prostatic utricle), opening on the colliculus breaks through, and the borders of the urethral
seminalis. The broad ligament of the embryo groove fuse, forming the penile, or cavernous,
encloses the wolffian and miillerian ducts in its urethra. The genital tubercles fuse on the mid
free edge. line, forming the scrotum. In the female, the
The terminal blind end of the primitive hind- clitoris arises from the indeterminate phallus,
gut (cloaca) divides into the rectum dorsally the urethral groove forms the vestibule, and the
and the urogenital sinus ventrally. The latter, genital tubercles become the labia of the vulva.
T H E M A M M A R Y G LA N D S
The mammary glands (mammae) are modi tively small cranial four are the thoracic mam
fied cutaneous glands. For the reason stated mae, the following four are the abdominal
earlier, their close physiological relationship mammae (median in size), and the relatively
with the reproductive organs, they are described large caudal two are the inguinal, or pubic,
here, rather than with the integument. Mam mammae. In some instances, especially during
mae are characteristic of mammals, and provide involution of the glands following lactation, the
nourishment to the newborn. In the male they relative sizes of the glands may digress from the
remain rudimentary throughout life, but in the typical pattern. Under these circumstances, the
female they are subject to conspicuous changes caudal abdominal mammae may rarely be
during pregnancy and during and after lacta slightly larger than the inguinal glands. Turner
tion. and Gomez (1934) examined 20 dogs and found
The mammary glands are typically arranged 10 glands in each of 16 dogs, 9 in 3, and 8 in 1.
in two bilaterally symmetrical rows extending Supernumerary glands are found in both tho
from the ventral thoracic to the inguinal region racic and abdominal regions. The mammary
(Fig. 15-29). The teats (papillae mammae) in glands of most larger domestic animals have
dicate the position of the glands in the male or been studied in great detail (Emmerson 1941,
in the non-lactating female. The number of Foust 1941, Turner 1952, Weber et al. 1955).
glands varies from 8 to 12, with 4 to 6 gland The mammae of the elephant, whale, monkey,
complexes on each side of the mid line. Most and small laboratory animals have been investi
commonly, there are a total of 10 glands, and gated (Schenke 1924, Speert 1948, Richardson
8 are more frequently seen than 12. In a few 1955). Study of the mammary glands of the dog,
cases, there are more glands on one side than however, has been comparatively neglected.
on the other. Four pairs of glands are more
commonly found in the smaller breeds (Kitt Structure
1882). When 10 glands are present, the rela The mammary gland consists of epithelial
799
- -Ovi d u c t
Epoophoron
Ovary- -
Infundibulum - '
Paroophoron
Gartner's duct
Bladder-
B. A d u l t Fern a I <
Pa r a d i d y m i s
Ductus d e f e r en s
Epididymis-
/ Uterus m a s c u l i n u s
Appendix epididymidis
' , Pro s t a t e
Efferent d uctules'
Rectum
Appendix testis /
- M e m b ra n o u s u r e t h r a
Testi
B la d d e r' - Pe n i s
D e s c e n de d t e s t i s
C. A d u l t Male Scrotum
F ig . 15-28. Schematic representations of indifferent stage in development of the genital system of the dog, and the genital
system in the adult female and the adult male dog.
800 Chapter 15. T he U r o g e n it a l S y s t e m and M am m ary G la n ds
Teat o r i f i c e _
Teat c a n a l - -
Teat sinus
Gland sin
glandular tissue (parenchyma), connective tis thin over the distal tip of the teat. It increases
sue (stroma), and the covering skin (cutaneous in thickness near the base. The corium contains
layer). elastic elements, smooth muscle fibers, and
The parenchyma, or secretory tissue, is pres blood vessels. The epidermis may be pigmented,
ent to a significant degree only during preg in which case the pigment is present in the ger
nancy, pseudopregnancy, the period of lacta minal layer. Although the distal blunt end of the
tion when pups are nursing, and for 40 to 50 teat is bare, the rest of it is covered by very fine
days following weaning. After this postpartum hairs which are accompanied by sebaceous
period, the alveoli and lobules are reduced to a glands. Turner (1939) found sweat glands at the
shrunken system of ducts with relatively few base of the teat.
remnants of lobules. The stroma appears rela
tively dense.
The number of ducts opening on a teat varies. Vessels and Nerves
In a study of the external teat openings of nine
mature dogs of different breeds, as few as 7 and The mammary glands are highly vascular.
as many as 16 ducts were observed on a teat. Veins are more extensive than arteries. The tho
The duct openings are located on the blunt end racic mammae receive their arterial blood sup
of the teat in an irregular, sievelike pattern. The ply from the perforating sternal branches of the
peripheral ducts tend to form a circle, whereas internal thoracic arteries. These penetrate the
the centrally placed ones form an irregular de thoracic wall through the intercostal spaces. In
sign. Turner (1939) described 8 to 14 ducts in tercostal and lateral thoracic arteries may also
one case, and 12 to 22 in another. Martin (1910) contribute blood to the thoracic glands. Ab
found from 8 to 12 ducts. dominal and inguinal mammae are supplied by
Each teat canal (papillary duct) occupies ap mammary branches of the epigastric arteries.
proximately one-third of the length of the teat. The cranial superficial epigastric artery arises
It is lined by stratified squamous epithelium. from the cranial deep epigastric artery, a branch
The epithelium usually lies in folds near the of the internal thoracic. It penetrates the ab
margin of the teat sinus. dominal wall approximately 2 to 4 cm. from the
The teat sinus (teat cistern, sinus lactiferus) mid-ventral line, mesial to the costal arch. It
extends from the teat canal into the paren sends mammary branches to the cranial ab
chyma of the gland. In large dogs, the sinus dominal gland and anastomoses with the caudal
system may be seen upon gross examination of superficial epigastric artery. The latter artery, a
the sectioned gland. The epithelium of the teat branch of the external pudendal, runs cranially,
canal changes gradually, in the teat sinus, from deep to the inguinal mamma which it supplies.
stratified squamous to columnar. The artery continues forward to supply the ab
In the middle portion of the teat (stroma) dominal mammae and terminates in numerous
there is an intermingling of diversely running superficial branches which anastomose with the
smooth muscle fibers and connective tissue ele end branches of the cranial superficial epigas
ments. The circular musculature radiates from tric artery.
the axis of the central zone of the teat into the The veins of the mammae in the dog parallel
area among the teat canals. The musculature the course of the arteries to a large degree. The
encircles the canals and joins or condenses into cranial and caudal superficial epigastric veins
the sphincters of the canals (sphincter papillae). are the major veins of the glands. The abdomi
Elastic fibers radiate among the teat canals, nal and inguinal glands drain into the caudal
forming an extensive network. Essentially, the superficial epigastric veins. The thoracic mam
tunica propria of the teat is composed of bands mae drain into the cranial superficial epigastric
of loose connective tissue, blood vessels, and veins, as well as directly into the internal tho
smooth muscle and elastic fibers. racic veins as far cranially as the fifth intercostal
According to Turner (1939), the teat and space.
gland sinuses unfold when filled with milk and Lymphatic drainage of the mammary glands
show no constrictions or circular folds between is also bilaterally symmetrical. By studies in
the two divisions of the sinuses. Each gland volving injection of solutions of Berlin blue and
sinus is separated from surrounding sinuses by India ink into the glandular tissue of each
connective tissue septa and has a distinct, sepa mamma in live, lactating bitches and observa
rate glandular area composed largely of minute tion of the flow of dye in the lymphatic vessels,
alveoli (Kappeli 1918). the findings of Stalker and Schlotthauer (1936)
The skin (integumentum commune) is very and of Baum (1918) were substantiated. Each
802 Chapter 15. T he U r o g e n it a l Sy st e m and M am m ary G la n d s
total amount of parenchyma. Changes in the of all of the glands on the affected side. If either
mammary gland during pseudopregnancy are the caudal abdominal or the inguinal mamma
essentially identical to those of pregnancy, ex contains tumors, both should be removed. The
cept that secretory activity at 60 days is less superficial inguinal lymph node is then also re
well developed. moved. Schlotthauer (1952) ascertained the
Approximately 10 days after parturition the lack of necessity for removal of the axillary
size of the mammae is greatly reduced, the lymph node when the thoracic and cranial ab
lobule-alveolar structures being affected sooner dominal mammae contain tumors. Metastasis
than the duct system. By 40 days the lobule- of mammary neoplasms to the axillary or acces
alveolar system is largely degenerated, and the sory axillary lymph nodes is infrequent in the
ducts are shrunken. After cessation of lactation, bitch. Spayed bitches rarely have mammary
in the dog, the mammary gland regresses to a tumors.
simple duct system.
Clinical Considerations
BIBLIOGRAPHY
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Anat. Rec. 6 2: 75-94. tation, mating, and maternal instincts in the adult dog.
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Francois-Franck, C. A. 1895. Recherches sur l’innervation Anat. Rec. 17: 203-219.
vasomotrice du penis; topographie des nerfs constric- -------------. 1919b. Experimental degeneration in the testis
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138, 744-816. Langley, J. N. 1896. The innervation of the pelvic and adjoin
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Gantt, W. H. 1938. Extension of a conflict based upon der in the fetal dog. Anat. Rec. 109: 1-12.
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Psychosexual Development in Health and Disease, Artificial Insemination of Farm Animals, edited by E. J.
edited by P. H. Hoch and Joseph Zubin. New York, Perry. 3rd Ed. New Brunswick, N.J., Rutgers Univer
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Gersh, I. 1937. The correlation of structure and function in and progesterone on male rabbit mammary glands;
the developing mesonephros and metanephros. Contr. varying doses of progesterone. Proc. Soc. exp. Biol.
Embryol. Carneg. Instn 2 6: 33-58. (N.Y.) 4 8: 83-86.
Gisler, E. 1922. Die Entwicklung der Milchdriise bei der MacCallum, D. B. 1926. The arterial blood supply of the
Katze. Inaugural Dissertation, Universitat Zurich. mammalian kidney. Amer. J. Anat. 38: 153-175.
Gruber, C. M. 1933. The autonomic innervation of the -------------. 1939. The bearing of degenerating glomeruli on
genito-urinary system. Physiol. Rev. 13: 497-609. the problem of the vascular supply of the mammalian
Gruenwald, P. 1942. The development of the sex cords in the kidney. Amer. J. Anat. 65: 69-103.
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and Bovine Ovarian Tumors. (Abst.) Ann. Rep. New glomeruli as related to kidney and to body weight, with
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Ed. Ithaca, N.Y. (Pub. by author.) Sampson, J. A. 1937. The lymphatics of the mucosa of the
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"Riser, W. H. 1947. Surgical removal of the mammary gland of the Udder of Cattle and Domestic Animals. Colum
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806 Chapter 15. T he U r o g e n it a l Sy st e m and M am m ary G la n ds
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CHAPTER 16
TH E E N D O C R IN E SYSTEM
By J. F. SM ITHCORS
807
808 Chapter 16. T he E n d o c r in e Sy stem
In spite of this lack of structural continuity example is the contraction of smooth muscle of
and the fact that they develop from diverse ele the blood vessels or uterus under the influence
ments, the several features shared by the endo- of secretions of the adrenal .medulla or of those
crines lend support to their being called a sys formerly thought to be secreted by the posterior
tem. Phylogenetically, the endocrines represent lobe of the hypophysis. Certain hormones may
a series of specialized tissues which have arisen exert an inhibitory effect, but these can also be
from forms in which all cells or tissues had endo included in this group.
crine function. Histologically, they share distinc The metabolic hormones have a specific influ
tive features of finer structure, all presenting one ence on certain chemical phases of metabolism.
basic structural type of irregular epithelial or Examples are the action of insulin from the pan
epithelioid parenchyma. Physiologically, they creatic islets on carbohydrate metabolism, and
present many highly specific and integrated the regulation of calcium mobilization by the
chemical relationships, the functional mecha secretion of the parathyroid glands.
nisms for which are similar. The influence of the The morphogenetic hormones are of special
endocrines is especially noticeable during the interest in anatomy because of the bearing they
period of growth and differentiation. In fact the have on growth and development of the organ
young animal cannot reach normal adulthood ism. Examples are the effect of the thyroid hor
without properly integrated endocrine function. mone on acceleration of maturity, and that of
All of the endocrines originate in large part the anterior lobe of the hypophysis upon growth.
from some type of embryonic epithelium, the The effect of all hormones is produced by chem
functional parenchyma being of epithelial or ical action. Those of the metabolic and morpho
epithelioid tissue. In the thyroid gland there is a genetic groups probably act more or less directly
typical single-layered epithelial arrangement, upon the target organs; those of the excitatory
probably because the thyroid is the oldest endo group act via the autonomic nervous system.
crine from a phylogenetic standpoint; thus it Most of the endocrines serve more than one
presents a more perfect structure. Most of the purpose, either simultaneously or in successive
other endocrines present an irregular distribu periods. The greatest effect of the hypophysis in
tion of polyhedral epithelial or epithelioid cells early development is on body growth. As the
arranged in clusters or cords. organism approaches maturity, the growth-
All of the endocrines have a rich blood supply, promoting factor diminishes, and the gonad-
in keeping with the fact that their secretions stimulating function assumes importance. The
originate from and are poured into the blood gonads then exert an influence on development,
stream. Certain of these glands were suspected especially of the secondary sex characters. Ab
to be endocrine in function by virtue of their normal functioning of the hypophysis before
vascularity before their hormonal action was maturity may result in dwarfism or gigantism;
confirmed. There is an intimate relationship be after maturity disorders of reproductive function
tween the secretory cells and the capillary walls, are among the most characteristic of disorders
the latter forming sinusoidal plexuses about the due to hypophyseal malfunction.
parenchymal cells. Although some of the endocrines, like the
As stated before, none of the endocrines have hypophysis, secrete more than one distinct hor
excretory ducts, their products being poured mone, certain of the differing effects mediated
directly into the blood stream. This character by an endocrine gland result from different ac
istic arose secondarily in the phylogenetic his tions of one hormone. The variable response
tory of these organs. They are believed originally may result from the same hormone acting on
to have been glands of external secretion which different tissues, or from alteration in the recep
then, in the course of their evolution, lost their tivity of the same tissue with differing condi
ducts. Rudiments of ducts of certain of the endo tions or at different times. It should be kept in
crines persist in the ontogenetic development of mind that the hormones are not the immediate
these glands in higher animals. cause of the action they mediate, but that they
The most useful classification of endocrine merely influence the inherent capacity of the
glands is made upon the basis of function, which protoplasm to react in a particular way.
permits their division into three major groups: The correlation of endocrine activity is essen
excitatory, metabolic, and morphogenetic. The tial for orderly development and function of the
excitatory secretions, the name hormone being animal body. Failure of proper endocrine func
chosen on the basis of this function, influence tion is soon evidenced by alterations in develop
the functional mechanism of specific tissues. An ment or function. Endocrine dysfunction may
G en era l C o n s id e r a t io n s 809
arise in several ways. Chemical deficiencies may the field of chemical regulation of the body un
result in the failure of proper endocrine secre doubtedly involves practically all of the body
tion. For example, lack of iodine results in dis cells, such a concept, although it should be kept
order of the thyroid (goiter), with attendant in mind, may be too broad for this discussion. A
alterations of metabolism and growth. Bio more useful concept, for the present, is that the
chemical conditions within the body may result endocrine system includes those restricted areas
in overactivity of an endocrine; oversecretion of of tissues that liberate, into the blood stream,
the parathyroid hormone results in withdrawal specific chemical substances, or hormones,
of calcium from the bones. Failure of develop which have a specific effect on remote tissues.
ment of an endocrine gland, or surgical removal The hormones are secreted and are effective in
(e.g., castration), may have complex effects on such minute amounts that they are believed to
body form and function. Trauma, as to the act similarly to catalysts. For example, epi
hypophysis in skull injury, may result in altered nephrine, the secretion of the adrenal medulla, is
growth patterns (e.g., dwarfism or obesity of normally present in the blood in concentrations
endocrine origin). Alteration attendant on a of one part in one to two billion. Its secretion
physiological state, as of the gonads and other rate may be increased by 20 times. The facts
organs in pregnancy, may initiate a series of that secretion of the hormones must be con
highly complex changes in body form and func tinuous for normal body function, and that
tion. degradation products of certain hormones ap
The effects of removal of certain of the endo- pear in the urine would, however, suggest modi
crines have long been known, although much fications of a purely catalytic action.
time elapsed before the mechanism of action was Although certain of the endocrine glands are
adduced. Castration has been practiced through formed early in development of the embryo of
out recorded history. This, together with the higher forms, it appears that the nervous system
chance removal or failure of development of soon dominates the integration of the proc
other endocrine glands, led to the accumulation esses of the organism. This deduction is phylo-
of data on which carefully planned experi genetically sound because the nervous system be
mental removal was based. The administration comes functionally significant in the evolutionary
of endocrine extracts or grafting of endocrine scale long before specific endocrines put in an
glands to normal or to clinically or experiment appearance. The hypophysis of the chick may be
ally deficient animals is one of the most effective removed surgically on the second day of incuba
tools in endocrine research. In this way it has tion without interfering with the early develop
been possible to assess rates of secretion and ment of the chick. In ontogeny the nervous sys
minimum effective and optimum amounts of tem is well developed before secretory elements
endocrine products in therapy, and gain more of the endocrine glands are differentiated; in
precise knowledge of the relation of the endo- fact, the latter are the last of the coordinating
crines to developmental, physiological, and mechanisms to develop. Despite the fact that
pathological conditions than could be adduced many individual tissues will differentiate in the
from mere removal. absence of nervous connections, their full poten
Chemical, histochemical, and autoradio cies cannot be realized.
graphic analysis has led to identification of the Although chemical regulation may be a more
active principles and synthesis of many hor primitive process than that afforded by the nerv
mones, and to the identification of these chem ous system, a true endocrine system, depending
ical compounds in specific cells of the various as it does upon a circulatory system for distribu
tissues. This has been a significant development, tion of its products, appears to be a late acquisi
because it is no longer necessary to administer tion. Phylogeny offers ample proof of this de
the fresh tissue or crude glandular extracts. Fi duction. The nervous system remains the prime
nally, genetic studies have led to an analysis of necessity in the responses of the organism to its
the role of the endocrines in the development of environment which require rapid mediation.
diverse constitutional types, especially in an Because many of the highly specialized organs
animal like the dog. of the body can be denervated without harming
It is apparent that the importance of the endo their normal function, increasing attention is
crine system goes far beyond a discussion of the being paid to humoral integration of these proc
series of organs as presented by most texts. It is esses. Metabolism is a prime example of a series
a narrow concept to limit it to the ductless glands of processes which depend primarily on chemical
which are distinct entities in the organism. Since integration.
810 Chapter 16. T he E n d o c r in e Sy stem
Importance of Dog in Endocrine Research nificance of the various parts of the gland. Thus
reference will be made to pars distalis, pars tu
The dog is widely used in endocrine research, beralis, pars intermedia, and pars nervosa. The
and the endocrines are undoubtedly involved in embryonic divisions of the hypophysis will be
more systemic conditions than is usually sus considered under the heading of Development.
pected. The endocrine aspects of tissues other
than those organs which are primarily endocrine Gross Anatomy
glands are discussed in a more general manner,
largely because specific information for the dog The hypophysis of the dog is a more or less
is not available. What is said of the more general flattened, ovoid body lying at the base of the
aspects of the dog endocrines is applicable to the brain in a concavity of the basisphenoid, the hy
other domestic animals. pophyseal fossa, which is a part of the sella
turcica. It is attached to the base of the brain by
THE HYPOPHYSIS means of a short hollow stalk, the infundibulum.
The third ventricle of the brain extends through
The hypophysis (Figs. 16-1, 16-2), although the stalk and may be evaginated slightly into the
not essential for life, is undoubtedly one of the substance of the pars nervosa. The infundibu
most important organs of the body (Dandy and lum is directed posteriorly, as is the remaining
Reichert 1925). One of its component parts, the portion of the hypophysis. Thus the long axis of
anterior lobe, has been termed the “master the entire structure lies more or less parallel to
gland” of the body. The common use of the term the floor of the brain.
pituitary gland originated from the belief that From a mid-sagittal section (Fig. 16-2) it can
this gland secreted a substance (Gr. pituita = be seen that the pars nervosa is a solid spherical
to oval outgrowth from the infundibulum. In
phlegm) into the nostrils. The hypophysis is of
color and consistency this portion of the hy
dual origin (epithelial and nervous), and of mani
pophysis is not unlike the brain. The pars distalis
fold function. From the standpoint of morphol
more or less completely surrounds the pars ner
ogy, its control over the growth of the body as
vosa, and is darker and less homogeneous in tex
a whole and the proportionate growth of body
ture. In most cases a small portion of the pars
parts is of particular interest.
nervosa is exposed posteriorly. The pars distalis
is separated from the pars intermedia by the ex
Terminology tensive hypophyseal cleft, or residual lumen of
Rathke’s pouch. The pars intermedia may be
A most confusing terminology has arisen in seen upon close inspection to be a narrow band
connection with this gland. The hypophysis of dark (vascular) tissue on the deep wall of the
(hypophysis cerebri, glandula pituitaria, or pi cleft, closely applied to the surface of the pars
tuitary gland or body) can be divided grossly nervosa. The pars tuberalis, also dark in color,
into two parts or lobes, anterior and posterior, is that portion closely applied to the stalk and
by the hypophyseal cleft, which is of greater de tuber cinereum, and appears to be continuous
velopmental than anatomical significance. On with the pars distalis. The significance of these
this basis the anterior portion would consist of relationships will be noted in discussion of the
the pars distalis (anterior lobe, pars anterior, or development of the hypophysis (Stockard 1941,
pars glandularis) and the relatively insignificant Stigliani, Cipriani, and Del Vivo 1954).
pars tuberalis (pars infundibularis), which sur The sella turcica is the bony structure partly
rounds the infundibulum (hypophyseal stalk). surrounding the hypophysis. Caudally it presents
The posterior portion would consist of the pars a prominent dorsum sellae with its well-devel
nervosa (posterior lobe, pars posterior, or pars oped posterior clinoid processes, but anteriorly
neuralis) and the small pars intermedia lying be there is only a slight bony eminence and poorly
tween the pars nervosa and the hypophyseal developed anterior clinoid processes. The dura
cleft. mater, the outer investment of the brain, is very
The terms anterior and posterior pituitary are intimately associated with the hypophysis and
somewhat ambiguous, particularly since the re the sella. Gross removal of the gland and the
lations in the dog make this human terminology meninges produces artifacts, sectioning in situ
inapplicable. In the following account each por being necessary to demonstrate the correct re
tion of the hypophysis is considered separately, lationships. The dura passes over the posterior
a viewpoint according with the functional sig clinoid processes and sides of the sella to the hy-
T he H y p o p h y s is 811
Ant. c e r e b r a l a.
Ant. i n t e r c o r o t i d
- M i d d l e c e r e b r a l a.
Po n s - B a s i l a r a.
F o r n i x -If;.
I n t e r t h a l a m i c a d h e s i o n - \ $ \ $ & \ ~ ------ C e re b ra l a q u e d u c t
A n te r io r com m issure
Optic chiasm a
■V . ^ \ \
Infundibulum '
NPa r s i n t e r m ed ia_
Hypophyseal c l e f t
' Pars d i s t a l i s >H y p o p h y s i s
\
pophysis, forming an incomplete diaphragma subject to variation between breeds and indi
sellae posteriorly. The dura then splits and viduals. Large dogs tend to have hypophyses
blends with the hypophyseal capsule dorsally as absolutely larger but relatively smaller than
far as the pars tuberalis. The ventral lamina dips those in small dogs. This is true for differences
into the sella and blends with both the hypo based on both age and breed. The hypophysis of
physeal capsule and the bony periosteum of the the female is usually larger than that of the male
sella. Anteriorly the dura leaves the surface of of the same breed and weight, and in pregnancy
the pars distalis at the junction with the pars it enlarges further (Hewitt 1950, Latimer 1941,
tuberalis and joins the dural covering of the White and Foust 1944).
brain. In life there is no subdural space sur
rounding the hypophysis, and the suggestion of Blood Supply
one in postmortem specimens has been shown to
The pars distalis receives several small anterior
be an artifact. The dura splits to form the caver
hypophyseal arteries from the internal carotid
nous and intercavernous venous sinuses lateral
and caudal to the hypophysis. The subarachnoid artery and the various components of the ar
space, also, does not surround the gland. The terial circle of Willis, especially the posterior
communicating arteries (Fig. 16-1). These small
arachnoid, one of the inner investments of the
arteries converge toward the hypophyseal stalk
brain, extends over the pars tuberalis as far as the
and enter the pars distalis. Here they break up
junction with the pars distalis, where it ends in a
into endothelial-lined sinusoidal channels which
blunt process similar to that found at the points
ramify throughout the glandular portion. Thus
of emergence of the roots of the spinal nerves
there are no arteries proper within the substance
(Schwartz 1936).
of the pars distalis. The venous drainage is simi
A minute epithelial structure, termed the
larly arranged; numerous small veins empty into
parahypophysis, is reported to be present in the
the venous circle and basilar plexus. The sinus
majority of cases as a protuberance on the sur
oids of the pars distalis continue into the pars
face of the pars distalis. This is a remnant of the
tuberalis. In addition, this portion of the hy
embryonic connection between the hypophysis
pophysis receives branches from the arterial
and the pharynx. In the brachycephalic and the
circle.
giant breeds the hypophyseal foramen through
The pars nervosa receives its meager blood
the basisphenoid may persist. In puppies a por
supply from the posterior hypophyseal artery,
tion of Rathke’s pouch remains associated with
which is the principal branch of the intercarotid
the nasopharynx, forming an epithelial structure artery. Upon reaching the apex of the pars ner
termed the pharyngeal hypophysis. Its presence
vosa it forms a superficial plexus which is re
has not been confirmed in the adult animal ported to be joined by vessels from the pars tu
(Kingsbury and Roemer 1940).
beralis. Capillaries enter the pars nervosa from
The sella turcica can be identified radiograph- the plexus. Venous drainage is by means of sev
ically at the lowest part of the floor of the cranial
eral veins which enter the cavernous sinuses.
cavity. Its position can be determined approxi
The parahypophysis has been reported to have a
mately in the living specimen as the midpoint of
separate blood supply of dual origin, a posterior
a line through the anterior borders of the vessel from the intercarotid and, on each side, a
zygomatic processes of the temporal bone. The branch from the internal carotid artery.
hypophysis may be removed surgically via a tem The infundibulum is supplied by the plexus of
poral approach. The dog is given hypertonic the pars nervosa, the sinuses of the pars tubera
saline intravenously to shrink the brain, and after lis, and by vessels from the arterial circle. The
the skull is trephined the animal is laid on its pars intermedia is relatively avascular. It re
back to cause the brain to drop away from the ceives branches from the stalk and the pars ner
floor of the cranial cavity. This lateral approach vosa. Although anastomoses have been demon
is possible because of the shallowness of the sella strated between the various components of the
turcica, and is the only method which is success hypophysis, in vivo experiments have cast doubt
ful in the puppy. Hypophysectomy of larger upon the functional value of these connections
dogs has also been accomplished by a buccal ap (Basir 1932, Dandy and Goetsch 1911, Green
proach through the soft palate and basisphenoid 1951, Morato 1939).
(Crowe, Cushing, and Homans 1910, Dandy
and Reichert 1925, Essex and Astarabadi 1953,
Nerves
Markowitz and Archibald 1956).
The size and weight of the hypophysis are The pars distalis receives unmyelinated sym
T he H y p o p h y s is 813
pathetic fibers from the carotid plexus by means period. In one gland, in pregnancy, there were
of numerous branches which radiate toward the 65 per cent acidophils. The proportion of baso
stalk along the hypophyseal vessels. These fibers phils is much lower in dogs than in man. A fourth
arise in the cranial cervical ganglion and have cell type has been described, with granules
their terminations between the epithelial cells. which are neither acidophilic nor basophilic,
A few fibers are reported to enter the pars dis and recently six cell types, including two classes
talis from the hypothalamic nuclei. These origi of basophils and a zeta cell, have been described
nate in the supraoptic, paraventricular, and tu- (Goldberg and Chaikoff 1952, Hartman, Fain,
beral nuclei, traverse the stalk, and terminate and Wolfe 1946, Purves and Griesbach 1957).
around the cells of the pars nervosa (Dandy The proportion of the acidophils to basophils in
1913, Green 1951, Knoche 1953, Yamada, the pars distalis of the most normal dog hy
Ozawa, and Endo 1956). pophyses is about 11 to 1, which corresponds to
The function of the nerves is not clear. It has the widest limit reported for the human. Most
been suggested that certain of these are secre counts in the more abnormal dog forms range
tory. Stimulation of the cervical sympathetic between 20 to 1 and 40 to 1. The basophils are
nerves is reported to elicit secretory activity of reported to become more numerous in proestrus.
the hypophysis. Communicating fibers exist be The pars distalis of the brachycephalic breeds
tween the oculomotor and optic nerves. The frequently shows one or more cysts, some of
stimulus of light via the retina and optic nerve which may be very extensive and lined with cili
upon hypophyseal function has been shown to ated epithelium. There is a low proportion of
be a factor in the reproductive activity (via hy basophils and an excess of connective tissue ar
pophyseal gonadotrophin stimulation) of various ranged in trabeculate fashion (Stockard 1941).
animals and birds. The structure of the pars tuberalis resembles
that of the pars distalis in that it is composed of
Microscopic Anatomy columns of cells separated by blood sinusoids.
However, the cells do not contain granules, but
The hypophysis of the dog presents an ex may form colloid-filled acini. Scattered nests of
tremely variable picture both with regard to the squamous epithelial cells may be present, which
arrangement of its component parts and the cy probably represent inclusions of embryonic
tology of each component. The dachshund pos buccal epithelium. The pars tuberalis extends
sesses one of the most normal hypophyseal farther along the stalk toward the diencephalon
patterns to be found in any of the pure breeds anteriorly than it does posteriorly (Green 1951,
of dogs. The pars distalis almost completely sur Kingsbury and Roemer 1940, Stockard 1941).
rounds the pars nervosa, and it is surrounded by The cells of the pars intermedia are arranged
a delicate capsule which blends with the dura. in columns and may form follicles. Fusion with
The meningeal investment, however, can be the pars nervosa is complete, numerous cell
separated from the pars distalis post mortem, cords protruding into the substance of the ner
leaving a glistening membrane over the surface vous portion. Some isolated islands of intermedia
of the latter (Stockard 1941). cells may be seen. There is no clear-cut histo
The cells of the pars distalis are arranged in logical differentiation that would serve as a basis
columns and small groups, separated by large for distinction between the pars tuberalis and
thin-walled vascular sinusoids. The three cell the pars intermedia. The hypophyseal cleft is
types described from the human pars distalis are extensive, in some cases extending almost com
present: chromophobes, acidophils, and baso pletely around the so-called posterior lobe (pars
phils. The granules of the acidophils and baso intermedia and pars nervosa) (Beato 1935).
phils stain with the acid and basic dyes, respec The pars nervosa is completely unlike the
tively; those of the chromophobes do not stain other components of the hypophysis. Its relative
distinctively (Herring 1908, Smith, Calhoun, avascularity gives it a whitish cast, differentiat
and Reineke 1953, Stockard 1941, Wolfe and ing it sharply from the highly vascular pars dis
Cleveland 1932, Wolfe, Cleveland, and Camp talis. The third ventricle of the brain extends
bell 1933). In a count of 48 glands from dogs of into the infundibulum and is indented slightly
all ages, the proportional cell count was found into the substance of the pars nervosa. Occasion
to be: acidophils 46 per cent, basophils 4 per ally the pars intermedia may invaginate into the
cent, and chromophobes 50 per cent. No age or distal portion of the pars nervosa and form a
sex difference was found, except a tendency for small cavity, a portion of the residual lumen.
chromophobes to decrease during the growth Otherwise the nervous portion is a solid, nearly
814 Chapter 16. T he E n d o c r in e System
spherical structure, almost completely sur slight indentation is made into its substance by
rounded by the other parts of the hypophysis. the extension of the third ventricle of the brain
The connection of the pars nervosa to the floor into the infundibulum. By means of the latter,
of the brain by way of the infundibulum may be the pars nervosa retains its connection with the
seen on sagittal section. Under low magnifica tuber cinereum. Although the pars buccalis and
tion the pars nervosa has a fibrous appearance. pars nervosa arise as separate entities, the two
Higher magnification reveals irregularly shaped parts are apparently dependent on each other
cells with numerous delicate processes. These for their subsequent development, as neither
are specialized neuroglia cells, termed pitui- develops when it is transplanted without a por
cytes. They are of ependymal origin and are not tion of the other (Kingsbury and Roemer 1940,
modified nerve cells (Dandy 1913, Green 1951, White and Foust 1944).
Hagen 1957, Knoche 1953).
Relation of Structure to Function
of the testis in the male (interstitial cell-stimulat (German shepherd, pointer) the hypophysis is
ing hormone, or ICSH). Injection of sex hor essentially normal in its gross and microscopic
mones decreases and castration increases the details, or at least free from major defects. It
gonadotrophic potency of the pars distalis. The may be inferred that the development of typical
role of the basophils is suggested by the fact that canine characteristics is dependent on the nor
castration results in an increase in the number mality of the hypophysis. Correspondingly,
of these cells. Also, few basophils are seen in many of the defects, or deviations from the nor
anestrus, but they increase in size and number in mal pattern, of body form may be associated
proestrus (Evans, Meyer, and Simpson 1933). with structural or functional disorders of the hy
Another hormone associated with the baso pophysis, particularly the pars distalis. It is prob
phils is the adrenocorticotrophic hormone able that some of the unusual characteristics of
(ACTH) which maintains the function of the certain breeds are of hypophyseal origin, and
adrenal cortex, and injection of which causes that their fixed qualities are dependent on the
cortical hypertrophy. The thyrotrophic hor inheritance of a particular histopathologic con
mone (TSH), associated with the acidophils, is dition of the endocrine system. The inheritance
similarly related to the thyroid. Mammotrophic of thyroid type, mentioned in connection with
hormones of the pars distalis are related to duct that gland, is probably intimately related to the
and lobule growth of the mammary gland, and thyrotrophic potency of the hypophysis.
lactogenic hormone (prolactin) is a factor in the Stockard (1941) has investigated the genetic
initiation of lactation. Hypophysectomy abol aspects of structural variation in the hypophysis
ishes lactation and causes atrophy of the mam of the dog and its relation to breed characteris
mary glands. In pregnancy there is an increase tics. The pars distalis of the dachshund has been
in size of the pars distalis coincident with an described as being essentially normal, whereas
increase in the number of chromophobes. that of the Boston terrier is usually cystic and
Other hormones affect metabolism of food otherwise abnormal, the low proportion of baso
stuffs, particularly that of carbohydrates; certain phils possibly being associated with the poor
injected extracts will produce the hyperglycemia breeding behavior. The pars distalis of the first
of pancreatic diabetes of dogs (Gregory, Drager, generation hybrids of these two breeds may be
Tsai, and May 1956). Additional hormones have extremely cystic, yet the dogs are of normal ap
been postulated for the pars distalis; certainly pearance and activity. This suggests that only a
the hypophysis is involved in many complex small amount of functional tissue is required if
body activities, but it may be superfluous to pro the secretion is of normal quality. In the second
pose a separate hormone for each activity (Brull generation hybrids those which physically re
and Louis-Bar 1953). sembled one or the other original parent stock
The pars nervosa contains three hormonal were found to possess the type of hypophysis
substances; one, a pressor principle, vasopressin, typical for that breed. This led Stockard to sug
causes elevation of blood pressure and is anti gest that certain breed and body types may be
diuretic. Another is primarily an antidiuretic attained only in the presence of a typical his
hormone, and the third, oxytocin, causes con topathologic condition of the endocrines. Ge
traction of smooth muscle, especially of that of netic factors are probably of primary impor
the uterus, and also causes hyperglycemia in the tance, the endocrine secretions presumably
dog (Van Dyke 1936). The oxytocic principle, in acting as intermediaries in the expression of
conjunction with estrogens, is apparently effec characters.
tive in stimulating the uterus in parturition. Further evidence of the role.of the pars dis
These hormones are probably elaborated by the talis in morphogenesis has been adduced by the
cells of the paraventricular and supraoptic nu injection of growth hormone into normal dogs
clei. No specific hormones of functional signifi (Putnam, Benedict, and Teel 1929). Growth
cance in mammals have been identified with hormone was injected into dogs of two breeds,
either the pars intermedia or the pars tuberalis. one normal (German shepherd) and the other
achondroplastic (dachshund), for a period of
Relation of Hypophysis to Morphogenesis over a year. In the dachshunds, acromegaly and
gigantism were produced although they retained
Through the mediation of the growth hor their short achondroplastic extremities. There
mone of the pars distalis the hypophysis exerts was overgrowth of the skin, which caused re
a vast influence upon the morphogenetic devel semblance of the head to that of the bloodhound.
opment of the dog. In the more normal breeds In this connection, it is of interest to note that
816 Chapter 16. T he E n d o c r in e Sy stem
certain acromegalic second generation basset growth of the body as a whole by regulating the
hound x bulldog hybrids (Stockard 1941) which proportionate growth of the body parts.
showed excessive overgrowth of the skin had a
high proportion of acidophils (600 to 1), and few THE THYROID GLAND
chromophobes were found in the hypophysis.
These animals resembled the acromegalic St. The term thyroid is derived from the resem
Bernard type, and the pars distalis presented an blance of the human gland to a shield. The Ger
almost perfect picture of the acidophilic adeno man term “Schildrusen” reflects this same
mas associated with human acromegalic gigan origin. Through its output of thyroxine the
tism. Acromegaly was the first disorder of the thyroid gland exerts a vast influence on most of
hypophysis to be recognized as such. It is cor the activities of the body. Loss or non-function
related with an overproduction of the growth of the thyroid results in a general lowering of
hormone in adult life, when skeletal maturity mental and physical activity. Besides having
prevents the generalized gigantism characteris general effects on metabolic function, the thy
tically produced in young animals under such roid has been shown to be related to the develop
conditions. The most marked changes are over ment of form and behavior in the various breeds
growth of the hands, feet, and face. A tumor of of dogs.
acidophils is usually present.
In the growth hormone-treated dachshund, Gross Anatomy
gigantism was manifested by a greatly increased
length of skull and vertebrae, but not of the ex The thyroid gland (Fig. 16-3) most commonly
tremities. All bones were thickened. Body consists of two separate lobes lying lateral to the
weight was twice that of the littermate control. first five to eight tracheal rings. In some dogs,
This, together with probable hypotonicity of the particularly the brachycephalic breeds, a glandu
muscles, caused the dog to assume a plantigrade lar isthmus is found on the ventral surface of the
attitude. The same conditions have been re trachea connecting the caudal poles of the two
ported in the bulldog after continued adminis lateral lobes. A small amount of fibrous tissue,
tration of growth hormone. In this case general probably representing an ontogenetic vestige of
ized splanchnomegaly was noted and lactation the isthmus, is found in others, but most com
was initiated. The failure of the achondroplastic monly no trace of an isthmus can be found. The
extremities of the dachshund to elongate is in two lateral lobes are more or less symmetrical in
dicative of the genetic nature of this deformity. size and shape; there is no constant difference
In the more normal breed of dog, the shepherd, between the mass of the two. In most dogs the
less tendency toward gigantism was noted. The cranial pole of the right lobe lies opposite the
characteristic overgrowth of the skin and en caudal border of the cricoid cartilage of the
largement of the skull were present (Putnam, larynx or the first tracheal ring. The correspond
Benedict, and Teel 1929). ing part of the left lobe usually lies one to three
Stockard (1941) has observed numerous struc tracheal rings caudal to the right. When an isth
tural disharmonies in various breeds of dogs. In mus is present, it lies horizontally, ventral to the
addition to the overgrowth of skin mentioned trachea, and the caudal poles of the two lobes
above, he cites skull achondroplasia and inade are more nearly in the same transverse plane
quate dental accommodation (bulldog), inade (Dye and Maughn 1928, French 1901, Gilmore,
quacy of the optic fossa to accommodate the Venzke, and Foust 1940, Halstead 1896, Mul
eyeball (Pekingese, griffon), and defective breed ligan and Francis 1951, Stockard 1941).
ing behavior (Boston terrier). Certain of these ab The lateral lobes are less intimately applied to
normalities may often be greatly exaggerated by the trachea in the dog than in man. They lie in
hybridization. It is of considerable significance fascia and are as closely attached to the overlying
that the one feature common to all of these indi musculature as to the trachea. The size of the
viduals was an abnormality of the hypophysis. two lobes is variable, the mass of one occasion
Inasmuch as these are breed characteristics and ally being as much as 50 per cent or more greater
are inherited as such, it is possible that the asso than that of the other. More conspicuous differ
ciated hypophyseal abnormality is the primarily ences between subjects are related to age, breed,
inherited character, the attendant structural and geographical origin. In general, proportion
modifications being secondary. In the normal ally larger thyroids are found in younger sub
animal, then, it is apparent that the role of the jects, and in the smaller, especially the brachy
hypophysis in morphogenesis is to control the cephalic, breeds. The thyroids are reported to be
T he T h y r o id G land 817
larger in females than in males, and a normal The thyroid lobes in most dogs can be palpated
transitory enlargement occurs during estrus and on the trachea immediately caudal to the angle
pregnancy. The absolute weight is greater in of the mandible. In short-necked dogs the posi
geographical areas where a relative deficiency tion is proportionally nearer to the sternum than
of iodine exists; in many of these cases, however, in long-necked individuals. Laterally, either lobe
the gland is pathologically enlarged. The normal is related to the sternohyoid, sternothyroid, and
range in thyroid weight in adult dogs is from 40 sternocephalicus muscles. The dorsal border,
to 400 mg./kg. body weight. Weights of 80 to usually thicker than the ventral, is in close prox
1600 mg./kg. have been reported in areas where imity to the carotid sheath carrying the carotid
iodine is deficient, undoubtedly representing, in artery, vagosympathetic nerve trunk, and inter
many cases, subclinical enlargement of the thy nal jugular vein. Closely associated also is the
roid. recurrent laryngeal nerve.
The thyroid lobes lie in the fascial compart
ment composed laterally of the deep lamina of Blood Vessels
the middle cervical fascia covering the overly
ing muscles, and medially of the prevertebral The thyroid is more richly supplied with blood
fascia. Inasmuch as this compartment communi vessels than is almost any other organ. The prin
cates relatively easily with the thoracic cavity, cipal blood supply to either lobe of the thyroid
surgical or other interference, or pathologic is the cranial thyroid artery. This is a short vessel
processes, may lead to widespread infection of that arises from the common carotid artery op
the cervical and thoracic regions. posite the larynx, runs caudally to the thyroid,
Usually two parathyroid bodies are associated and supplies a variable number of branches to
with each thyroid lobe (see The Parathyroid the lobe. One branch usually gives off several
Glands, infra). The “external” parathyroid most twigs that enter at the dorsal border of the
commonly lies in the fascia at the cranial pole of cranial pole, one of which supplies the parathy
the thyroid. It may be entirely separate from the roid body in this region. Several twigs from an
thyroid tissue, or may be indented into the cra other branch enter the ventral border of the lobe
nial pole of the thyroid external to the capsule. In near its middle. One ramus, larger than the rest,
either case, the “external” parathyroid can be usually runs along the ventral border of the lobe
left in situ in thyroidectomy. The “internal” and continues to the thoracic inlet in close asso
parathyroid body lies beneath the thyroid cap ciation with the recurrent nerve. It anastomoses
sule on the medial aspect of the lobe; in some with the caudal thyroid artery when the latter is
cases it is embedded in the thyroid parenchyma present.
(Fig. 16-4). In no case is it possible to remove The caudal thyroid artery, when present, is a
the thyroid without removing the “internal” small vessel which usually arises by a common
parathyroid. The location of the parathyroids trunk with its fellow from the brachiocephalic
varies, and in some cases they are not at corre artery between the origin of the common carotid
sponding positions on the right and the left lobe arteries. It may arise from other vessels in this
of the thyroid. region, and may occasionally be the major supply
The surgical approach to the thyroid under to the thyroid. The vessel courses toward the
pathologic or experimental conditions is by way thyroid lobe in contact with the trachea and the
of a ventral mid-line incision. The mobility of the corresponding border of the esophagus parallel
lobes permits easy separation from their site and to the recurrent nerve. In the middle third of
ligation of the thyroid vessels. Care must be the neck it anastomoses with a branch of the
taken to leave the “external” parathyroids with cranial thyroid artery.
their blood supply intact. Unwitting removal of In puppies given thiouracil both the cranial
these with the thyroid lobes in early experimental and the caudal thyroid arteries were developed
work led to the faulty conclusion that thyroid to an extraordinary degree. Each was as large as
ectomy was incompatible with life (Dye and the common carotid artery and transmitted a
Maughn 1928, French 1901, Halstead 1896). pulse which was palpable externally.
Functional “thyroidectomy” can now be per Beneath the fibrous capsule of the thyroid lobe
formed with the use of thiourea compounds the vessels form a vascular plexus. This ulti
which inhibit the production of the thyroid mately gives rise to capillaries which form a
hormone, or by administration of radioactive complete network about each follicle, the cap
iodine which accumulates in the thyroid tissue illary endothelium being contiguous with the
and destroys it. follicular epithelium. Numerous arteriovenous
818 Chapter 16 T he E n d o c r in e S ystem
■Thyroid c a r t i l a g e
Thyro id branch of
C ra nia / l a r y n g e o i rt. ^
M. c r ic o th y r o id e u s
M. S t e r n o t h jr o id e u s
C r a r t h q r o i d a.r V.-
Cauda I la r y ng ea I n.
Pa r o t h u r a i d g l a n d -
P B t a t h y r o i d g la n d
M Sternohyoideus
C a u d a l t h y r o i d v.---------
V th y ro id e a ima
M s t e r n c c e p h a i i c u s ■-------
Caud th y ro id v.
In t ju g u la r v Caud- th y r o id O-
Common c a r o t i a a. -- - - Int. j u g u l a r v
with histologic features typical of the breed. in that of the French bulldog, Boston terrier, and
Follicle diameter ranges from 30 to 160 microns, Pekingese. The Brussells griffon and Maltese
with epithelial height from 8 to 20 microns. Two poodle, with flat faces and monkey-like heads,
types of cells, colloid and chief, are found in the have thyroid glands histologically similar to
follicles, but these may be functionally the same. that of the fetal or newborn puppy. These have
Colloid is present in about 95 per cent of the many very small follicles and excessive extra-
follicles, with varying degrees of colloid vacuoli follicular tissue of epithelial character.
zation. In general, the larger the follicle, the
lower the epithelial height. In the fetus many Development
undifferentiated cells with numerous mitoses are
found. Numerous small follicles are found at The thyroid primordium in the dog fetus is a
birth, all of which contain colloid. The average thickened area of epithelium in the mid-pharyn
follicle size increases from about 30 microns at geal floor extending to the caudal limit of the
birth to 75 microns at 16 weeks. After this age aortic sac. Cells bud off from this plate causing it
there is little change in the histologic pattern to extend lateral to the aortic sac and project
(Venzke 1940). convexly into the pharyngeal cavity. The attach
In addition to the follicular cells, large and ment to the pharynx becomes narrowed to a
small groups of epithelial cells with no distinct slender stalk consisting of a layer of cells sur
lumina are found between the follicles. These rounding a potential or actual lumen. A vestige
have been variously termed interfollicular, para of this thyroid stalk may persist as the thyroglos-
follicular, or parenchymatous cells. They are sal duct. Small groups of cells may become de
frequently found in the follicle wall and are tached from the developing thyroid mass and
believed to be concerned with the formation of descend with the aortic sac. This explains the
new follicles. By alternately leaving the follicle presence of thyroid material within the peri
and entering into new follicle formation, they aortic fat bodies. This accessory thyroid material
maintain a cycle of follicular metamorphosis is found along the aorta of all dog embryos and
(Zechel 1931). It is possible that a particular in about 50 per cent of adult dogs. The thyroid
follicle is able to function only a short time. The expands laterally and fuses with the ultimobran-
interfollicular cells are most common in the chial bodies which arise from the fourth
thyroids of young dogs and are rare in old sub branchial pouch. These aid in the formation of
jects. These cells are more numerous in dogs the thyroid parenchyma. The interfollicular cells
with poorly developed thyroids, such as the probably have their origin from the ultimobran-
Boston terrier, than in those with more histo chial bodies. An isthmus, usually transitory, is
logically normal thyroids. They are also more formed from the medial portion of the thyroid
numerous in areas of regenerating thyroid tissue plate. The isthmus persists in animals living in
(Nonidez 1932a and 1932b, Stockard 1941, areas of iodine deficiency, and was found to be
Vicari 1932 and 1937). well developed in puppies given thiouracil. Fol
Stockard (1941) emphasizes that the thyroids licles are formed by breaking up of the cell cords
of mongrel dogs are extremely variable and of the thyroid plate into spheres whose walls
are not suited for reliable histologic studies. represent the original surface of the plate. In
The dolichocephalic breeds in general possess this period of rapid expansion and breaking up
histologically normal thyroids, using the normal into follicles, groups of cells may become isolated
human thyroid as a basis of comparison. In the and form accessory thyroid tissue.
German shepherd, for example, the follicles are The follicles of the fetal thyroid, solid at first,
large and regular, with moderately active epi contain colloid at birth. Areas of relatively un
thelium, as determined by cell height and the differentiated cells remain, giving rise to new
presence of secretion droplets. Nests of inter follicles later in the normal cycle of follicular de
follicular cells are found in the sparse extrafollic- struction and regeneration. After pathologic or
ular tissue. The English bulldog, as a representa experimental destruction of a portion of the
tive of the brachycephalic breeds, shows a most gland, areas of regeneration are found which are
distorted histologic picture. Characteristic are similar to the state observed in the fetus (Godwin
the small, irregular, almost adenomatoid follicles 1936, 1937a).
suggestive of a demoralization of follicular de
velopment. There is an excessive amount of ex- Accessory Thyroid Tissue
trafollicular tissue. The epithelium is active, but
is not suggestive of the hyperthyroidism evident The origin and occurrence of thyroid material
T he T h y r o id G la n d 821
within the periaortic fat bodies has been men related to reproductive function (Halstead 1896,
tioned. Three fat bodies are uniformly present Marine 1932).
at the base of the aorta. In 50 per cent or more of Various patterns of behavior which are typical
individuals these contain small red-brown bod of certain breeds of dogs may be related to thy
ies of typical thyroid tissue ranging in size from roid activity. The German shepherd is a highly
that of a pinpoint to 1 cm. or more in diameter. active dog which reacts strongly to minimal
Other bodies are found in the mediastinum and stimuli. The basset hound is a more lethargic
pericardium, pedunculated or sessile, and en type of animal which reacts less strongly to ex
veloped by serous membrane. These are sup traneous stimuli but is less easily distracted from
plied with blood vessels from the brachioce reacting to an effective stimulus. This is reflected
phalic trunk or by pericardial branches of the in the staying qualities of the latter in the hunt.
internal thoracic artery. Occasionally thyroid tis Crosses of these two breeds show intermediate
sue may be found in the abdomen of normal behavior (Stockard 1941). In all of these animals
dogs (Godwin 1936, Swarts and Thompson activity is associated with the thyroid pattern
1911). and can be reversed or accentuated by thyroid
Accessory thyroids are commonly found dis ectomy or by thyroid therapy. The rate of thy
tributed in the cervical connective tissue from roxine secretion is governed by the thyrotrophic
the region of the larynx to the thoracic cavity. hormone of the hypophysis. Lack of iodine, a
These may number from 2 to 10 or more. In necessary constituent of the thyroid hormone,
cases of thyroid hyperplasia, or following thy increases the secretion of thyrotrophin so that
roidectomy, the accessory thyroids enlarge to the thyroid attempts to compensate for the poor
several times normal size. The effectiveness of quality of its secretion by hypertrophy and hy
thyroidectomy depends on the relative absence perplasia. Injections of thyroxine decrease the
of accessory thyroid material. This is not a fac thyrotrophic potency of the hypophysis. Iodin-
tor when thyroid-inhibiting drugs are given, ated casein (thyroprotein) has an effect on the
since these accessory thyroids are likewise in body similar to that of thyroxine (Borgman and
hibited. Reineke 1949, Smith, Calhoun, and Reineke
1953).
Relation of Structure to Function
Relation of Thyroid to Morphogenesis
In the adult animal the thyroid gland, through
its output of thyroxine, functions in maintaining In addition to its effect on metabolism, the
the proper rate of metabolism of the body tis thyroid in the young animal has a special rela
sues. Thyroxine probably acts as a catalyst in the tion to morphogenesis which is of particular in
oxidative processes of the tissues. The rate of terest in the development of diverse dog breeds.
output varies to meet the changing require The thyroid exerts a vast influence upon growth.
ments of the individual. Thyroid deficiency in Various parts of the body are affected unequally.
the adult, whether produced experimentally or In the process of normal growth from birth to
through pathologic processes, produces a myxe maturity, the ratio of body length and of bone
dematous condition with puffiness, drying, and length and volume to the body weight decreases;
thickening of the skin, falling-out of the hair, visceral and skeletal weights are proportionally
muscular weakness, mental sluggishness, and decreased, and muscular weight is proportion
lowering of the metabolism. Hernia may occur ally increased. The relative size of the head de
as a result of weakening of the abdominal mus creases, and body weight is both absolutely and
cles. Certain of these conditions which occur in relatively increased per unit of body length. The
the presumably normal process of aging may be relative increase in length of the long bones nor
a manifestation of a lowering of the functional mally occurs at a faster rate than that in their
activity of the thyroid. diameter.
The degree of alteration of body structure and Thyroidectomy in the young animal reverses
function depends on the age at which thyroid these relationships. In general, the body is af
activity becomes deficient. Cretinism, which fected more than the skeleton, and different
develops in thyroid-deficient young, is a more bones are not affected to the same degree. The
severe condition than the myxedema which is ratio of body length and of bone length and vol
manifested in older animals. Physical growth ume to body weight is greater in thyroidecto-
and mental development are both markedly re mized animals than in normal littermates, al
tarded. The function of the thyroid gland is also though the absolute values are considerably less.
822 Chapter 16. T he E n d o c r in e System
Visceral and skeletal weights are relatively vention of goiter has made possible considerable
greater, and muscular weight is proportionally reduction in these figures.
less. The head remains relatively large. The rela Symptoms of hypothyroidism may occur with
tive increase in diameter of the long bones is ac out enlargement of the gland in simple goiter;
centuated in the cretin because subperiosteal in this case the active tissue of the gland is re
deposition of bone is not altered while growth is placed by inactive tissue. In colloid or paren
retarded at the epiphyses. Neither is the forma chymatous goiter, which is the most common
tion of membrane bone in the skull interfered type in dogs, the gland attempts to compensate
with, although increase in length of the skull for iodine deficiency by overgrowth. Goitrous
bones is retarded. This causes the head to tend thyroids weighing as much as 74 grains have
toward the brachycephalic type. The skull be been recorded. This type of goiter is found in
comes broader from before backward, and prog dogs given thyroid-inhibiting drugs. In congeni
nathism of the lower jaw is apparent. Although tal goiter of puppies the thyroid may be so large
the mandible is shortened in proportion to the as to interfere with parturition. In these cases
upper jaw, the shortening of the basicranium the lateral lobes and isthmus form an indistin
gives the mandible an undershot appearance guishable mass in the caudal cervical region.
(Dye and Maughn 1928, Halstead 1896). Cretinism is a manifestation of failure of the thy
Stockard (1941) has shown that the physical roid to develop and is most common in the young
characteristics of certain breeds of dogs are asso of hypothyroid parents.
ciated with the histologic pattern of the thyroid. Hyperthyroidism may take the form of malig
The dolichocephalic breeds, which most closely nant goiter or, more rarely, exophthalmic goiter.
approximate the primitive type of dog, have a Malignant neoplasms of the thyroid of old dogs
histologically normal thyroid, whereas the are common. They may metastasize to various
brachycephalic breeds in general present a dis internal organs, the lungs being the principal or
torted thyroid pattern. First generation hybrids gans affected. Normal accessory thyroid masses,
of these two types usually have uniformly nor especially those in the periaortic fat bodies, may
mal thyroids and are good physical types. In be confused with metastatic nodules on gross
second generation crosses the thyroid condition examination (Davis 1938, Marine 1932, McClel
of the animal can be freely predicted from the land 1941).
physical type, and vice versa. Behavior of these
hybrid animals can also be related to the nature THE PARATHYROID GLANDS
of the thyroid. Many of the second generation
hybrids present bizarre physical characters as The parathyroid glands, so called because of
sociated with abnormal endocrines and abnor their anatomical relation to the thyroid, were
mal behavior. early believed to be remnants of embryonic thy
roid tissue. The term, “thyroid glandules,” and
Pathology the German name, “Epitheliokorperchen,”
(small epithelial bodies) also suggest a lack of
The thyroid gland of the dog is subject to all understanding of the significance of these struc
of the pathologic conditions known in man, and tures at the time the terms were applied. The
for this reason the dog has been extensively uti only relation of the parathyroids to the thyroid
lized in such studies. It had early been observed is anatomical; they are distinct entities in both
that wherever goiter was endemic in man the origin and function (French 1901). The impor
same condition was common in dogs of the re tance of the parathyroid secretion in the well
gion. The incidence of goiter in dogs in inland being of the animal is indicated by the fact that
cities (Cleveland, Chicago) has been reported as abrupt total extirpation of all parathyroid tissue
high as 90 per cent, whereas in the same period is almost invariably fatal in dogs.
(early 20th century) the incidence in coastal
cities (Baltimore) was less than 10 per cent. At Gross Anatomy
one time it was rare to find a dog with normal
thyroids in certain districts of Switzerland and The parathyroid glands (Figs. 16-3, 16-4)
of other European countries. It has been sug are small oval bodies associated with the thyroid.
gested above that the data on thyroid size of They vary in number and size; usually there are
dogs from such areas are indicative of a relatively two principal parathyroids on each side. Al
high incidence of subclinical thyroid enlarge though they vary also in topographic relations
ment. Knowledge of the role of iodine in the pre and intimacy of contact with the thyroid lobe,
T he P a r a t h y r o id G la nds 823
they can be distinguished as “external” and “in parathyroidectomy difficult, or in some cases im
ternal” on the basis of their position with refer possible. Accessory parathyroids may be found
ence to the capsule of the thyroid, and whether within the thyroid lobe, or externally in the
they are lateral or medial to the associated lobe region of the larynx, the carotid sheath, the an
of the thyroid. On an embryologic basis they are terior mediastinum, and within or associated
designated as parathyroids III or IV because with the thymus (Godwin 1937b).
they take origin from the third and fourth pha
ryngeal pouches (French 1901, Godwin 1936 Microscopic Anatomy
and 1937b, Vicari 1932).
Parathyroid III, commonly referred to as “ex The parathyroids are homogeneous, compact
ternal,” is a flattened oval body, about 2 to 5 masses of epithelial cells arranged in irregular
mm. long, most commonly found in the connec cords. The gland is very vascular, with sparse
tive tissue about the cranial pole of the thyroid connective tissue. Nearly all of the cells are of the
lobe. Exceptionally this gland may be as large as “chief cell” type, the type believed to be the
1 cm. or less than 1 mm. in length. It may be source of the parathyroid hormone. A few larger,
indented into the thyroid lobe, but is always ex granular, bluish-staining cells are scattered irreg
ternal to the capsule of the latter. Frequently it ularly or collected in groups. Oxyphil (acidophil)
is found lateral to the thyroid lobe, occasionally cells described for the human parathyroid are
as far caudally as the caudal pole of the thyroid. absent in the dog. The arrangement of the cells
In general the size of the external parathyroid is not the same in all of the parathyroid bodies
varies in proportion to the size of the dog. In from one dog. Differences may also be noted be
the giant breeds it is large and plump; in the toy tween the glands of different members of the
breeds it is minute in size. In dolichocephalic same breed or between dogs of different breeds.
dogs it is less plump. Although it is not practical Few constant differences which are character
to estimate the total amount of parathyroid tis istic of a breed can be found, but a study of vari
sue present in the dog, the external parathyroids ous glands suggests differences in functional
average about 5 mg. in weight in large dogs activity at least.
(Mulligan and Francis 1951). Various derangements of structure, such as
Parathyroid IV, commonly referred to as cysts and embryonic rests, are found, particu
“internal,” is a small mass of tissue of inconstant larly in poorly formed second generation hybrids
size and shape. It is generally, smaller, rounder, of certain breeds (Boston terrier x dachshund,
and flatter than the external parathyroid. It is basset hound x bulldog). Because of other con
usually found on the tracheal surface of the thy spicuous endocrine disturbances in these ani
roid lobe, beneath the capsule, but occasionally mals, it is not possible to state whether parathy
it may be found on the lateral surface. It may be roid distortions have a causal relationship to the
found deep within the thyroid parenchyma. physical deformities found. The frequency of
Parathyroid tissue may be distinguished from poor tooth structure in these mongrels suggests
thyroid by its darker, more homogeneous con a possibility of parathyroid influence (Stockard
sistency (Godwin 1937b). 1941). In the basset hound and bulldog it was
The blood supply to the external parathyroid held probable that the modifications of the skele
is most commonly a separate branch from the ton and overgrowth of skin were related to the
cranial thyroid artery. In some cases only a num parathyroids.
ber of smaller twigs more intimately associated
with the thyroid are supplied to the parathyroid. Development
If they are supplied by a separate branch, the
external parathyroids with their blood supply Parathyroid III can first be identified in the
can be left intact when the thyroid is removed. 7.5 mm. dog embryo as a thickening on the dorso-
The internal parathyroids are supplied by minute anterior face of the third pharyngeal pouch. A
ramifications of the arterial supply to the thyroid duct is formed connecting the parathyroid with
parenchyma. As stated earlier, it is impossible to the pharynx. The parathyroid primordium is
remove the thyroid lobe without removing the associated with that of thymus III, and in separa
internal parathyroid. The parathyroids have tion of these structures accessory parathyroids
their venous and lymphatic drainage in common may be formed. This would explain the presence
with the thyroid. of parathyroid tissue within or associated with
A variable number of accessory parathyroids the thymus. Clusters of cells sprout away from
are found with such frequency as to make total the developing mass and may give rise to the
824 Chapter 16. T he E n d o c r in e Sy stem
other accessory parathyroids in the extensive that these are effective only if small amounts of
growth shifts which occur in the differentiation parathyroid tissue are present.
of the organs derived from structures in this In the normal animal parathormone maintains
region. The parathyroid shifts medially, ven an exchange between serum calcium and the
trally, and caudally, and loses contact with the calcium stores of the body. The first effect of an
pharynx. The principal parathyroid III, after excess of hormone is a resorption of the trabec
separating from thymus III, usually comes to lie ulae in the marrow cavities of the hollow bones
on the lateral aspect of the thyroid near its cranial and a rise in serum calcium (Jaffe and Bodansky
pole. Cysts are frequently associated with the old 1930, Learner 1929). Conversely, the first effect
duct. of a deficiency of the hormone is a fall in serum
Parathyroid IV, arising from the fourth pha calcium. Tetany occurs when the serum calcium
ryngeal pouch, is at first connected to the level falls to about one-half the normal value. In
pharynx by an open duct, which later becomes a about 5 or 6 per cent of dogs enough accessory
solid cord of cells. It undergoes a period of fusion parathyroid tissue is present to maintain life after
and inclusion with the thyroid. Fragments may parathyroidectomy. In all others death even
give rise to accessory parathyroids within the tually occurs by tetany-induced asphyxia. The
thyroid parenchyma. The duct may become proportion of dogs surviving can be increased by
cystic. previous partial parathyroidectomy or by tem
In man the positions of parathyroids III and porary feeding of calcium to allow a compensa
IV are the reverse of those in the dog, parathy tory mechanism to develop, probably by
roid IV being more cranial. Their developmental hyperplasia of accessory parathyroid tissue.
history, however, is similar (Godwin 1937b). The effects of parathyroidectomy are more
serious in puppies than in older dogs. Fatal tetany
in puppies may be induced by ligation of the
Physiology cranial thyroid artery. In the adult one functional
parathyroid body is adequate to meet the normal
The chief function of the parathyroid, through demands of the body, but tetany may occur in a
its output of parathormone, is to maintain opti dog with intact parathyroids, especially during
mum calcium and phosphorus levels of the blood, times of high calcium demand (pregnancy and
especially during growth, reproduction, and lactation). Growth, reproduction, pregnancy,
lactation, when there is an increased calcium de and lactation excite the tetanic state after para
mand. Although abrupt total removal usually thyroidectomy, but this may be avoided if the
results in fatal tetany, dogs can be maintained in serum calcium level is maintained by proper
an apparently normal state after parathyroidec feeding (Dragstedt, Sudan, and Phillips 1924,
tomy with substances not chemically related to Kozelka, Hart, and Bohstedt 1933). Parathor
parathormone. Tetany is a neuromuscular condi mone injections are, of course, also effective, as
tion directly related to the low calcium level are grafts of parathyroid tissue if the grafts are
brought about by parathyroidectomy. It is char successful (Shambaugh 1936). Non-fatal tetany
acterized by high body temperature which so in the pregnant bitch almost invariably results in
increases the respiratory rate that the carbon abortion through asphyxia of the pups. Cataracts
dioxide level of the blood falls so low that the are frequent, and dogs are more susceptible to
blood becomes more alkaline. This interferes infection after parathyroidectomy.
with the ionization of the already low amount of
calcium in the blood. Because of the lack of the Pathology
normal sedative action of calcium on the nervous
system, the animal becomes hyperexcitable and The similarity of the tetany syndrome associ
enters an epileptiform state. The muscles go into ated with the parturient state in the intact dog
tetanic contraction; death is usually due to con to that following parathyroidectomy suggests
tinuous contraction of the respiratory muscles further investigation of the role of the parathy
which results in asphyxia (Bensley 1947, Marine roid in the former condition.
1914, Reed, Lackey, and Payte 1928). Dogs are also subject to spontaneous osteitis
Vitamin D has been demonstrated to maintain fibrosa, a condition similar to that found in man.
the serum calcium level and permit normal bone It is characterized by an excessive mobilization
growth in puppies after parathyroidectomy. of calcium at the expense of the skeleton. The
Tetany may be prevented by the administration bones are thin, curved, and shortened. The mar
of calcium salts, although it has been claimed row is replaced by fibrous tissue. Existing bone is
T he P a r a t h y r o id G la n d s 825
P r e v e r t e b r a l fa scia Thtjroid g l a n d
M, longus col 11 P a r a t h y r o i d g l a n d (IV)
Postcava i
P h re n ic o o b d o m in o f a
S u p r a r e n a l o. 1 , c e l i a c a.
I
P h r e n i c b r . o f P h r e m c o o b d o m m o l a .s /A o r t a
' I ' i /
' I
Diaphragm-
Cranial
Kidnei j m e s e n t e r i c a.
Left adrenal
gland
Abdominal
b r o n c h e s of * -
Phremcaabdomi nal aw. - "Renal a. v v
U reter
I/
R iq h t t e s t i c u l a r a .w . ■*
M. psoas minor
Fic.. 16-5, Th» adrenal glands. ventral aspecl
82 6 Chapter 16. T he E n d o c r in e System
resorbed and transformed, the haversian canals The left adrenal lies further caudally, corre
being invaded by fibrous processes. There is en sponding to the position of the left kidney. It is
largement of both the upper and lower jaws, medial to the cranial pole of the kidney and is
together with distortion of the cranial bones. separated from the postcava and aorta by fatty
The syndrome can be produced on a calcium- tissue. The left adrenal is markedly constricted
poor diet, but a more specific condition is pro in its central portion, the cranial half being broad
duced by an excess of parathormone. This con and flat. The caudal portion of each of the
dition is found in puppies, suggesting a con adrenals tends to be relatively long and narrow,
genital origin, or in senile dogs, suggesting a and in cross section is more or less oval.
deranged metabolism. Experimentally, the long Cranially the ventral surface of each adrenal
time injection of parathormone in dogs produces is covered by peritoneum, the organs otherwise
progressive decalcification and resorption of being surrounded by loose connective tissue and
bone with fibrous replacement. Other features fat. The caudal pole of each adrenal is buried in
of osteitis fibrosa, with deformities similar to perirenal fat. In fat subjects the cranial half of
those in clinical cases, are present. Calcium may the left adrenal may be mistaken for the entire
be deposited in the soft tissues, particularly in gland, so completely is the caudal pole hidden
the kidney, liver, lungs, heart, and arteries. Uri from view (Baker 1937, Randolph 1950, Stock
nary calculi may be formed (Leberman 1940). ard 1941).
The adrenal is enclosed in a fibrous capsule,
which in turn is invested by the loose connective
THE ADRENAL GLANDS
tissue and fat. On the surface of the capsule are
found nerves and ganglia from adjacent plexuses,
Like the hypophysis, the adrenal glands are
but their relationship to the adrenal is superficial.
composed of two major divisions which are
From the capsule numerous septa penetrate the
separate entities both developmentally and func
tionally. Because of their structural relations, the cortex, dividing it into oval or oblong compart
outer cortex and inner medulla of the adrenal ments and serving as a framework for the gland
have usually been considered parts of one organ. parenchyma. The medulla is similarly divided
In some of the lower animals the cortex and into cell groups by these septa.
On section of the adrenal the cortex and me
medulla are anatomically separate bodies. The
term adrenal with reference to the gland as found dulla can be distinguished grossly by the color
in domestic animals is preferable to the human and consistency of the two parts. The cortex is
term suprarenal because of the difference that firm and yellowish; the medulla is softer and has
posture makes in the relative position of the a brown pigmentation. In general, the cortex is
glands. The importance of the adrenal in the of uniform thickness; hence the shape of the
economy of the animal is shown by the fact that medulla conforms to that of the gland. Fre
the cortex is essential for life, and the medulla, quently, however, evaginations of one part into
although not indispensable, is apparently of the other can be seen with the naked eye. Rarely,
particular significance in the mechanism by the medulla may extend entirely to the capsule
which the animal responds in emergencies. at one point.
The position of the adrenals can be best ascer
tained externally by determining the position of
Gross Anatomy the kidneys, which is possible in many cases by
palpation. Because of location of the adrenals
The adrenal glands (Fig. 16-5) of the dog are medial and slightly cranial to the cranial pole of
generally flattened, bilobed organs situated the corresponding kidney, the right adrenal is
cranial and medial to the kidneys on either side located opposite the last thoracic or first lumbar
of the vertebral column. They are asymmetrical vertebra, lying at the level of the cranial mes
in both position and shape. The right adrenal enteric artery. The left lies opposite the second
usually lies between the medial surface of the lumbar vertebra, along the aorta between the
cranial pole of the right kidney and the lateral roots of the cranial mesenteric and renal arteries.
aspect of the postcava. A portion of the cranial The right adrenal is in contact with the right crus
pole of the right adrenal is directed sharply later of the diaphragm.
ally and backward, giving it a distinct hook, or The preferred experimental surgical approach
comma-shape, and is separated from the right is by a paralumbar incision. The relations of the
crus of the diaphragm by a small amount of fatty diaphragm limit the anterior extent of such an
tissue. incision. Because of the rich blood supply to the
T he A d r en a l G la n d s 82 7
adrenals it is necessary to tie off the principal groups and blood vessels of its three zones. Other
blood vessels of each gland before it is excised. fibers from the capsular plexus, some of which
may be myelinated as they pass through the
Blood Vessels and Nerves cortex, are distributed to the cell groups of the
medulla. Here they are arranged in dense
The adrenal develops in a highly vascular area plexuses about the chromaffin cells and blood
and probably has a richer blood supply, in pro vessels. The medulla and capsule may contain
portion to its size, than any other organ except ganglion cells which may not, for the most part,
the thyroid. The adrenal receives one or more be functionally related to the adrenal but may
branches from all of the major arterial trunks that represent portions of the celiac plexus which
pass near it. Thus the adrenal arteries may arise were carried in by the ingrowing medullary cells
directly from the aorta (middle adrenal artery), (Alpert 1931).
and from the phrenic (cranial adrenal artery), Lymphoid nodules are sometimes found in the
renal (caudal adrenal artery), accessory phrenic medulla, and irregular groups of lymphoid cells
and lumbar arteries, less frequently from the may be found in the cortex. Lymph vessels have
cranial mesenteric or celiac artery. In most not been demonstrated except for those about
cases the cranial portion of the gland is supplied the larger veins.
by the phrenicoabdominal artery; the caudal
pole is supplied by the other vessels named. Microscopic Anatomy
In the capsule the adrenal arteries form a
poorly defined plexus from which the entire The outer layer of the capsule of the adrenal is
gland is supplied. From the plexus the blood is made up of white fibrous tissue beneath which is
directed through three different channels to the a smooth inner layer of reticular connective tis
capsule, cortex, and medulla. The small capsular sue. Within the capsule are found connective
arteries form an irregular capillary network in tissue cells and some smooth muscle fibers, as
the substance of the capsule and empty into the well as blood vessels and nerves. From the inner
deeper capsular venous plexus. The cortical layer of the capsule septa pass in toward the
arteries form a capillary network about the cell center of the gland. Smaller septa extending both
columns of the cortex and empty into the venous from the capsule and from the larger septa
tree of the adrenal at the periphery of the me further subdivide the parenchyma of the adrenal
dulla. Each cortical cell is in contact with one to and support individual cell columns of the cortex
four capillaries. The medullary arteries run from and cell groups of the medulla. The connective
the capsule through the cortex and form a capil tissue framework of the cortex is much denser
lary plexus in the medulla. Blood sinuses are than that of the medulla, producing a sharp line
formed in the medulla and the inner zone of the of demarcation between the two parts of the
cortex. Blood is returned from the medulla either gland.
by way of the venous tree or by the central veins Cortex. It is customary to divide the adrenal
of the adrenal. The larger branches of the venous cortex into three distinct zones or layers, regard
tree, as well as satellite veins of the adrenal less of how poorly defined these layers may be.
arteries, empty into the adrenal vein. The right From the periphery inward these are termed the
adrenal vein joins the postcava, the left joins the zona glomerulosa, zona fasciculata, and zona
left renal vein. Valves are present in the capsular reticularis. Although these three zones can be
veins, but none are found in the large branches recognized in the dog adrenal, ordinary methods
of the venous tree. Arteriovenous anastomoses do not always reveal a sharp distinction between
have been described in the loose connective adjacent zones. The peripheral zona glomeru
tissues about the adrenal. This suggests a means losa, representing about 25 per cent of the
of control of blood flow through the gland cortex, is composed of coiled cell columns which
(Brondi and Castorina 1953). have a sigmoid or U-shaped arrangement next to
The adrenals are richly innervated, mainly by the capsule. For this reason, in the dog adrenal,
fibers from the splanchnic nerves, but also by it has more aptly been termed the zona arcuata.
fibers from the celiac ganglion and the first three The middle zona fasciculata, representing about
or four ganglia of the abdominal sympathetic 60 per cent of the cortex, is arranged in long
chain. These fibers form a plexus in the capsule, anastomosing columns of cells the long axes of
consisting of both medullated and non-medul- which run at right angles to the capsule. Adja
lated nerves. Fibers from this plexus enter the cent columns are separated by capillaries, and
cortex and form small plexuses around the cell the cells of this zone, especially, contain large
828 Chapter 16. T he E n d o c r in e S ystem
amounts of lipoid material. The cells of the inner tively late in fetal development that the defini
zona reticularis, representing the remaining 15 tive relationships of the two are obtained. The
per cent, are arranged in small groups sur cortex, which is the first to develop, arises from a
rounded by capillaries. Portions of this zone may budding of peritoneal mesothelial cells between
be evaginated into or even form islands of corti the dorsal mesentery and the genital ridge.
cal substance in the medulla, or the medulla may These cortical primordia soon become highly
extend into the zona reticularis in places. In spite vascularized and relatively large. The definitive
of this irregularity, a sharp distinction can always structure of the cortex is not attained until after
be made between cortical and medullary sub birth (Randolph 1950).
stances either in fresh sections or in stained prep The chromaffin cells of the medulla arise from
arations (Bennett 1940, Gruenwald and Konikov the neural crest and form masses of cells which
1944, Nicander 1952, Smith, Calhoun, and invade the already formed cortical primordium.
Reineke 1953). This process is probably not complete until after
In general the dolichocephalic breeds have birth, nor is the process always complete in an
an adrenal cortex showing the characteristic individual. Frequently invaginations or even is
arrangement of the three layers. In the brachy- lands of medullary substance may appear in the
cephalic breeds the two inner zones are clearly cortex, or groups of cortical cells may be carried
distinguishable from the outer but are not distin in with the ingrowing medullary cell masses. Al
guishable from each other. Some hybrid animals though a delicate capsule is found about the cor
show extreme disorganization of all the com tical primordium, the dense capsule of the com
ponents of the gland. These findings are in con bined gland is not formed until late in embryonic
formity with Stockard’s (1941) report on the life.
other endocrines. The close association of the cortex and the
Medulla. In most dogs the medulla represents gonads in development provides the basis for
10 to 20 per cent of the entire adrenal. The cells one explanation of the functional interrelation
of the medulla are arranged in small round or ships of these organs in the individual. The simi
oval groups separated by loose connective tissue lar origin of the medullary cells and the sym
and blood vessels. Nerve cells are found less fre pathetic ganglion cells also suggests an explana
quently in the medulla of the dog than in most tion of the sympathomimetic action of the
other animals. The typical brown staining reac medulla, although the two originate from dif
tion given by medullary cells with the salts of ferent types of neural crest cells. Groups of
chromium has given rise to use of the term chromaffin tissue which do not become incorpo
chromaffin cell. This reaction is considered to be rated with the medullary cell masses may form
indicative of the presence in these cells of the paraganglia about the abdominal aorta. These
medullary secretion, epinephrine, the intensity apparently have a function similar to that of the
of staining reflecting the rate of secretion. Cer adrenal medulla, and their presence may account
tain aggregates of cells having the same reaction for the minor consequences attending removal
have been described along with the adrenal of the medulla. These masses of chromaffin tis
medulla as belonging to the chromaffin system. sues include the aortic and carotid bodies and
In the dog there is a large paraganglion situated the chain of paraganglia.
dorsal to and extending cephalad of the bifurca
tion of the abdominal aorta. It can be demon Structure in Relation to Function
strated by covering that region with cotton
soaked in potassium bichromate solution, which Cortex. Dogs do not live longer than 15 days
renders the paraganglion brownish. Chromaffin after complete bilateral adrenalectomy. Preg
cells are also found scattered in the sympathetic nant animals or those in estrus or pseudopreg
ganglia and in a number of organs. It is assumed nancy survive longer than do other females or
that they have a function similar to that of the male animals. It has been shown that progester
adrenal medulla, but the advisability of using the one, the hormone of the corpus luteum, is the
term chromaffin system has been questioned principle responsible for extending the life of
(Malmejac, Neverre, and Bianchi 1957). these animals. An interrelationship of the adrenal
cortex and the ovary is indicated by an increase
Development in the weight of the adrenal in estrus. The adrenal
weight is relatively increased in pregnancy if the
The cortex and the medulla of the adrenal net body weight (total weight less weight of
gland arise separately, and it is not until rela uterus and contents) is taken as the basis for
T he A d ren al G lands 829
comparison. The medulla : cortex ratio is 1:7 in When injected, its main actions are to raise blood
estrus and 1:8 in pregnancy, compared with a pressure by causing constriction of the blood
ratio of 1:5 in diestrus. vessels and to inhibit the action of the intestinal
Following loss of the adrenal cortex, dogs tract. Metabolism and the capacity to perform
show loss of appetite and weight, general weak work are increased.
ness, and, finally, shock-like coma. In Addison’s Under normal conditions, however, epineph
disease of man, a condition due to degeneration rine is secreted in such minute amounts and is
of the adrenal cortex, similar symptoms with ex destroyed in the body so quickly that its signifi
tensive skin pigmentation are found. Pigmenta cance appears doubtful. Since the rate of secre
tion of the buccal mucosa has been described. tion of the adrenal medulla is increased sharply
These symptoms can be prevented and even by sympathetic (splanchnic) stimulation, it is
comatose animals restored to normal by the ad possible that the medulla plays some part in the
ministration of extracts of the adrenal cortex. emergency response of animals. It is obvious that
Estrus, pregnancy, and lactation will occur nor this response mechanism is of considerable im
mally in treated adrenalectomized bitches and portance to an animal with habits like the aver
growth can be maintained in puppies (Hadlow age dog, but dogs have led apparently normal
1953, Rigdon and Swann 1953, Rogoff 1944, lives after total removal of the medulla.
Rogoff and Stewart 1926, 1927, 1928a, and
1928b).
The functions of the several hormones of the Pathology
adrenal cortex in the normal animal are related
to carbohydrate, mineral, and water metabolism. While insufficiency of the adrenal cortex may
The cortex also appears to have some effect in be an accompaniment or the cause of a number
maintaining the integrity of the nervous system of disease syndromes, only sporadic cases of ad
and the kidney. Of special interest is the relation renal lesions have been reported. In a series of
of the adrenal cortex to sex functions indicated 65 dogs examined by the author, the adrenals of
above. The persistence of sex drive in castrate three specimens were found to bear extensive
subjects, especially if the operation is performed caseous nodules, a figure comparable to the
after maturity, has been attributed to sex hor number of lesions of the other endocrine glands.
mone-like substances produced by the adrenal Undoubtedly more such lesions would be found
cortex. In man adrenal tumors are associated if they were looked for.
with sexual precocity or with masculinization of A syndrome comparable to Addison’s disease
adult females. Injections of cortical extracts have (adrenal insufficiency) in man can be produced
caused precocious sexual maturity in rats. The at will in dogs and cats by subtotal vascular oc
common origin of the adrenal cortex and the clusion of the adrenal venous drainage (Rogoff
gonads from the celomic epithelium suggests 1944). The symptoms of this condition in the dog
fundamental relationships between these or (loss of appetite, severe intoxication, and some
gans. It is not improbable that the abnormal times coma and death) resemble the symptoms
breeding behavior of certain breeds of dogs may of various other systemic diseases. With this fact
be related to the adrenal cortex. The function of in mind, it would not be unreasonable to believe
the adrenal cortex is controlled by the adreno- that the adrenal may be either primarily or sec
corticotrophic hormone of the hypophysis. ondarily involved in a wide variety of conditions
Medulla. The adrenal medulla is not essential (Hadlow 1953, Rigdon and Swann 1953). Stock
for life and is not related, other than topograph ard (1941) has emphasized that the endocrine
ically, to the cortex. The medullary cells produce glands of dogs dying of a wide variety of diseases
a hormone, epinephrine, which has striking phar are not reliable specimens for histological study.
macological effects but is probably of minor sig The use of adrenal cortical extract and cortex-
nificance in the normal economy of the animal. stimulating agents in shock and toxic syndromes,
Epinephrine produces the same effect on struc in both man and animals, especially the dog,
tures receiving sympathetic innervation as does further jeopardizes the validity of findings based
stimulation of the sympathetic fibers themselves. on such specimens.
830 C h apter 16. T he E n d o c r in e System
EN D O C R IN E FU N CTIO N S O F O TH ER ORGANS
damage by x-rays, the seminiferous elements of ICSH at puberty. With the production of the
may be completely degenerated, whereas the male hormone the secretion of ICSH is inhibited
interstitial tissue may be normal or increased in as a result of the reciprocal action of the male
amount, and the animal may be normally mas hormone on the hypophysis. This interaction
culine. Most convincing proof is the histochemi- provides a very necessary and nicely regulated
cal finding of substances with the chemical control of these processes. Castration, with its
properties of testosterone in the interstitial cells attendant loss of male hormone, results in an in
but not in other elements of the testis. The crease in the gonad-stimulating potency of the
greater resistance of the interstitial tissue to un hypophysis.
favorable influences is characteristic of the Spermatogenesis is stimulated by another
mesenchymal tissues in comparison with the gonadotrophic hormone of the pars distalis,
more highly specialized epithelial tissues. which is identical with the follicle-stimulating
Castration before puberty results in persist hormone (FSH) in the female. The secretion of
ence of the juvenile state, not in the development this hormone is also regulated by the level of cir
of characteristics of the opposite sex as is some culating male hormones. Injection of large
times thought. The metabolic rate decreases, amounts of testosterone will increase the libido
favoring the accumulation of fat. The secondary of males, but may injure the seminiferous tu
sex characters do not appear, and the accessory bules.
sex glands do not become functional. The penis The adrenal cortex also produces substances
remains small; this may be a disadvantage in geo possessing androgenic activity but not in suf
graphical areas where urinary calculi commonly ficient quantity normally to substitute for the
occur, since they cannot pass as easily through secretion of the interstitial tissue. Degradation
the urethra. The long bones grow longer than products of the male hormone appear in the
they normally do, because the epiphyseal plates urine of normal males; the amounts present are
fuse later than they do in intact animals. an index of the quantity produced, and thus are
If castration is performed after maturity, the roughly proportional to the virility of the animal.
regression to a neuter state is in keeping with the Female sex hormones, produced by the testis
plasticity of the various structures and functions and the adrenal cortex, also appear in the urine
concerned. Most obvious is the loss of reproduc of normal males. Large amounts of female sex
tive (fertilizing) ability, although sperm present hormone are distinctly harmful to the male re
in the excurrent ducts make the animal capable productive tract. The amounts normally pro
of at least one fertile service after castration. The duced are detoxified by the liver.
metabolic rate drops, but the results on activity
of the animal may be modified by its habits. Fat
is usually deposited. Libido may remain at a high ENDOCRINE FUNCTION OF THE OVARY
level because of the psychic elements involved.
The skeletal system is least modified by late cas Like the testis, the ovary has a dual function,
tration. producing both germ cells and sex hormones,
The interactions of the testis with the hy both functions being under control of the pars
pophysis in the dog are undoubtedly the same as distalis of the hypophysis. The gonadal-hy
in other species of continuously breeding males. pophyseal relationship in the female, however,
The descent of the testes into the scrotum in is more complex than that in the male. This
early life appears to be governed by the gona would seem to be related to the additional func
dotrophic hormones of the hypophysis. Clinical tions in the female of preparing the uterus for
use of such hormones is sometimes successful in implantation of the fertilized egg, maintenance
causing testicular descent in cryptorchid dogs. of pregnancy, and the preparation of the mam
The functional integrity of the interstitial tissue mary glands for nourishment of the young.
and consequently the production of male hor The bitch differs from other domestic animals,
mone depend on the secretion of the interstitial including the cat, in being monestrous. Most
cell-stimulating hormone (ICSH) of the pars dis breeds have but two heat periods a year; estrus
talis. The early functioning of the interstitial tis does not recur in the absence of a fertile mating,
sue in the embryo apparently is due to the pres as is the case in polyestrous animals. That full re
ence of a similar hormone secreted by the dam productive capacity is not attained at puberty is
and circulated via the placenta. After birth the indicated by the fact that the first heat periods
interstitial tissue remains quiescent until the hy of young animals tend to be irregular, and fewer
pophysis begins to secrete appreciable amounts young are born to females mated at that time.
832 C h ap ter 16. T he E n d o c r in e System
This decrease in number is due to the fact that ovary, but its use might more properly be lim
fewer follicles mature in the first few periods ited to the estrogenic substances.
than characteristically do when the animal is ma The follicular hormone liberated into the
ture. This is evidence of the imperfection of the blood stream has several functions. It is respon
endocrine mechanism in early reproductive life. sible for the outward manifestations of sexual
Breeders usually prefer to allow a female to at desire, mediated largely by the central nervous
tain nearly full size before breeding for the first system. It is a powerful stimulant to the repro
time, but it is probable that such females pro duction, growth, and activity of the epithelial
duce fewer offspring in their lifetime than those cells lining the reproductive tract. The number
which are allowed to breed earlier. Dogs are the and height of ciliated cells of the oviducts in
most variable of the domestic animals in their crease. The increased secretory activity of the
breeding behavior. Dogs of small breeds, which tubal and uterine epithelium is made possible by
attain full size earlier, tend to reach puberty a concomitant increase in vascularity of the un
earlier than do dogs of the larger breeds. They derlying stroma. The stratified squamous epithe
tend to show more frequent reproductive cycles, lium of the vagina increases its rate of growth
but litter size is usually smaller than in many of and comification, and the cells of the cervical
the larger breeds. The limitation on litter size is epithelium produce large amounts of mucus
probably a matter of natural selection, since the which covers the vaginal epithelium. This mucus
metabolic demands of puppies of all breeds are serves to protect the wall of the vagina from in
quantitatively similar. The poor breeding be jury during coitus. Comification of the vaginal
havior of certain breeds of dogs, notably the bull epithelium of rodents is so characteristically
dog type, is associated with a defective endo produced by estrogenic hormones that this is
crine constitution. used as a test of their potency in the standardiza
As the follicle begins to develop under the in tion of commercial preparations. It should be
fluence of follicle-stimulating hormone (FSH) noted that comification is an indirect effect re
from the hypophysis, a cavity appears in the sulting from the increased growth of the epithe
growing follicle which fills with fluid. The ovum lium which removes the surface cells from their
is at first surrounded by a single layer of squa blood supply, and hence causes their death. The
mous or cuboidal follicular cells. As the number rate of contraction of the smooth muscle of the
of cell layers increases by mitotic division of the oviduct and uterine horns increases, regulating
cells, vacuolated areas appear which represent the transport of the ovum on its journey from
foci of follicular fluid formation. Fluid is secreted the ovary to the uterus.
by the cells surrounding these vacuoles more In addition to its effects on the reproductive
rapidly than cell division can produce cells to fill tract, the female sex hormone is responsible for
the spaces; hence cavitation ensues. In this proc the appearance of the typical secondary sex char
ess the ovum, surrounded by a few layers of cells, acters of the female. These include the pattern of
the cumulus oophorus, or germ hill, is pushed to fat distribution and of hair, earlier cessation of
one wall of the follicle. The innermost cells form bone growth, and higher voice. The growth of
a columnar layer, the corona radiata, separated the duct system of the mammary gland typically
from the ovum by a clear membrane, the zona occurs with the advent of sex cycles at puberty.
pellucida. Just as an increasing concentration of tes
The fluid-secreting cells become arranged in tosterone inhibits further production of the
a follicular epithelium (membrana granulosa) gonadotrophic hormone of the hypophysis, so
surrounding the cavity. As the follicular cavity estradiol decreases the production of FSH.
enlarges the fluid comes to contain increasing Ovariectomy increases, and injection of large
amounts of estradiol, an estrogenic hormone amounts of estrogen decreases, the gonado
produced by the cells of the follicular epithelium trophic potency of the pars distalis. Thus the
and other elements of the follicle. The term es administration of estrogens to a non-breeding
trogen is applied to the various biologically ac animal will produce the phenomena associated
tive forms of this hormone and their degradation with the reproductive cycle but may depress
products, whether produced by the ovary, pla follicular development, and continued adminis
centa, adrenal cortex, or other tissues, and is tration may result in sterility. The growth-stim
synonomous with the more general term, female ulating property of the estrogens has an impor
sex hormone. The latter term is sometimes tant bearing on neoplasia; the estrogens,
loosely used to indicate other hormones of the whether of endogenous or exogenous origin, are
E n d o c r in e F u n c t io n o f th e P an crea s 833
among the most potent carcinogenic compounds of the pars distalis the corpora lutea persist until
known. They are especially related to the pro near the end of pregnancy. In a number of spe
duction of mammary tumors, which are of com cies, it is known that near the end of preg
mon occurrence in the bitch. nancy the corpora lutea produce another hor
As the follicle increases in size, a connective mone, relaxin, which causes relaxation of the
tissue capsule forms from the interstitial tissue of symphyseal ligament, thus facilitating passage
the ovary. This is differentiated into an outer of the fetus through an otherwise unyielding
dense layer, the theca externa, and an inner layer birth canal. During pregnancy, gonadotrophic
of epithelioid cells, the theca interna, in which and sex hormones are also produced by the pla
an extensive network of capillaries develops. centa and influence the course of fetal develop
This arrangement is suggestive of an endocrine ment.
function. If fertile mating does not occur, the bitch
With the increase in concentration of estradiol usually undergoes a period of pseudopregnancy,
in the follicular fluid, as the fluid increases in a condition which resembles pregnancy except
amount, there is a decrease in production of FSH that no young are developing in the uterus. Per
and an increase in the production of another sistence of the corpora lutea in a functional state
gonadotrophic hormone of the anterior lobe of is the immediate cause of this condition. Not
the hypophysis, the luteinizing hormone (LH). only is there a proliferation of the endometrium,
Ovulation, or release of the ovum by rupture of as in pregnancy, but the mammary glands also
the wall of the mature (graafian) follicle, appears develop, frequendy to the secretory state. Ter
to depend on a balance of the concentrations of mination of pseudopregnancy, like that of preg
estradiol and the gonadotrophic hormones. The nancy, is preceded by a regression of the corpora
histological changes in the follicle which precipi lutea.
tate ovulation seem to be centered around the
production of an avascular area in the follicle ENDOCRINE FUNCTION OF THE PANCREAS
wall, and the pressure of the follicular fluid. A
marked increase in intrafollicular pressure just The exocrine functions of the pancreas have
prior to ovulation causes the follicle to bulge at been considered in Chapter 13, on the Digestive
this avascular area. Ovulation is probably not the System. More significant, from the standpoint of
cataclysmic process it is frequently thought to the vital importance of the pancreas in the econ
be; rather the fluid appears to ooze out. The omy of the animal, is the endocrine function of
ovum usually enters the infundibulum of the the pancreatic islets of Langerhans. These are
oviduct and begins its tubal descent. small aggregates of cells which are delimited
The opening in the ruptured follicle is sealed from the exocrine acini by thin connective tissue
by the more viscous portion of the follicular fluid membranes. Their rich vascularity suggested an
and by the formation of a blood clot, or corpus endocrine nature before this function was estab
hemorrhagicum. About this time the theca in lished. The cells stain more palely than the acini,
terna cells begin to proliferate and fill the cavity and, although in ordinary preparations they all
of the ruptured follicle. They become even more look alike, several cell types maybe distinguished
epithelioid in nature and accumulate a lutein by the use of special stains. The two chief types
(yellow) pigment, thus the name corpus luteum are the alpha cells, and the more numerous beta
(plural—corpora lutea). Most of the lutein cells cells, which are believed to be the source of the
originate by proliferation of the follicular epithe islet secretion, insulin.
lial cells. The lutein cells produce a hormone, That it is the islets that produce insulin is indi
progesterone, which is vital for the maintenance cated, aside from their great vascularity, by the
of pregnancy. Under the influence of proges fact that their degeneration results in the clinical
terone the uterine glands secrete the “uterine condition of diabetes, which can be relieved by
milk,” which is essential for the early nourish injection of insulin, and the fact that hyper-
ment of the fertilized ovum. It moreover sensi insulinism occurs when tumors of the islets are
tizes the endometrium so that it will produce the present. Most of the exocrine portion may occa
maternal placenta. The muscular activity of the sionally be destroyed, as by obstruction of the
uterus diminishes, and the production of FSH by duct, in which case the islets may remain fairly
the pars distalis is inhibited, thus preventing the normal and continue to produce insulin.
recurrence of estrous cycles during pregnancy. The pancreatic islets are formed from cell
Under the influence of the luteotrophic hormone masses which, along with acinar cells, arise from
834 C h ap ter 16. T he E n d o c r in e S ystem
the terminal ends of the pancreatic ducts. The ogenic hormone of the posterior lobe of the hy
islets lose duct connections, and thus come to pophysis. This, together with the level of circu
secrete only into the blood stream. It is apparent lating blood sugar, maintains a fine balance of
that some relatively undifferentiated duct ter insulin secretion.
minations in the adult may give rise to new aci
nar and islet tissue.
Insulin is essential in the metabolism of carbo ENDOCRINE FUNCTION OF THE GASTRO
hydrates. Glucose is absorbed in the small intes INTESTINAL TRACT
tine and carried to the liver via the portal system.
It may be converted by the liver into glycogen While specific endocrine tissues in the gastro
and stored there, or it may pass to the tissues of intestinal tract have not been identified, there is
the body to be used as a source of energy or be convincing physiologic evidence that the mucosa
stored as muscle glycogen. Glucose as such is of the stomach and small intestine secretes spe
present in the blood as the form in which carbo cific hormones. Extracts have been prepared
hydrates are transported to the tissues, to pro from the gastric mucosa which, circulating in the
vide energy or to be converted into glycogen. blood stream, elicit gastric secretion. It is there
Only as glycogen can carbohydrates be stored, fore inferred that certain phases of gastric secre
chiefly in the muscles and liver. The transforma tion are not stimulated by the chemical action of
tions of glucose to glycogen to glucose are medi the foodstuffs, but by liberation into the blood
ated by enzymes which require insulin for their stream of this hormone, gastrin, which in turn
proper function. In the absence of insulin these stimulates secretion of the gastric glands. An
enzymes cannot function; thus absorbed glucose other factor is essential in the maturation of red
remains as such in the blood and is excreted by blood cells.
the kidneys when the blood level becomes too The entrance of the acid contents of the stom
high. The presence of sugar in the urine accounts ach into the duodenum stimulates the produc
for the term diabetes mellitus (mel = honey), the tion of a hormone, secretin, by the cells of the
attraction of bees to urine of such individuals duodenal mucosa. Passing through the blood
having been noted by the Romans. The correla stream, this hormone elicits the secretion of the
tion of the pancreas with this condition, how pancreatic enzymes. Other hormones produced
ever, was not made until the late 19th century, by the duodenal mucosa cause contraction and
when a laboratory caretaker noted swarms of in emptying of the gall bladder, secretion of the in
sects about the urine of dogs which had been testinal glands, and inhibition of the motility of
pancreatectomized for other purposes. the stomach.
The study of diabetes by extirpation of the
pancreas led to an understanding of the funda
mental nature of this condition, but the picture ENDOCRINE FUNCTION OF THE KIDNEY
was complicated by the loss of the digestive en
zymes, especially pancreatic lipase, along with Areas of the kidney cortex which are deficient
the endocrine secretion. Thus such animals also in blood (renal ischemia) produce a secretion,
show serious disturbances of fat metabolism. renin, the action of which causes elevation of
The finding that alloxan, a urea derivative, selec systemic blood pressure (hypertension). The
tively destroys the beta cells of the islets has immediate factor in its production seems to be a
made study of uncomplicated diabetes possible. reduction in the amount of available oxygen. The
It is apparent that the principal factor which epithelioid nature of the cells of the juxtaglomer
controls the secretion of insulin in the normal ular apparatus and the macula densa would sug
animal is the glucose level of the blood circulat gest that these may be the site of renin pro
ing through the pancreas. High blood sugar re duction, but it is not clear whether the changes
sulting from the absorption of glucose from the observed in these cells are a cause or are an
intestine stimulates the production of insulin and effect. The small amounts of renin produced by
the storage of glycogen. Low blood sugar result the normal kidney apparently function in the
ing from an increased utilization of glucose by maintenance of homeostasis of the blood and
the tissues, as in exercise, or with a decreased thus of the body fluids as a whole. Low blood
rate of absorption, decreases the rate of insulin pressure results in a decreased flow of arterial
secretion and initiates the mobilization of stored blood, and thus decreases the amount of avail
glycogen to maintain the blood glucose level. able oxygen to the kidney. Renin may be a factor
The action of insulin is counteracted by a diabet in restoring blood pressure to normal.
B ib l io g r a p h y 835
Modell, W. 1933. Observations on the structure of the blood ence of adrenal extracts on the survival period of adrenal-
vessels within the thyroid gland of the dog. Anat. Rec. 55: ectomized dogs. Amer. J. Physiol. 84: 660-674.
251-269. --------------- 1928b. Studies on adrenal insufficiency; the influ
Morato, M. J. X. 1939. The blood supply of the hypophysis. ence of “heat” on the survival period of dogs after adre
Anat. Rec. 74: 297-320. nalectomy. Amer. J. Pliysiol. 86: 20-24.
Mulligan, R. M., and K. C. Francis. 1951. Weights of thyroid Ross, W. D., and V. H. K. Moorhouse. 1938. The thyroid nerve
and parathyroid glands of normal male dogs. Anat. Rec. in the dog and its function. Quart. J. exp. Physiol. 2 7 :209-
110: 139-143. 214.
Nicander, L. 1952. Histological and histochemical studies on Schwartz, H. G. 1936. The meningeal relations of tlie hypoph
the adrenal cortex of domestic and laboratory animals. ysis cerebri. Anat. Rec. 67: 35-51.
Acta anat. (Basel) 14 (Suppl. 16): 1-88. Shambaugh, P. 1936. Autotransplantation of parathyroid gland
Nonidez, J. F. 1931. Innervation of the thyroid gland; origin in the dog. Arch. Surg. 32: 709-720.
and course of the thyroid nerves in the dog. Amer. J. Anat. Smith, E. M., M. L. Calhoun, and E. P. Reineke. 1953. The
48: 299-329. histology of the anterior pituitary, thyroid and adrenal of
--------------- 1932a. Origin of the parafollicular cell, a second tliyroid-stimulated purebred English bulldogs. Anat. Rec.
epithelial component of the thyroid gland of the dog. 117: 221-240.
Amer. J. Anat. 49: 479-505. Stigliani, R., P. L. Cipriani, and R. Del Vivo. 1954. Topografia
--------------- 1932b. Further observations on the parafollicular dell’ipofisi del cane. Arcli. De Veechi Anat. pat. 2 2 :309-
cells of the mammalian thyroid. Anat. Rec. 53: 339-347. 321.
Purves, H. D., and W. E. Griesbach. 1957. A study on the cy Stockard, C. R. 1941. The genetic and endocrine basis for dif
tology of tlie adenohypophysis of the dog. J. Endocr. 14: ferences in form and behavior. Amer. Anat. Memoir 19.
361-370. Philadelphia, Wistar Institute of Anatomy and Biology.
Putnam, J. T., E. B. Benedict, and H. M. Teel. 1929. Studies in Swarts, J. L., and R. L. Thompson. 1911. Accessory thyroid
acromegaly; experimental canine acromegaly produced tissue within the pericardium of the dog. J. med. Res. 29:
by injection of anterior lobe pituitary extract. Arch. Surg. 299-308.
18: 1708-1736. Van Dyke, H. B. 1936. The Physiology and Pharmacology of
Randolph, K. H. 1950. Growth changes in the adrenal gland the Pituitary Body. Vols. 1 and 2. Chicago, University of
of tlie dog from birth to two years of age. M. S. Tliesis, Chicago Press.
Iowa State University. Venzke, W. G. 1940. Histology of the thyroid glands of dogs 16
Reed, C. I., R. W. Lackey, and J. I. Payte. 1928. Observations weeks of age. Proc. Iowa Acad. Sci. 46: 439-441.
on parathyroidectomized dogs, with particular attention Vicari, E. M. 1932. Thyroid and parathyroid size in various
to the regional incidence of tetany and to the blood min pure-bred dogs and their hybrids, with histological find
eral changes in this condition. Amer. J. Pliysiol. 84: 176- ings. Anat. Rec. 52 (Suppl.): 40.
188. --------------- 1937. Observations on the nature of the parafol
Rienhoif, W. F., Jr. 1938. The lymphatic vessels of the thyroid licular cells in the thyroid gland of the dog. Anat. Rec. 68:
gland in the dog and in man. Arch. Surg. 23: 783-804. 281-285.
Rigdon, R. H., and H. G. Swann. 1953. Morphologic changes White, J. B., and H. L. Foust. 1944. Growth changes in body
in tlie dog’s adrenal gland following anoxia. Proc. Soc. organs; growth changes in the pituitary of the normal dog.
exp. Biol. (N.Y.) 82: 111-115. Amer. J. vet. Res. 5: 173-178.
Rogoff, J. M. 1944. Experimental pathology and physiology of Wolfe, J. M., and R. Cleveland. 1932. Cell types found in the
the adrenal cortex; production of Addison’s disease in anterior hypophysis of the dog. Anat. Rec. 52: 43-44.
laboratory animals. Arch. Path. 38: 392-409. Wolfe, J. M., R. Cleveland, and M. Campbell. 1933. Cyclic
Rogoff, J. M., and G. N. Stewart. 1926. Studies on adrenal in histological variations in the anterior hypophysis of the
sufficiency; control animals not subjected to any treat dog. Z. Zellforsch. 17: 420-452.
ment. Amer. J. Physiol. 78: 683-710. Yamada, H., S. Ozawa, and R. Endo. 1956. Histological studies
--------------- 1927. Studies on adrenal insufficiency; the influ on the mammalian pituitary gland with special reference
ence of pregnancy upon the survival period in adrenalec- to the innervation. Bull. Tokyo med. dent. Univ. 3 :51-65.
tomized dogs. Amer. J. Physiol. 79: 508-535. Zechel, G. 1931. Follicular destruction in the normal thyroid
--------------- 1928a. Studies on adrenal insufficiency; the influ of the dog. Surg. Gynec. Obstet. 52: 228-232.
CHAPTER 17
TH E SENSE O RG AN S
A N D IN TE G U M EN T
Various end-organs or sensory receptors serve to or auditory ossicles serves to connect the tym
receive information and transmit it to the central panic membrane with the oval window of the
nervous system. The receptors in the organs of internal ear. The internal ear, contained within
special sense (eye, ear, nose, tongue, and skin) the petrous temporal bone, is composed of a
are modified in accordance with the type of spiral tube or cochlea (for hearing) and the
stimuli they receive. The information transmit vestibular apparatus (for balance).
ted to the brain by these variously modified end- The organ of smell (organum olfactus) is lo
organs results in the sensations of sight, hearing, cated within the nasal cavity as a portion of the
smell, taste, touch, pressure, heat, cold, and nose. The functions of the nose include olfaction
pain. and air conduction.
The eye, or organ of sight (organum visus), is The organ of taste (organum gustus) is repre
located within the cavity of the orbit. Certain sented by taste buds distributed over the tongue
accessory structures such as the muscles, fasciae, and occasionally on adjacent areas. The micro
eyelids, conjunctivae, and the lacrimal apparatus scopic taste buds are associated with the papil
are associated with the bulb of the eye (bulbus lae, which are seen as projections of a connective
oculi). tissue core covered with stratified squamous
The ear, or organ of hearing (organum vestibu- epithelium.
locochleare), is divided into an external, middle, The common integument (integumentum
and internal ear. The size, shape, and position of commune) is the protective covering of the body.
the external ear of the dog vary with the breed. It consists principally of layers of dense connec
The middle ear, with its tympanic cavity en tive tissue termed the corium and an outer layer
closed by a bony bulla, communicates with the of epithelium termed the epidermis. Within the
pharynx via the pharyngotympanic tube. Within integument are glands, hair follicles, smooth
the middle ear cavity a chain of movable bones muscles, and sensory nerve endings.
By ROBERT GETTY
The vertebrate eye (oculus) and the muscles smaller breed, the orbit may not necessarily in
which provide for its movement are contained crease in diameter proportionately. Ocular
within the orbit. The extraocular muscles (mus- pathology and injury are common in the dog.
culi bulbi), conjunctiva (tunica conjunctiva),
palpebral ligaments (ligamenta palpebrales), and THE EYELIDS
periorbital fat (corpus adiposum orbitae) suspend
and cushion the eye. There is considerable vari The eyelids (palpebrae) may be thought of as
ation in the size of the eyes between breeds. Al integumentary curtains which protect the ante
though a large dog may have a larger eye than a rior surface of the eyeball. They not only are
837
838 C h apter 17. T h e Sen se O rgans and In teg u m en t
capable of excluding light from the eye, but they and lower lids will be seen to have a small slitlike
also contain auxiliary glands which supplement orifice near the pigmented edge of the lid only
the fluids from the larger orbital glands. In addi a few millimeters from the medial canthus. These
tion to the upper (dorsal) and lower (ventral) openings (puncta lacrimalia) through which the
eyelids, a so-called third eyelid is found, which lacrimal fluid drains from the lacrimal lake (lacus
in man has become the vestigial plica semilu lacrimalis) are located in the lacrimal fossa of the
naris conjunctivae (Prince et al. 1960). The two medial orbital wall. The medial canthus does not
eyelids (palpebra dorsalis et ventralis) converge rest directly against the eyeball, but is separated
and join to form the m edial and lateral angles from it by the lacrimal lake. The lacrimal punc-
or canthi (angulus oculi medialis et lateralis). The tum of each lid represents the beginning of the
medial canthus is somewhat more obtuse. The dorsal and ventral lacrimal ducts (canaliculi lac-
upper eyelid is more movable than the lower rimales). The lower lid shows a slight protuber
one. When the lids are open, the interval be ance (papilla lacrimalis) at the site of this orifice.
tween them is known as the palpebral fissure. The lacrimal gland (glandula lacrimalis) (Fig.
Both eyelids are covered with normal, hair- 17-2), which is located ventral to the supraorbi
bearing skin on their outer or cutaneous surface. tal process and medial to the orbital ligament,
The inner surface is covered by a thin mucous liberates its secretion into the dorsolateral part
membrane or modified fascial sheath called the of the conjunctival sac. The lacrimal fluid that
conjunctiva. drains via the lacrimal ducts into the lacrimal sac
The conjunctiva consists of two parts, a bulbar (saccus lacrimalis) enters the nasolacrimal duct
and a palpebral portion. The palpebral conjunc (ductus nasolacrimalis), located within the naso
tiva (tunica conjunctiva palpebrarum) lines the lacrimal canal, to empty into the nasal cavity.
inner surface of the lids, and the bulbar (ocular) The eyelashes (cilia) are located along the
conjunctiva (tunica conjunctiva bulbi) covers the edge of the upper eyelid. Well-defined eyelashes
anterior part of the eyeball except the cornea are absent on the lower lid. Associated with the
(Fig. 17-1). Thus the conjunctiva is seen grossly hair follicles of the cilia are sebaceous glands (of
as a smooth, slick surface which is reflected in a Zeis) and rudimentary sweat glands (of Moll).
continuous manner from the inner surface of the Long, coarse hairs are also observed growing in
lids onto the eyeball. The partially enclosed po a clump above the medial canthus of the eye.
tential space between the palpebral and bulbar
conjunctivae is known as the conjunctival sac Nictitans
(saccus conjunctivae). The deepest part of the
conjunctival sac where an angle is formed be The third eyelid (palpebra tertia), or nictitat
tween the two conjunctivae presents an area of ing membrane, in the dog is located at the me
abrupt reflection of the conjunctiva and is called dial angle of the palpebral aperture (Fig. 17-3).
the conjunctival fornix (fornix conjunctivae). It follows the curvature of the eyeball and is
The conjunctiva is continuous with the skin therefore correspondingly convex on the outside
superficially along the line of the lid edges, the and concave on the inside. The edge of the mem
palpebral rim (rima palpebrarum). Histologi brane may be either pigmented or unpigmented,
cally, the conjunctiva consists of connective tis and both surfaces are covered with epithelium
sue beneath epithelium which contains colum which is continuous with the bulbar conjunctiva
nar cells and goblet cells. Lymphatic nodules on one side and the palpebral conjunctiva on the
are also present. other. This third eyelid protrudes from the me
One can observe through the conjunctiva on dial canthus of the eye. Embedded within the
the inner surface of the eyelids the tarsal (Mei third eyelid is a flat, more or less T-shaped hya
bom ian) glands (glandulae tarsales). The orifices line cartilage (Fig. 17-4). Glandular tissue sur
of the glands, which sometimes number forty rounds the cartilage, particularly along the
(Prince et al. 1960), lie along the inner lid mar shaft. This single superficial gland of the third
gins. Their secretion is more viscous than lacri eyelid is largely molded to the cartilaginous
mal fluid and thus aids in preventing the more process of the nictitating membrane. The gland
watery fluid from the lacrimal gland from over is referred to as a nictitans gland when it is asso
flowing. The connective tissue surrounding the ciated with the third eyelid. The gland, which is
tarsal glands is dense, but the plate of connective relatively small, is surrounded by fat and is at
tissue (tarsal plate) is not as well developed in tached to the surrounding area by connective
the dog as in man. tissue. According to Prince et al. (1960), the cells
If the eyelids are partially everted, both upper of the gland react positively to the P.A.S. test
T he E ye, O r b it , a n d A dn exa 839
Lens
C o r n e a l e p ith e liu m ------------- J
— — — P u p illa ry a p e rtu re
S u b s t a n t ia p r o p r ia o f c o rn e a
C o rn e a l m e s o th e liu m
P o s te rio r c h a m b e r
-Z o n u lar lig a m e n ts
(Fib ers of s u s p e n s o ry lig a m e n t)
S p h in c te r o f ir i s -------------
C ilia ry p ro c e s s
D i l a t o r o f i r i s ------------------
C ilia r y b o d y
A n te r io r c h a m b e r -— —
T ra b e c u la e — — "
C ilia r y m uscle
P a lp e b ra l co n ju n ctiva
R e tin a
O c u la r c o n ju n c tiv a x
C h o ro id
S c le r a l ven ous plexus
S c le r a
F ig . 17-1. Longitudinal section of anterior part of eye showing attachments of cornea, iris, and lens.
Temporal f o s s a
Zygomotic p ro ce ss
Orbital ligament
Sclera
Cornea
___________ Pupil
A - ^ ^ s ^ _ _ _ p a lpebra terfia
------------------ L a c r i m a l gland
" Mandible
'M a s s e t e r m.
and are classified as seromucoid. The secretion sertion of the dorsal rectus muscle. The dorsal
of the gland passes to the conjunctival sac oblique muscle is innervated by the trochlear
through minute ducts. In some species, particu nerve.
larly the pig, the gland presents both a superficial The ventral oblique muscle of the eye origi
and a deeper portion. The deeper gland is called nates ventral to the lacrimal fossa near the origin
the deep gland of the third eyelid, or Harder’s of the medial pterygoid muscle and dorsal to the
gland (Harder 1694). The dog, however, does not maxillary foramen. The muscle is short, passes
have a deep gland as described by Harder. One laterally, and inserts on the sclera of the eyeball
can also observe, on the bulbar surface of the in the region of the insertion of the lateral rectus
third eyelid, a mass of diffuse, slightly raised muscle. The inferior oblique muscle is supplied
lymphoid tissue (Fig. 17-5). This tissue maybe- by the oculomotor nerve. The dorsal, medial,
come inflamed and should not be confused with and ventral straight muscles of the eye are also
the gland of the third eyelid. The nictitating innervated by the oculomotor nerve. The lateral
membrane and a small elevation known as the straight muscle and the retractor receive inner
lacrimal caruncle (caruncula lacrimalis) are lo vation from the abducens nerve. Thus the vol
cated in the medial canthus of the eye. This pro untary muscles within the orbit are supplied
jection within the lacrimal lake is thought by by the third, fourth, and sixth cranial nerves.
some to be a modified remnant of the eyelid in The smooth musculature of the orbit includes the
tegument and is covered by stratified epithelium dorsal and ventral palpebral muscles and the
containing modified sweat and sebaceous glands. ciliary and iridial musculature within the eye
According to Prince et al. (1960), in addition to ball. The specific innervation is discussed under
the modified sweat glands there are also some the respective headings.
modified lacrimal glands of the tubulo-alveolar
type. It has been suggested that the caruncula THE LACRIMAL GLAND AND APPARATUS
lacrimalis may aid in preventing foreign objects
from entering the lacrimal puncta. The lacrimal apparatus consists of a secretory
portion, the lacrimal gland and its ducts; the lac
EXTRAOCULAR MUSCLES rimal lake, the lacrimal canaliculi and sac, and
the nasolacrimal duct. The lacrimal gland is a
The extraocular muscles are described in large, modified skin gland which has become
Chapter 3 with the muscles of the head. The specialized as a tubulo-alveolar structure. It con
seven muscles of the orbit include dorsal, ven forms to the contours of the surrounding tissue,
tral, medial and lateral rectus muscles, dorsal and and macroscopically it may be confused with
ventral oblique muscles, and the retractor bulbi muscle. It is fairly well demarcated, however,
muscle. The four rectus muscles are flat and have being spatula-shaped and located on the dorso
their origin around the optic canal (foramen), lateral side of the globe within the periorbita. It
orbital fissure, and pterygoid crest. The muscles is light red, flat, and slightly lobulated. It lies just
of the eyeball insert into the sclera by means of below the tip of the supraorbital process of the
aponeurotic tendons. The retractor bulbi (oculi) frontal bone and is attached by a tendinous por
in the dog is a muscle which surrounds the optic tion to the zygomatic bone. Small ducts, which
nerve and interdigitates with the straight mus are not readily seen, open into the conjunctival
cles of the eye. It forms four distinct bundles lo sac at the superior fornix. Histologically, it is
cated between the recti. Occasionally the mus made up of both serous and mucous acini ar
cle may appear as a complete cone with the apex ranged in lobules. According to Trautmann and
at the orbital fissure and the base at its insertion Fiebiger (1957), the secretory cells often contain
into the globe. The dorsal oblique muscle of the fat droplets in all domestic animals.
eye originates medial to the optic foramen and
passes forward medial and dorsal to the medial ZYGOMATIC GLAND (ORBITAL OR
rectus muscle, separated from it by a vascular DORSAL BUCCAL GLAND)
plexus of the orbital vein. The muscular belly of
the dorsal oblique muscle gives way to a slim In addition to the lacrimal gland and the gland
tendon, which passes over a cartilaginous plate of the third eyelid, the dog has a zygomatic gland
(trochlea) held to the dorsomedial orbital wall (Fig. 17-2) which is also located in the orbital
under the zygomatic process of the frontal bone. region. The zygomatic gland is slightly lobulated
The tendon then turns laterally and inserts on and pyramidal in shape, the apex lying beneath
the sclera of the eyeball in the region of the in the zygomatic arch of the temporal bone. It is
T he E ye, O r b it , a n d A dn exa 841
S c l e r a _______ . Iris
. P u p il
----------Lateral canthus
M e d ia l c a n th u s
E x te n t o f c a rtila g e o f P a lp e b ra t e r t i a
P a lp e b ra te r tia (P a lp e b r a l s u r f a c e )
in contact with the periorbita above, but sep palpebral ligaments are intimately connected
arated from the pterygoid muscle below by fat. with the palpebral conjunctiva on their deep
Its secretions pass through ducts which enter face. The dorsal and ventral tarsi are slightly
the mouth near the last upper molar tooth. thickened areas of fibrous tissue. They are
curved, flattened, elongated, semilunar plates
MUSCLES OF THE EYELID which tend to stiffen and give form to the edge
of the palpebrae. The dorsal tarsus is somewhat
The muscles of the eyelids are four in number: larger and stronger. Both are suspended in the
orbicularis oculi, retractor anguli oculi, corru- palpebral ligament.
gator supercilii (superciliaris), and levator palpe The corrugator supercilii muscle has two parts.
brae superioris. The medial portion is the largest. It is a thin,
The orbicularis oculi muscle is located close to small muscle, which spreads out in the upper
the edge of each eyelid, and is seen as a flattened eyelid, blending with the orbicularis oculi mus
ring of muscle fibers extending around the palpe cle. The smaller, lateral portion of the muscle
bral fissure. It is well developed in the dog and is fuses in part with the lateral retractor muscle.
attached to the orbital wall by the medial (nasal) The muscle arises at the mid line from the tem
palpebral ligament. Laterally, the orbicularis poral or nasofrontal fascia. The action of the
oculi muscle blends with the relatively small re corrugator supercilii muscle is to assist in elevat
tractor anguli oculi muscle, which serves as an ing the upper lid.
equivalent to a lateral palpebral ligament. The The levator palpebrae superioris muscle arises
action of the muscle is to close the palpebral fis deep within the orbit. It is a long, narrow band
sure. of muscle dorsal to the dorsal rectus muscle. The
The retractor anguli oculi muscle is approxi anterior tendon of insertion of the levator palpe
mately 1 to 2 cm. long and about 3 to 5 mm. brae superioris muscle fans out and fuses with
wide, varying with the breed of the dog. It ex the dorsal palpebral ligament. The origin of the
tends from the lateral canthus of the eye to the muscle is the dorsal margin of the optic canal.
temporal fascia. It is partially superficial to and The contraction of this muscle raises the upper
partially continuous with the orbicularis oculi lid. The lower eyelid depresses feebly when the
muscle. The medial homologue of this retractor orbicularis oculi muscle is relaxed.
muscle is the nasal palpebral ligament. The lat
eral retractor lengthens the palpebral fissure. BLOOD AND NERVE SUPPLY
The aponeurotic sheath of the orbicularis
oculi muscle on its deep face is fused with the The sensory nerve supply to the palpebral re
palpebral ligament, a portion of the palpebral gion is via the ophthalmic branch of the trigem
fascia. This fascia is in the form of a membranous inal nerve to the upper lid and via the maxillary
cone surrounding the contents of the orbit and branch of the trigeminal to the lower lid. The
continuous with the periosteum at the optic orbicularis oculi, corrugator supercilii, and re
canal and orbital margins. The palpebral liga tractor palpebrarum muscles are innervated by
ment is formed in the upper lid by the tendon of the palpebral branch of the facial nerve. The
insertion of the levator palpebrae superioris mus levator palpebrae superioris muscle is inner
cle, the deep sheath of the orbicularis oculi mus vated by the oculomotor nerve. The blood sup
cle, and the process from the orbital septum. It ply to the eyelids is via the malar and superficial
is continuous medially with the medial palpebral temporal arteries. The conjunctiva is supplied
ligament; anteriorly, it inserts on the superior by the palpebral and anterior ciliary arteries,
tarsus and the skin of the palpebral rim. Later and drainage is via the palpebral veins.
ally, it blends with the sheath of the retractor
palpebrarum muscle and the periosteum of the THE PERIORBITA AND THE ORBITAL FASCIA
zygomatic process of the malar bone, and dorso-
caudally it is continuous with the orbital sep The periosteum of the orbit is usually referred
tum. The palpebral ligament of the lower lid is to as the periorbita. At the margin of the orbit
somewhat similar, extending from the medial the periorbita thickens and becomes continuous
palpebral ligament to the sheath of the retractor with the periosteum of the external surface of
palpebrarum muscle and the periosteum of the the skull. Orbital fat fills the portions of the orbit
zygomatic process of the malar bone in a thick not occupied by other structures. The fascia
ened, slinglike band. The ventral palpebral liga bulbi (Tenon’s capsule) is a fibrous layer be
ment also inserts on the ventral tarsus. Both tween the orbital fat and the eyeball itself. It is
T he E ye, O r b it , and A d n exa 843
attached firmly to the sclera around the point portions, the choroid, the ciliary body, and the
of entrance of the optic nerve and is also at iris. The internal membrane of the eye contains
tached firmly anteriorly to the sclera near the the peripheral end-organs serving the sense of
corneoscleral junction. Thus the contents of vision and is referred to as the tunica interna,
the orbit are enclosed within a conical, fibrous tunica nervia, or retina.
periorbital membrane. The extraocular muscles
of the eye of the dog are also enclosed in fibrous Fibrous Coat (Cornea and Sclera)
and fascial sheaths.
The eyes of the dog are located in the orbits The fibrous coat is the outer tunic of the eye
with a minimum angle of 2 0 ° between the visual ball. It is divided into two parts. The cornea
axes (Fig. 17-6), according to Prince et al. (1960). comprises approximately a sixth of the tunic,
This angle varies considerably between breeds. and the sclera the remaining five sixths.
The eye is composed of segments of two asym The cornea is the transparent distal portion
metrical spheres giving it the form of an oblate which is almost circular and, like the globe as a
spheroid. The anterior, transparent segment whole, varies in dimension between the breeds.
(cornea) has a radius of curvature one-third The vertical meridian has the smaller dimension.
smaller than the posterior, opaque segment The cornea is thicker in the center than at the
(sclera). The canine eye is relatively large for the periphery. This is the reverse of that found in
size of the animal and shows considerable varia man. The transition from the dense, scleral
tion between breeds. The anterior-posterior di fibrous structure to the transparent corneal
ameter, or sagittal diameter through the poles, structure is comparatively abrupt. The cornea is
is the greatest diameter, whereas the transverse largely avascular, but it is supplied abundantly
diameter is slightly greater than the vertical di with sensory nerves derived from the ciliary
ameter. The eyeball (bulbus oculi) lies in a bony nerves.
cavity which is bounded laterally by a fibrous Microscopically, the cornea is composed of
band of tissue and a few bands of smooth muscle five layers: (1) Epithelium. The anterior surface
called the orbital ligament. This ligament passes of the cornea is covered with a stratified squa
from the supraorbital process of the frontal bone mous epithelium which is continuous peripher
to the zygomatic bone. The shape of the bulbus ally with the conjunctiva. The corneal epithelium
is that of a hollow sphere having a wall com exhibits great powers of regeneration and is
posed of three concentric tunics. exceedingly sensitive. (2) Bowman ’s membrane.
In order to facilitate orientation on the eye, The epithelium is divided from the substantia
special designations as to direction and position propria by a homogeneous membrane. This
are used. The terms “distal” or “corneal” (ante membrane is very thin in the dog. It is con
rior pole) and “proximal” or “cerebral” (poste sidered to be a modification of the substantia
rior pole) are used for direction and position. propria, and although it is fairly tough, it is said
The two poles are connected by the optical axis. not to have any powers of regeneration (Prince
A meridian is formed by a peripheral connecting et al. 1960). (3) Corneal stroma. The main por
line between two poles. Thus there may be a tion of the cornea, the substantia propria, is com
horizontal as well as a vertical meridian. A line posed of modified connective tissue arranged in
around the greatest width of the sphere, which the form of many lamellae. Small flattened cells
is perpendicular to the optical axis, is called the can be seen between the lamellae and are likened
equator. Thus the bulbus can be divided into to those of the Haversian system of bone (Wolff
quadrants by planes or sections through the 1961). (4) D escem et’s membrane. Descemet’s
main meridian. membrane sharply delineates the substantia
propria from the endothelial layer. This mem
THE GLOBE OF THE EYE (EYEBALL) brane is approximately four times the thickness
of the posterior layer of endothelium. (5) Endo
The wall of the eyeball is composed of three thelium. The most posterior layer of the cornea
layers: the external, middle, and internal mem is endothelium, which is continuous with that
branes of the eye. The external membrane of the of the anterior surface of the iris.
eye is dense and fibrous and hence is called the The opaque sclera constitutes the posterior
tunica externa (tunica fibrosa). It is composed five-sixths of the fibrous tunic. It is dense in
of two portions, anteriorly the cornea and pos consistency and in general has a dull white color.
teriorly the sclera. The middle membrane, tunica At the equator the sclera is thin. It is much
media (tunica vasculosa), is composed of three thicker in the ciliary region and around the optic
844 C h apter 17. T he Sen se O rgans and In tegum en t
nerve. The rectus muscles insert anteriorly, entrance of the optic nerve, the choroid shows
whereas the retractor oculi muscle inserts near a triangular area of iridescent luster (tapetum)
the equator of the globe where the sclera is which has a distinctive color for each species
thin. Tenon’s capsule attaches near the corneo (Nicolas 1925). The tapetum of the dog varies
scleral junction. in brightness and in color from green through
Histologically, the sclera presents both fibrous yellow, and gold to pink. The fibrous tapetum
and elastic fibers. In addition to receiving the found in herbivores is composed largely of un
attachments of the muscles of the eyeball, it is dulating collagenous fiber bundles, whereas the
perforated also by the passage of nerves and cellular tapetum, as seen in the carnivores, is
blood vessels. The intrinsic nerves and vessels made up largely of irregular cells. The tapetum,
enter the proximal (posterior) pole; however, the located behind the retina, lies between the
vortex veins leave the eye anterior to the equa choriocapillaris (layer of small vessels) and
tor. The lamina cribrosa is a thin portion of the the larger vessel layer of the choroid. In the
sclera which is perforated by numerous holes dog the larger vessels are prominent, but fre
through which pass the axons that form the quently the choriocapillaris is difficult to identify,
optic nerve. for the vessels are not plentiful. The choroid con
The long and short posterior ciliary arteries sists largely of a plexus of vessels. Macroscopi
and nerves pass through the sclera in a circle cally, the dog’s tapetum varies in shape among
around the optic nerve. The anterior ciliary the breeds; it may be triangular to semicircular.
vessels, which are given off the muscular ar It may be in contact with the optic disc and ex
teries that supply the extraocular muscles, pene tend a little more than halfway to the periphery,
trate the sclera just caudal to the corneoscleral or sometimes there is a tapetum-free zone
junction. This junction, which is sharply de around the optic papilla. The tapetum is less
fined macroscopically, appears microscopically colorful and bright in the dog as compared to
as an area of gradual transition. The term the cat. The vessels of the choroid, which con
“limbus” is sometimes applied to this junction. verge to form the four venae vorticosae, pene
Trabeculae are found at the limbal iris angle. trate the sclera in the region of the insertion of
Small channels are seen in the trabecular net the retractor bulbi muscle.
work which lead into the scleral venous plexus Anterior to the choroid the ciliary body rep
(Fig. 17-1). The latter drain the aqueous humor resents a thickened, middle portion of the uveal
from the anterior chamber of the eye. These tract. The ciliary body and the choroid are con
vessels are thought to function in a manner simi tinuous with each other at an undulating line
lar to the canal of Schlemm in man. The larger known as the ora serrata (Fig. 17-7). The ciliary
vessels appear to have connections with the body is continuous anteriorly with the iris, and
perilimbal vascular system. on its inner surface it is associated with the non-
nervous continuation of the retina. Therefore
the ora serrata marks the most forward extension
Vascular Tunic of the nervous elements of the retina. The ex
ternal surface of the ciliary body which borders
The middle coat of the eye (tunica vasculosa the sclera is smooth; however, its internal surface
oculi) is often referred to as the uvea or uveal is thrown up into a number of radiating folds
tract and consists of three continuous parts from called the ciliary processes. These processes are
behind forward: the choroid, the ciliary body, pyramidal and radiate from their bases, which
and the iris. The uvea is firmly attached to the are directed toward the lens. Upon close exami
sclera in a ring about the optic nerve and also is nation short ciliary processes are seen irregularly
attached firmly to the sclera at the corneoscleral distributed between long ciliary processes. The
junction. Between these two areas the attach ciliary processes usually number seventy to
ment is very delicate. eighty. It can also be observed that the ciliary
The choroid coat forms approximately the body is wider on the temporal side, and that it
posterior two-thirds of the uvea and is deficient is wider from the pupil edge to the point where
posteriorly over the lamina cribrosa, allowing it becomes the ora ciliaris retinae.
the exit of the optic nerve. The choroid, which The ciliary muscles are unevenly distributed.
lies between the sclera and the optical portion The fibers are seen at the junction of the cornea
of the retina, is the thickest portion of the tunica and sclera and extend to the ciliary processes
media. toward the ora ciliaris retinae. These muscles
At varying distances, dorsal to the point of control accommodative focusing.
T he E y e , O r b it , a n d A dn exa 845
Iris ^
Lens
A n te rio r cham b er
PoSTerior cham ber--------
C ilia ry process
C iliary b o d y — —
Vitreous b od y — Retina
P res p h en o id bone
F ig . 17-6. Frontal section through orbit and ocular adnexa.
-Sclera
Choroid
Cornea
--------------Retina
Lens
F ig . 1 7 -7 . Posterior view o l eye with lens sectioned, revealing ciliary processes and iris.
846 C h ap ter 17. T he Sen se O rgans and In teg um en t
Frontal sinus
_ --Cerebrum
Levator palpebrae superioris
^D ura mater
Vitreous b o d y ^ ^
^ Dorsol rectus m.
Li
„ ^O ptic nerve
Cornea
___--R e tra c to r oculi m.
Anterior chamber
_ _ _ -V e n tra l rectus m.
Posterior chamber
_ —— Pterygoideus medialis m
Ciliary body — " "
------- Palatine bone
~ ~ — Nasopharyngeal canal
Ventral oblique m.
^ 'O r a l pharynx
Lens
Michaelson, I. C. 1954. Retinal Circulation in Man and Ani
The lens of the eye is a transparent, biconvex, mals. Springfield, 111, Charles C Thomas.
and laminated structure. Its anterior surface is Nicolas, E. 1925. Veterinary and Comparative Ophthalmol
ogy. London, H. and W. Brown.
covered by epithelium and is bathed by aqueous Prince, J. H. 1956. Comparative Anatomy of the Eye. Spring
humor, while its posterior surface is in contact field, 111, Charles C Thomas.
with the vitreous body. The circumferential line Prince, J. H , C. D. Diesem, I. Eglitis, and G. L. Ruskell. 1960.
between the anterior and posterior surfaces is Anatomy and Histology of the Eye and Orbit in Domestic
Animals. Springfield, 111, Charles C Thomas.
known as the equator. It is at the equator that
Smelser, G. K. (ed.) 1961. The Structure of the Eye. New York
the epithelial cells are transformed into the and London, Academic Press.
elongated lens fibers which enable the lens to Smythe, R. H. 1956. Veterinary Opthalmology. London, Bail-
grow slightly throughout life. Enclosing the lens liere, Tindall and Cox.
is an elastic capsule to which the zonular fibers --------------- 1961. Animal Vision. Springfield, 111. Charles C
Thomas.
or suspensory ligament of the lens is attached. Trautmann, A , and J. Fiebiger 1957. Fundamentals of the
histology of domestic animals. Ithaca, N.Y. Comstock
BIBLIOGRAPHY Press.
Walls, G. L. 1942. The Vertebrate Eye. Bloomfield Hills,
Harder, J. J. 1694. Glandula nova lachrymalis una cum duct Michigan, Cranbrook Institute of Science.
unexeritoria in Cervis und Damis ad., Acta Eruditorum, Wolff, E. 1961. The Anatomy of the Eye and Orbit. London,
Lipsiae. H. K. Lewis.
THE EAR
By ROBERT GETTY
The ear is divided, for purposes of discussion, (auris media) consists of the tympanic cavity, the
into three portions: the external ear, the middle tympanic membrane, and the three auditory
ear, and the internal ear. The external ear (auris ossicles with their associated ligaments and mus
externa) consists of the auricle, or pinna, and cles. The middle ear cavity is connected with the
the external auditory meatus. The middle ear pharynx by way of the auditory or Eustachian
848 C h ap ter 17. T he Sen se O rgans and In tegum en t
tube. The internal ear (auris interna) includes the scapha. A relatively dense, irregularly quad
cochlea and semicircular canals and is enclosed rangular plate of cartilage known as the tragus
in the petrous portion of the temporal bone (Bast forms the lateral boundary of the initial portion
and Anson 1949). It consists of a membranous of the ear canal lying opposite the anthelix. The
and an osseous or periotic labyrinth (Gray 1907). tragus (Fig. 17-10) curves caudomedially and
The internal ear is the organ for both hearing with the proximal end of the antitragus com
and equilibrium, whereas the external ear and pletes the caudal boundary of the opening into
middle ear represent a sound-collecting and the ear canal. The antitragus is a thin, elongated
conducting apparatus (Honda 1908; Miller and piece of cartilage caudal to the tragus and sepa
Witter 1942; Getty et al. 1956). rated from it by an important notch, the incisura
intertragi ca.
EXTERNAL EAR The antitragus may be divided into two limbs:
the medial cornu and the lateral cornu. They are
The pinna (auricula) of the external ear is a caudally demarcated by the antitragic incisure.
funnel-like plate of cartilage which serves to The apex of the lateral cornu ends in a sharp
receive air vibrations and transmit them via the process, the styloid process of the antitragus.
ear canal to the tympanic membrane (eardrum) Just distal to this the caudal border presents the
(Fig. 17-9). The tympanic membrane is enclosed cutaneous helicine pouch. A prominent feature
in the deep portion of the external acoustic of the caudal border of the auricular cartilage is
meatus and forms the lateral wall of the middle the caudal process of the helix. Proximal to the
ear. The pinna is covered on both sides with skin caudal process in the region of the cutaneous
which is tightly attached to the perichondrium. pouch is the deep caudal incisure or antitra-
The pinnae are highly mobile and can be con gohelicine fissure.
trolled independently. The shape of the pinna is The anterior border of the auricular cartilage
characteristic of the breed. Some are small, is nearly straight. At the junction of the proximal
erect, and V-shaped as in toy terrier breeds; and middle thirds the spine of the helix or distal
some may be slightly tipped as in collies and crus of the helix is formed by an abrupt incisure
Irish terriers; others may be large and pendant of this border. The medial crus of the helix is
as in the hounds. separated from the tragus by the tragohelicine
The auricular cartilage is pierced by many incisure. The lateral crus of the helix arises
foramina which permit the passage of blood anterior to the medial crus and extends around
vessels. The skin covering the inner or concave the medial crus to overlap the anterior border of
surface of the pinna is firmly attached, and thus, the tragus.
when the ear is traumatized, hemorrhage may The groove or furrow bounded by the ant
occur between the skin and the cartilage. The helix, tragus, and antitragus, which continues
auricular cartilage is attached to the external into the external auditory meatus, is termed the
acoustic process of the temporal bone by means cavum conchae. The concha is the area of the
of a small annular cartilage. This is usually about auricular cartilage between the scapha and the
2 cm. long, presenting a lumen of 5 to 10 mm. in cartilaginous external acoustic meatus. A shallow
diameter. The description that follows is based groove is present on the medial surface of the
upon the pendant type of ear held erect. The auricular cartilage opposite the anthelix. This
isolated cartilages will be described separately. is the anthelicine sulcus. The bend in the pinna
The opening of the ear canal faces dorsolater- of lop-eared animals occurs distal to the anthelix
ally. The apex of the pinna points dorsally, the in the scapha. The skin lining the scapha and
convex or outer surface faces medially, and the concha shows pigmentation characteristic of the
concave or inner surface faces laterally. Thus breed. It has, in most specimens, a decreasing
the margins of the pinna are anterior (rostral) amount of hair from the distal to the proximal
and posterior (caudal) in the description. The parts. A few very fine hairs are found at the
elastic cartilage is thin and pliable, and at its entrance of the cartilaginous external acoustic
proximal end it thickens where it is rolled into meatus. The rather prominent transverse ridges,
the form of a tube. The term “helix” is applied as well as the longitudinal ridges, are simple skin
to the slightly folded free margin of the cartilage. folds which frequently continue toward the apex
A low transverse ridge, the anthelix (Fig. 17-10), of the ear. Cartilage is not discernible in them
is present on the medial wall of the initial proxi upon histological section. The protective hair of
mal part of the ear canal. The concave triangular the skin becomes fine and scanty in the conchal
area between the helix and the anthelix is the cavity.
T he E ar 849
F ig . 17-9. Transverse section through head showing ear canal. (Modified after Sis.)
850 C h ap ter 17. T he Sen se O rgans and In tegum en t
Scapha
P o s t e r io r b o r d e r of h e l ix
A n t e r i o r b o r d e r of hel ix
F o r am e n f o r b l o o d v e s s e l s
Poste rior p ro c e s s of h e l i x
P osterior in c is u re
■Tuber cl e o f a n t h e l i x
A n t i t ra g ic i n c i s u r e r ||| \
^ S p i n e of he lix
S tylo id p r o c e s s of) '' ^ ( d i s t a l crus o f h e lix )
a n ti tragus)
Med . c o r n u of a n t i tra g u s -
Trayohelicine in cisure
Lat. c r u s o f h e l i x
I nte rfracj i c in cisu re
T r a g us
A uricula r c artilag e
Interposed between the auricular cartilage in the deeper dermal layers. Nielsen (1953) is
and the external acoustic process is the annular of the opinion that the tubular and sebaceous
cartilage. This is a narrow band of cartilage glands in the ear are similar in structure to those
rolled to form a tube (Fig. 17-10). The proximal of the skin and that the normal ear secretion, the
end of the cartilage overlaps the osseous ex cerumen, is a product of both glandular types.
ternal acoustic process, with which it articulates Trautmann and Fiebiger (1957) also believe the
by means of ligamentous tissue. The ear canal, secretion (ear wax) to be a mixture of both types
therefore, is divided into lateral cartilaginous of glands. These authors also describe the great
and medial osseous parts. The space between vascularity of the subcutaneous tissue of the
the incomplete tubes of the auricular and an osseous meatus.
nular cartilages and between the annular carti
lage and the external acoustic process provides THE MIDDLE EAR
for freedom of movement of the pinna.
A small cartilaginous plate, the scutiform The tympanic cavity (cavum tympani) (Fig.
cartilage (Fig. 17-10), is located in the muscles 17-11) contains the auditory ossicles, the chorda
anterior and medial to the ear. This cartilaginous tympani nerve, muscles, and the auditory tube,
plate is shaped somewhat like a boot with the which communicates with the nasal pharynx.
heel directed away from the mid line. It is an The middle ear is lined with a mucous mem
isolated cartilage interposed in the preauricular brane that is, in general, covered with a two-
muscles, thus forming no part of the external ear. layered, columnar, ciliated epithelium, accord
Deep to the scutiform cartilage lies a fatty ing to Krolling and Grau (1960). The cavum
cushion, the corpus adiposum auriculi. This fatty tympani is divisible into a dorsal part, the epi-
pillar extends over a portion of the superficial tympanicum; a middle, mesotympanicum; and
surface of the temporal muscle and around the a ventral, hypotympanicum. The last corre
base of the auricular cartilage. sponds to the bulla tympanica. The tympanic
The bony external auditory meatus is lined membrane may be divided into two parts: the
with a thin cutaneous membrane which con pars flaccida and the pars tensa. The pars flac-
tains, in carnivores, large alveolar glands, accord cida is a small, triangular portion which lies
ing to Ellenberger and Baum (1943). The mus between the lateral process of the malleus and
cles of the external ear are described with the the margins of the tympanic incisure. The pars
muscles of the head on pages 140 to 144. The tensa constitutes the remainder of the mem
tympanic membrane (eardrum) which separates brane. The external aspect of the tympanic
the external ear from the middle ear is a thin, membrane is somewhat concave, owing to trac
semitransparent sheet oval in shape, and con tion on the medial surface by the manubrium
cave when viewed from the external aspect. Its of the malleus. The most depressed point, which
long axis is horizontal. The membrane is thin is opposite the distal end of the manubrium, is
centrally and becomes thicker near its periphery. termed the umbo membranae tympani. A light-
colored streak, stria malleolaris, may be seen
Glands of the External Auditory Meatus running dorsocaudally from the umbo toward
the pars flaccida when viewed from the external
The external auditory meatus presents a cu side. This is caused by the manubrium being
taneous lining which includes stratified squa partly visible through the tympanic membrane
mous epithelium, sebaceous and tubular glands, along its attachment. The manubrium is em
and hair. The conchal cartilage is covered with bedded in the tunica propria and is covered by
skin which, according to Fraser (1961), presents the epithelium lining the membrane. This in
fewer hair follicles on the concave inner surface turn is fastened to a collar of bone in the exter
than on the external surface. Both types of nal acoustic meatus. This bony collar is incom
glands are present in the cartilaginous and bony plete dorsocaudally, forming the tympanic
portions of the auditory meatus of the dog. incisure.
The sebaceous glands form a superficial The epitympanic recess is dorsal to a frontal
glandular bed immediately below the epithelial plane through the osseous external acoustic
surface, whereas the tubular glands are found meatus. It is the smallest of the three portions
in the deeper connective tissue layers. The and is occupied almost entirely by the head of
sebaceous glands are frequently associated with the malleus and the incus at their articulation.
hair follicles, whereas the tubular ceruminous The tympanic cavity proper is that portion
glands are located below the sebaceous glands adjacent to the tympanic membrane. It is ir-
852 C h ap ter 17. T he Sen se O rgans and In tegum en t
Semic i r c u l a r d u c t s x
Petrous te m p o r a l bone
phatic s p a c e o f vestibule
es
Endolym phatic
ncus
S cala ve stib u
Malleus
C o chle ar d u c t-
Scala t y m p a n i- -
E x t er n c
Dura m a te r - " '//■ a c o u s tic meatus
C ochlear aqueduct
Round window
Tympanic membrane
A u d ito ry tube
F ig . 17-11. Diagram of middle ear and inner ear. (From Getty et al. 1956.)
T he E ar 853
regularly quadrangular in shape, being flattened plexus arising from the tympanic branch of the
laterally by the tympanic membrane which glossopharyngeal nerve lies on the promontory
forms its wall. In the posterior portion, but fac and supplies the tympanic mucosa. Other nerves
ing anteriorly, is the secondary tympanic mem which contribute to this plexus are the small,
brane closing the round window (fenestra superficial petrosal and the caroticotympanic
cochlea) (Fig. 17-11). nerve.
The ventral portion, the part within the The auditory or Eustachian tube (tuba audi-
tympanic bulla, may be compared in shape to tiva) is a short canal which extends from the
the interior of an egg shell, having an elliptical nasal pharynx to the anterior portion of the
opening on the side which faces dorsally. It tympanic cavity proper. Its short bony wall is
communicates with the tympanic cavity proper formed anteriorly by the squamous part, and
through this opening. The long axis of the ventrally its floor is formed by the tympanic part
tympanic cavity is about 15 mm. in length and of the temporal bone. The lateral wall, which is
at an angle of about 45° with the sagittal plane about 8 mm. long, is nearly twice the length of
in a caudolateral direction. The width and depth the medial wall. The tube is oval in cross section
are about equal, measuring 8 to 10 mm. The with its greater diameter 1.5 mm. The medial
tympanic membrane is slanted ventromedially. wall of the membranous part of the tube is sup
Holz (1931) states that when viewed from the ported by a plate of hyaline cartilage, the an
front, a plane through the annulus tympanicus terior end of which curves medially to form a
is in general at a 57° angle with the frontal plane short hook.
in the dog. The tensor veli palatini muscle (Fig. 17-13)
Holz describes a membrana Shrapnelli (pars arises in the groove of the petrous temporal bone
flaccida of the tympanic membrane) which helps ventrolateral to the tensor tyrnpani muscle. It
to form the lateral boundary of an upper tym supports the lateral wall of the auditory tube.
panic pouch (Prussak’s pouch). This pouch lies The branch of the fifth cranial nerve which sup
dorsal to the lateral (short) process of the mal plies the tensor tyrnpani muscle enters the
leus and is bounded medially by the neck of the tympanic cavity in association with the tendon
malleus. In man there are relatively small open of origin of the tensor veli palatini muscle.
ings which communicate with the epitympani-
cum and the mesotympanicum, while in most Bones of the Middle Ear
animals Prussak’s pouch communicates freely (Auditory Ossicles)
with the tympanic cavity, according to Holz
(1931). In the dog, however, he describes a The auditory ossicles are three small bones
complete separation between Prussak’s pouch which transmit air vibrations from the tympanic
and the tympanic cavity. membrane across the cavity of the middle ear
On the medial wall of the tympanic cavity is a to the inner ear. The most lateral and largest of
bony eminence (promontorium) (Fig. 17-12), the three bones is the malleus (Fig. 17-11). The
which houses the cochlea; it lies opposite the most medial is the stapes. The handle of the
tympanic membrane medial to the epitympanic malleus attaches to the tympanic membrane.
recess. The oval or vestibular window (Fig. 17- The base of the stapes is attached to the margin
11 ), which is occupied by the base of the stapes, of the vestibular window. Between the malleus
is located on the dorsolateral surface of the and stapes is the incus.
promontory just medial to the pars flaccida. The The malleus consists of a head, a wide thin
ostium of the auditory tube (ostium tympanicum neck, and a manubrium or handle. The handle
tubae auditivae) is the anterior extremity of the is three-sided in cross section. The side em
tympanic cavity proper. The tendon of the bedded in the substance of the tympanic mem
tensor tyrnpani muscle (Fig. 17-13) descends brane is wider and smoother than the other two;
ventrolaterally through an arch in a thin lamina it is also slightly concave longitudinally. At the
of bone which overlies the muscle. It inserts on base of the manubrium, extending medially and
the muscular process of the malleus. The ossicles slightly anteriorly, is the muscular process of the
form a short chain across the dorsal part of the malleus. This is provided with a tiny hook at its
tympanic cavity. end to which the tensor tyrnpani muscle at
The tympanic nerve (chorda tyrnpani) (Fig. taches. The anterior process (Fig. 17-14, A) or
17-12), after leaving the facial nerve, passes long process is largely embedded in the tym
through the tympanic cavity medial to the panic membrane. It extends directly forward
malleus to join the lingual nerve. The tympanic from the neck of the malleus, arising at the same
854 C h apter 17. T he Sen se O rgans and In tegum en t
Vestib ular n.
F a c i a l n. i C o c h l e a r n.
' i
Scala ty m p a n i In ternal acoustic m e a tu s
O s s e o u s s p i r a l lQ m in a ^ f0 0 ^ ^
S e c o n d a r y s p i r a l I a m i na
Lam ina of m o d i o l u s — A
Cupula
Cut e d g e
of promontory
Chorda
tym pani
Tympanic
Tympan ic m e m b r a n e 1M a n u b r i u m of m alleu s
(p a r s tensa)
Fic,. 17-12. Sculptured medial view of the right middle ear and cochlea. (From Getty et al. 1956.)
T he E ar 855
Anterior s e m ic ir c u la r canal
L a t e r a l wall of v est ib ul
Floccular fossa
s L a t e r a l s em i c i r c u l a r c a n a l
F a c i a l n.
s S t a p e d i u s m.
Base of s t a p e s /
Lo n g c r u s of i n c u s s x
F acia l n.
f 'x Tym p a n o h y o i d
T e n s o r t y m p a n i m. ^ 1 c a r t i lage
Maj or superficial} u l o h y o i d e u s m.
p e t r o s a l n . j ^ ' i'i 1'
ik
/ 'A u d ito ry tu b e ^
' Te n s o r v e l i p a l a t i n i m.
F ig . 17-13. Sculptured medial view of the right middle ear showing auditory ossicles and their muscles. (From Getty et al. 1956.)
856 C h ap ter 17. T h e Sen se O rg an s and In te g u m e n t
level as the muscular process. Opposite the mus tympanic cavity. The body of the incus is
cular process at an angle of about 90° with the attached to the roof of the epitympanic recess
anterior process is the short, lateral process. This by the dorsal ligament of the incus. The posterior
is the most dorsal attachment of the manubrium ligament of the incus attaches the short crus of
to the tympanic membrane. The head of the the incus to the fossa incudis. An annular liga
malleus articulates with the body of the incus ment attaches the base of the stapes to the
in the epitympanic recess, the most dorsal por cartilage which lines the oval window. In addi
tion of the tympanic cavity. tion, there are interosseous ligaments which join
The incus (Fig. 17-14, C), measuring about 4 the ossicles together (Fig. 17-15).
mm. long and 3 mm. high, is much smaller than
the malleus. Its shape has often been likened to Muscles of the Ossicles
a human bicuspid tooth with divergent roots.
The incus lies caudal to the malleus in the Two tiny muscles (Fig. 17-13) are associated
epitympanic recess. The crura are located on with two of the ossicles. The tensor tympani is
each side of a transverse ridge which forms the spherical with its base in the fossa tensor
caudal limit of the recess. The short crus points tympani. The short tendon of insertion is at
caudally into the fossa incudis dorsal to this tached to the hook on the apex of the muscular
ridge. The long crus is also directed caudally, process of the malleus. Contraction of this mus
but presents a small bone, the os lenticularis, cle tends to draw the handle of the malleus
which extends anteriorly and somewhat medially medially, tensing the tympanic membrane.
from its distal end. In some instances this con Innervation is by a twig from the mandibular
nection ossifies to form the processus lenticularis. division of the trigeminal nerve. The stapedius
The stapes (Fig. 17-14, D) consists of a head, muscle is the smallest skeletal muscle in the
neck, two crura, a base, and a muscular process. body, and its origin is in the fossa musculae
It lies in a horizontal plane, the base facing medi stapedis. The body of the muscle lies largely
ally. The base articulates with the cartilage medial to the facial nerve. Its tendon of insertion
which covers the edge of the vestibular or oval attaches to the muscular process of the stapes
window (fenestra vestibulae). The stapes is the (Fig. 17-13). Contraction of the stapedius mus
innermost ossicle and is the smallest bone in the cle moves the anterior end of the base of the
body, being approximately 2 mm. in length. The stapes caudolaterally. This muscle is innervated
crura are hollowed on their concave or opposed by the stapedial branch of the facial nerve. The
sides. A cross section of a single crus appears as cavum tympani is lined with a thin mucous
a narrow semicircle of bone. There is a thin membrane that is partly covered with ciliated
connective tissue, obturator or stapedial mem epithelium. Some portions of the mucous mem
brane (membrana stapedis) which connects one brane are lined with a single layer of squamous
crus to the other. The anterior crus is slightly epithelium, according to Krolling and Grau
longer than the posterior crus. Arising from the (1960).
posterior crus near the neck is a minute muscular
process which provides attachment for the sta THE INNER EAR
pedius muscle (Fig. 17-14, D).
The internal ear (auris interna) consists of
fluid-filled ducts and sacs, the membranous
Ligaments of the Ossicles labyrinth, contained within an osseous labyrinth.
The structures of the inner ear may be separated
Several ligaments attach the ossicles to the into three parts: the cochlea, the vestibule, and
wall of the tympanic cavity. A short but fairly the semicircular canals. The anterior part is the
well-defined lateral ligament of the malleus con cochlea, the organ where mechanical stimuli are
nects the lateral process of the malleus to the converted to nerve impulses which, upon reach
margins of the tympanic notch. The dorsal liga ing the brain, result in audition. The posterior
ment of the malleus is a somewhat diffuse mass part consists of the three semicircular canals,
of ligamentous tissue which joins the head of each in a different plane. The third part, the
the malleus to a small area on the roof of the osseous vestibule, contains the utricle and sac
epitympanic recess. The anterior ligament of the cule. The semicircular canals and utricle are
malleus (Fig. 17-13) is a short ligament attach directly concerned with equilibrium. Perilymph
ing the anterior process of the malleus to the occupies a narrow space between the endo-
osseous tympanic ring just ventral to the canal by lymph-filled membranous labyrinth and the
which the chorda tympani nerve leaves the osseous labyrinth.
T he E ar 857
— Manu b ri u m
M anubrium -
A B
Short c r u s ~ - ^ ^ jj
^ Base
Posterior crus
A rtic u la r surface) JjL>
for m alleusj TK A n t e r i o r ^ [ X ^ V r - - ~ O b t u r a t o r membrane
crus— °f stapes
Lonq c ru s -^ ^ ^ ^ k ^ M Q
1 Head — xM u s c u l a r p r o c e s s
q Os l e n t i c u l a r i s '
D
F i g . 1 7 -1 5 . Auditory ossicles o f right ear, ventral aspect (tympani bulla removed). (From G etty e ta l. 1956.)
T he E ar 859
anterior canals. In sculptured specimens the an face of the apex of the parotid salivary gland. It
terior semicircular canal is seen to surround the leaves the parent vessel at a right angle approxi
floccular fossa, a small but deep depression on mately 0.5 cm. before the internal maxillary ar
the medial side of the petrous temporal bone. tery arises. But in 2 per cent of the ears examined
This depression is occupied by the paraflocculus by Sis (1962) the great auricular arose further
of the cerebellum. caudad, at a point where the hypoglossal nerve
The ampullated end of the posterior canal and passes over the external carotid artery. The great
the nonampullated end of the lateral canal are auricular artery gives off a small artery, the stylo
united for a short distance caudal to the vesti mastoid artery, as well as arteries to the parotid
bule. and mandibular salivary glands and to the neck.
Its three larger branches, lateral, intermediate,
Membranous Labyrinth and medial auricular rami (Fig. 17-18), ramify
on the convex face of the concha and anastomose
The membranous labyrinth (labyrinthus mem- with each other at the apex of the ear. The con
branaceus) does not completely fill the hollow cave surface is also supplied by smaller branches
system within the osseous labyrinth. Thus it is which freely encircle the auricular cartilage as
slightly smaller, but similar in shape. The fluid well as receiving branches from the convex side
perilymph surrounds it, and connective tissue which have passed through small foramina in
trabeculae support and attach it to the osseous the auricular cartilage.
wall. Spaces comparable to those in the sub Sis (1962) found that the vascular pattern on
arachnoid space exist among the trabeculae. the convex surface of the pinna varied. The lat
There are three regions which correspond to eral auricular ramus leaves the great auricular
those of the osseous labyrinth. They are the as a common trunk and divides into two vessels.
membranous cochlear duct, the membranous One vessel continues toward the apex of the
semicircular ducts, and the membranous vesti pinna, and the other anastomoses with the inter
bule (Fig. 17-16). The last, however, differs mediate auricular ramus. Sis found that the in
from the osseous vestibule, since the membra termediate auricular ramus always left the great
nous part is composed of two saclike structures, auricular halfway between the medial and lateral
the utricle and saccule, which occupy the lumen auricular rami. It did not, however, always con
of the osseous part. The membranous cochlea is tinue as a main branch to the apex of the pinna.
united to the sacculus by the ductus reuniens. In 50 per cent of the ears examined the inter
The cochlear duct is roughly triangular in cross mediate auricular ramus began as a small artery
section. The fibrous basilar membrane forms the supplying the postauricular muscles. It contin
floor of the cochlear duct, separating the endo- ued as a small branch until it received large
lymph of this duct from the perilymph of the anastomotic trunks from the lateral and medial
scala tympani. The very thin vestibular mem auricular rami approximately midway up the
brane forms the roof of the cochlear duct, sep pinna. From this point it extends in a straight
arating its cavity from that of the scala vestibuli. course to the apex of the ear. The medial auricu
The spiral organ (of Corti), the organ of hearing, lar artery is the last branch given off the great
lies upon the basilar membrane and consists of a auricular before it continues dorsad along the
thickened, specialized epithelium. nuchal crest as the occipital branch. The medial
auricular ramus runs along the medial margin of
the pinna to the apex, where it anastomoses with
BLOOD AND NERVE SUPPLY the intermediate ramus. The lateral, intermedi
ate, and medial auricular rami are accompanied
The great auricular artery is the chief blood by satellite veins.
supply to the external ear (Fig. 17-18). It arises A small branch, the deep auricular artery,
from the external carotid artery, which is the supplies the numerous muscles at the base of the
direct continuation of the common carotid. The auricular cartilage. It is given off between the
external carotid artery, after giving off several intermediate and medial auricular rami. Some
significant collateral branches, divides at the branches also pass through the foramina of the
caudal border of the mandible, ventroanterior pinna to supply the skin on the concave side.
to the annular cartilage of the ear, into superfi The superficial temporal artery, one of the ter
cial temporal and internal maxillary arteries. minal branches of the external carotid, arises at
The great auricular artery arises from the ex the caudodorsal border of the masseter muscle.
ternal carotid artery on the deep (medial) sur The anterior auricular artery (Fig. 17-18), which
860 C h ap ter 17. T h e Sen se O rg an s and In te g u m e n t
- - Z y g o m a t i c process
o f temporal bone
Cochlea^
Petrous temporal bone
I nt . a c o u s t i c m e a t u s -
Anterior s e m ic irc u la r canal
C ochlear n -
— Lateral s e m ic irc u la r canal
C ochlear a g u e d u c t-
Posterior s e m i c i r c u l a r c a n a l
V estibular agueduct-
F ig . 1 7 -1 6 . Phantom diagram o f right inner ear in situ, dorsal aspect. (From G etty et al. 1956.)
T he E ar 861
Cupula of cochlea
Oval window
Anterior am pul I a - ^
Lat. a m p u l l a - Si te of secondary
sp iral lamina
x Cochlear aqueduct
Round wi nd o w
I mm. Vesti b ul e
Vesti b u l a r a q u e d u c t
Anterior / Common c r u s
sem i c i r c u l a r c a n a l '
F i g . 1 7 -1 7 . R ight osseous labyrinth drawn from a la te x cast, lateral aspect. (From G etty et al. 1956.)
862 C h ap ter 17. T he Sen se O rgans and Integum en t
arises from the superficial temporal, is largely superficial temporal vein before the latter joins
a cutaneous branch to the skin of the lateral sur the internal maxillary vein. A cutaneous anasto
face of the base of the ear. Its terminal branches motic branch between the anterior and posterior
anastomose with the medial auricular artery. At auricular veins lying on the medial (convex) sur
this point a small arterial trunk can be seen to face of the base of the ear has been observed by
cross the margin of the auricular cartilage to Sis (1962).
supply the medial portion of the concave surface The ear of the dog is supplied by several cra
of the concha. The remaining portion of the nial nerves as well as a cutaneous cervical branch
concave surface of the auricle is supplied by from the ventral branch of the second cervical
branches of the auricular rami which freely en nerve. A branch of the second cervical (great
circle the margin of the auricular cartilage or auricular nerve) supplies largely the base of the
pass through the foramina from the convex side concha and the skin of the back of the neck. The
(Sis 1962). retroauricular nerve, a branch of the facial,
The anterior (medial), intermediate, and pos courses caudad and dorsal under the platysma
terior (lateral) auricular veins arise near the apex to become superficial at the base of the ear to
of the ear and parallel the comparable satellite supply the postauricular muscles. The auriculo-
arterial rami. The intermediate auricular vein palpebral nerve, another branch of the facial,
usually does not accompany the corresponding arises under the parotid gland ventral to the
arterial ramus as closely as the satellite medial conchal cartilage and dorsal to the internal maxil
and lateral rami accompany the corresponding lary vein. It courses dorsoanteriorly to divide
arterial rami. Differences can be observed in the into palpebral and anterior auricular branches.
veins between the ears of the same dog. Tribu The anterior auricular nerve supplies the pre
taries enter the vessels from the convex surface, auricular muscles. The auriculotemporal nerve,
while others enter by passing from the concave a branch of the trigeminal, lies dorsal to the cau
surface through the foramina of the auricular dal part of the masseter muscle, covered in part
cartilage or encircling its border. The posterior by the parotid gland. Branches from this nerve
auricular vein, which closely parallels the mar supply the skin in the anterior portion at the
gin of the pinna, enters the internal maxillary base of the ear and skin of the temporal region as
vein just dorsal to the mandibular salivary gland. well as the parotid salivary gland.
The anterior auricular vein terminates in the The labyrinthine or internal auditory artery,
T he N a sa l C a v it y 863
a branch of the basilar, is the sole supply to the daria, and mucosa of the tympanic cavity of man and the
labyrinth. The labyrinthine artery divides into dog. Z. Zellforsch. Bd. 39: 447-469.
--------------- 1955. Histologic studies of the autonomic and
the rostral vestibular artery, which supplies the cerebral innervation of the inner ear and its vessels in the
crista of the posterior semicircular canal and the case of man and the dog. J. cell Res. 42: 1-18.
common crus and a branch which runs in the Bast, T. H., and B. J. Anson. 1949. The Temporal Bone and the
vestibular aqueduct. A separate branch, which Ear. Springfield, 111., Charles C Thomas.
apparently leaves the labyrinthine artery, di Ellenberger, W., and H. Baum. 1943. Handbuch der vergleich
enden Anatomie der Haustiere. Berlin, Springer.
rectly supplies the cochlea. Shambaugh (1923) Fraser, G. 1961. The histopathology of the external auditory
states that the venous blood from the labyrinth meatus of the dog. J. comp. Path. 71: 253-258.
is collected into trunks. The larger one leaves the Getty, R., H. L. Foust, E. T. Presley, and M. E. Miller. 1956.
internal ear along the cochlear aqueduct and Macroscopic anatomy of the ear of the dog. Amer. J. vet.
Res. 17: 364-375.
drains the cochlea. The smaller one leaves along Gray, A. A. 1907. The Labyrinth of Animals, Vol. I. London,
the vestibular aqueduct. I. and A. Churchill.
The acoustic nerve supplies the internal ear. Holz, K. 1931. Vergleichende anatomische und topographische
It divides into two branches in the depths of the Studien liber das Mittelohr der Saugetiere. Z. Anat.
internal acoustic meatus. These are the cochlear Entwickl-Gesch. 94: 757-791.
Honda, Y. 1908. Gehororgan des Hundes. Inaugural Disserta
nerve, which supplies the cochlea, and the ves tion. Erlangen, Junge und Sohn.
tibular nerve, which supplies the remaining part Krolling, O., and H. Grau. 1960. Lehrbuch der Histologie und
of the membranous labyrinth. For further de vergleichenden mikroskopischen Anatomie der Haus
tailed histological studies of the innervation of tiere. Berlin, Paul Parey.
the inner ear and its vessels the reader is referred Miller, M. E., and R. Witter. 1942. Applied anatomy of the ex
ternal ear of the dog. Cornell Vet. 32: 64-86.
to the work of Andrzejewski (1954, 1955). Nielsen, S. W. 1953. Glands of the canine skin—morphology
and distribution. Amer. J. vet. Res. 14: 448-454.
Shambaugh, G. E. 1923. Blood stream in the labyrinth of the
ear of dog and man. Amer. J. Anat. 32: 189-198.
Sis, R. F. 1962. Polytetrafluoroethylene in Reconstructive Sur
gery of the Canine External Acoustic Meatus. M.S. The
BIBLIOGRAPHY sis, Iowa State University, Ames.
Trautmann, A., and J. Fiebiger. 1957. The Histology of Do
Andrzejewski, C. 1954. The finer histology of the nervous tis mestic Animals (translated and revised by R. Habel and
sue in the membrana tympani, membrana tympani secun E. Biberstein). Ithaca, N. Y., Comstock Pub. Asso.
T H E NASAL CAVITY
The nasal cavity is divided into right and left longed anteriorly as the cartilaginous nasal sep
halves by the nasal septum (septum nasi). Each tum (cartilago septi nasi).
half may be divided into two regions: the respir NASAL TURBINATES
atory region (regio respiratoria) and the olfactory The nasal turbinates, covered by nasal mu
region (regio olfactoria). Longitudinally, the cosa, occupy the major portion of each half of
nasal cavity extends from the nostrils or nares the nasal cavity. They include the nasoturbinate,
to the posterior openings or choanae. Each nasal maxilloturbinate, and ethmoturbinates (see pp.
cavity is further divided by the nasal conchae 24 to 31 and Figures 1-17 to 1-21).
(conchae nasales) into dorsal, middle, ventral, The nasoturbinate, or dorsal nasal concha
and common nasal meatuses (Figs. 1-21,17-19). (concha nasalis dorsalis), is composed of the
The structure of the external nose is described crista nasoturbinata, a thin shelf of bone, and
with the respiratory system on pages 713 to 718. the scroll of endoturbinate I. Although the osse
The bony nasal septum (septum nasi osseum), ous portion of the nasoturbinate ends at the level
which divides the nasal cavity into two portions, of the second premolar tooth, a mucosal fold
is composed of a perpendicular plate which continues into the vestibule of the nose.
joins the vomer below and the septal processes The maxilloturbinate, or ventral nasal concha
of the frontal and nasal bones above. It fuses (concha nasalis ventralis), occupies the anterior
posteriorly with the cribriform plate and is pro part of the nasal cavity, extending from the level
864 C h ap ter 17. T he Sen se O rgans and Integum en t
of the first to the third premolar teeth. Each The nasal cavity is richly supplied with blood
maxilloturbinate is formed by an intricately vessels and nerves. The nerves which supply the
folded series of lamellae which begin as a com nasal mucosa are the olfactory nerves, and
paratively tightly wound structure on the me branches from the ophthalmic and maxillary
dial surface of the maxilla. divisions of the trigeminal nerve. According to
The ethmoturbinates (ethmoturbinalia) are Read (1908), the olfactory nerves supply about
composed of numerous delicate bony scrolls half of the ethmoturbinates, one-third to one-
which attach to the external lamina and cribri half of the mucosa of the nasal septum, and the
form plate. It is customary to divide the ethmo roof as well as the lateral wall of the nasal cavity.
turbinates into four long, deeply lying endo-
turbinates (endoturbinalia I to IV) and six
smaller, more superficially located ectoturbi-
nates (ectoturbinalia I to VI). For details of the NASAL MEATUSES
arrangement of this ethmoidal labyrinth see
page 24. The first endoturbinate forms the bony The turbinates fill the nasal cavities so com
scroll for the concha nasalis dorsalis, while the pletely that only a narrow sagittal cleft remains
second endoturbinate forms the concha nasalis between the median perpendicular septum and
medialis (Graeger 1958). the maxilloturbinate of each side. This space is
In the dog, one scroll of the ethmoturbinate known as the common nasal meatus (meatus
extends for a short distance into the funnel-like nasi communis).
opening of the frontal sinus (Figs. 12-22 and 17- The dorsal nasal meatus (meatus nasi dorsalis)
20). Olfactory nerves ramify in this scroll, ac is a cleft between the shelflike nasoturbinate and
cording to Read (1908). They also extend for the nasal bone. It is formed by endoturbinate I.
some distance into the mucosa covering the bony As a narrow channel it almost disappears at the
wall of the sinus opposite the cribriform plate. level of the third premolar tooth, where it widens
The epithelium on the olfactory part of the sinus and continues between the scrolls of the ethmo
has a brown color. turbinates.
Dorsal meatus
Common
meatus - - M i d d l e m eatu s
Vomeronasal
o rg a n ' — Canine tooth
F ig . 17-20. Sagittal section of skull with nasal septum removed, revealing endoturbinates I to IV.
Frontal sinus
Ectoturbinate
(Ethmoidal
labyrinth)
Endoturbinate
Periorbital fat
- —-Lamina lateralis
I— molar
/ 1Hard palatex
To respiratory pharynx N Lamina transversalis
F ig . 17-21. Transverse section of left nasal cavity at level of first molar tooth.
F ig . 1 7 -2 2 . Transverse section o f left nasal cavity at level of third prem olar tooth.
866 C h ap ter 17. T he Sen se O rgans and Integum en t
The middle nasal meatus (meatus nasi me narrows prior to joining the vomeronasal organ.
dius) is the space between the nasoturbinate and In the region where the duct joins the organ, a
the maxilloturbinate. It extends to the cribri semicircular hyaline cartilage envelops it on the
form plate, and its osseous base is formed by lateral side and then arches over the organ to
endoturbinate II. Posteriorly, the middle nasal continue along only the medial surface as the
meatus divides into a dorsal or ethmoidal part vomeronasal cartilage (cartilago vomeronasalis).
which communicates with the ethmoturbinates The nerve to each vomeronasal organ origi
and a ventral part which continues into the naso nates from the medial side of the olfactory bulb.
pharyngeal meatus. It passes through one of the foramina of the
The ventral nasal meatus (meatus nasi ven cribriform plate (Fig. 1-9) at a point approxi
tralis) is the passageway between the maxillo mately halfway between the ventral and dorsal
turbinate and the bony palate or floor of the borders of the plate. Of the two branches which
nasal cavity. Posteriorly, it blends with that part emerge from the cribriform plate on each side,
of the middle meatus which continues into the one divides into two (approximately 2.5 cm.
nasal pharynx via the nasopharyngeal meatus. from its emergence) and supplies twigs which
ramify throughout the length of the organ. The
NASOLACRIMAL DUCT other branch terminates in the anterior portion
of the vomeronasal organ. A comparatively large
The nasolacrimal duct (ductus nasolacrimalis) branch of the palatine nerve can be traced to
lies initially within the bony lacrimal canal the posterior ventral portion of the organ, and
(canalis lacrimalis). It begins in the lacrimal bone branches of the nasopalatine nerve via the
(Fig. 1-27) at the fossa for the lacrimal sac, passes sphenopalatine nerve also supply it. According
ventroanteriorly through the lacrimal bone, and to Read (1908), the vomeronasal organ is inti
continues in a canal or groove on the medial sur mately connected with the olfactory sense, and
face of the maxilla. It opens ventral to the basal McCotter (1912) states that the vomeronasal
lamina of the maxilloturbinate scrolls. The open nerves are branches of the olfactory nerves. The
ing of the duct is difficult to reach clinically from presence of the nervus terminalis in the dog was
the nasal fossa. first described by McCotter (1913). He observed
that the vomeronasal nerves coursed almost hori
NASOPALATINE DUCT zontally across the medial aspect of the olfactory
bulb to its caudal border. Here the nerves
The nasopalatine duct (ductus nasopalatinus) formed a fine plexus and turned dorsolaterally.
passes through the palatine fissure and connects Connected with this plexus, a single trunk of
the nasal and oral cavities. It also communicates fibers extended posteroventrally on the medial
with the vomeronasal organ at its dorsal or nasal surface of the olfactory peduncle where it ap
orifice. The oral orifice of each duct lies lateral peared to enter the brain some distance from the
to the incisive papilla behind the central incisor olfactory bulb. Thus, according to McCotter,
teeth. The paired ducts run caudodorsally, at a the filaments of the nervus terminalis can be
45° angle with the palate, to open in each nasal seen separating from the vomeronasal nerves.
fossa. Kadowaki (1959) described the histology The histology of the vomeronasal organ has
of the nasopalatine duct in the fetal dog. been described by Kadowaki (1959). He found
that it was largely lined with olfactory neuro
VOMERONASAL ORGAN epithelium on its medial wall and with a thinner
cylindrical epithelium on its lateral wall.
The paired vomeronasal organ (organum
vomeronasale), or organ of Jacobson, is an iso PARANASAL SINUSES
lated area of olfactory membrane located in the
anterior base of the nasal septum as a tubular The paranasal sinuses are air-filled cavities
pocket partially enclosed by a scroll of cartilage which are often invaded by the nasal turbinates
(Figs. 10-1, 17-19). It is connected with the (Figs. 1-35 to 1-37). They are located in the
nasopalatine duct. In almost all groups of mam maxilla, frontal, and sphenoid bones. All para
mals the vomeronasal organ persists as a func nasal sinuses are small at birth, enlarge with age,
tional organ and communicates with the oral and are lined by a mucoperiosteum which may
cavity or the nasal cavity, or with both cavities. include olfactory elements. The sinuses may be
Dorsal to its origin in the mouth, each naso divided into several compartments which drain
palatine duct expands considerably and then directly or indirectly into the nasal cavity. A
T he N a sa l C a v it y 867
description of the paranasal sinuses is given on cribriform plate. The olfactory epithelium is ex
page 49. tremely thick in comparison with the areas pre
senting the simple ciliated epithelium. Kawata
INNERVATION OF THE NASAL CAVITY and Okano (1961) emphasized that considerable
differences exist in olfactory morphology be
The nasal fossa is lined by a mucoperiosteum tween the dog and the cat.
which is richly supplied by blood vessels and The terminations of the sensory fibers to the
nerves. The nerves which supply the nasal mu nasal and oral cavities of the dog have been ex
cosa include the olfactory nerve (I) and branches tensively investigated. It has been reported that
of the ophthalmic and maxillary divisions of the the sensory fibers are chiefly formed in the tunica
trigeminal (V). The olfactory nerves are rela propria of the mucous membrane (Abe 1954).
tively large and numerous in the dog. About half The author also noted, however, that some sen
of the ethmoturbinal folds and all the folds of sory fibers penetrated further into the epithelium
mucosa adjoining the cribriform plate are olfac to end as intraepithelial fibers.
tory, according to Read (1908). The anterior
ethmoidal branch of the ophthalmic nerve in ARTERIES OF THE NASAL CAVITY
nervates the septum (Fig. 10-1) and olfactory
The external ethmoidal artery enters the cra
folds. Read states that the sphenopalatine nerve nial cavity and anastomoses with the internal
innervates the mucosa anterior to the ethmo ethmoidal artery to form the ethmoidal rete on
turbinal folds, the maxillary sinus, the lateral the cribriform plate. Branches from the rete pass
wall of the nose, and the maxilloturbinal folds. through the cribriform plate to supply the cau
Histologically, the epithelium of the olfactory dal portions of the ethmoturbinates and a por
region consists of three kinds of cells: the sup tion of the nasal septum. See discussion on page
porting or sustentacular cells, the receptor or
316 and Figures 4-22 and 4-24.
olfactory cells, and the small stellate, basal cells.
The sphenopalatine artery arises in the ante
Serous glands are found in the submucosa. The
rior part of the pterygopalatine fossa and passes
axons of the olfactory cells come together in the
through the sphenopalatine foramen into the
submucosa to form the olfactory nerves. The ol
nasal cavity accompanied by its satellite nerve
factory nerves pass through the foramina of the
and vein. It divides into several branches and is
cribriform plate to the olfactory bulb.
distributed to the mucous membrane of the floor
Kawata and Okano (1959) examined the sen
and lateral wall of the nasal cavity, the maxillo
sory innervation of the ethmoturbinates of the
turbinates, the maxillary sinus, and part of the
dog and demonstrated olfactory cells. They also
ethmoturbinates. For details see page 309 and
demonstrated olfactory cells in the mucous mem
Figures 4-24 and 4-25.
brane of a limited area of the septum nasi of the
horse and the dog. They describe a special ol BIBLIOGRAPHY
factory cell whose peripheral part is thick, pre
senting a swollen tip shaped like a pineapple. Its Abe, Y. 1954. Fine structure of nasal and oral cavities in dog
proximal end rapidly tapers into a fiber of the and their sensory innervation, especially on the intraepi
olfactory nerve. The olfactory cells are distrib thelial fibers. Tohoku J. exp. Med. 60: 115-128.
Graeger, K. 1958. Die Nasenhohle und die Nasennebenhohlen
uted between the sustentacular cells. The shape beim Hund unter besonderer Berucksichtigung der Sieb-
and length of the olfactory cells are variable, beinmuscheln. Dtsch. tierarztl. Wschr. 65: 425-429,468-
depending upon their site. The above-mentioned 472.
authors arbitrarily divided the region of the Kadowaki, S. 1959. On the nerve supply of the nasopalatine
duct and the Jacobson’s organ of dog in the later fetal
ethmoturbinates macroscopically into three stage. Arch. Hist. Jap. 17: 437-458.
equal portions, I, II, and III, from anterior to Kawata, S., and M. Okano. 1959. Histological analysis of sen
posterior. The distribution of the olfactory cells sory nerve of the ethmoid bone of the dog. Arch. Hist.
is poor in part I, moderate in part II, and rich in Jap. 17: 609-615.
--------------- 1961. Histological analysis of sensory nerve of the
part III. The lamina propria presents numerous
nasal cavity of the cat. Wien, tierarztl. Mschr. (Festschrift
olfactory glands, blood vessels, and a rich plexus Schreiber 1960) 68-81.
of nerve bundles. McCotter, R. E. 1912. Vomeronasal nerves. Anat. Rec. 6 :299-
In the region of the cribriform plate the com 318.
paratively thin nerve bundles join to form large ------------ — 1913. The nervus terminalis in the adult dog and
cat. J. comp. Neurol. 23: 145-152.
nerve trunks. The nerve fibers are largely un
Read, E. A. 1908. A contribution to the knowledge of the ol
myelinated. Myelinated nerve fibers are some factory apparatus in dog, cat, and man. Amer. J. Anat. 8:
times seen passing through the openings of the 17-47.
868 C h ap ter 17. T he Sen se O rgans and In tegum en t
T H E ORGAN O F TA STE
The tongue of the dog consists largely of stri be found behind the vallate papillae among the
ated muscle and intermuscular connective tissue conical papillae. The taste buds are situated on
which attaches the mucous membrane to the the summits of these papillae (Fig. 17-26).
muscular mass. The tongue of the dog has great These papillae are covered by a stratified
mobility. A median groove is located on the dor squamous epithelium which is slightly cornified.
sum of the tongue, and five types of lingual papil The epithelium is very thin. The blood in the
lae have been described. These papillae, which vessels beneath the epidermis gives the fungi
are important structures of gustation, are lo form papillae a rose color in the living state. The
cated for the most part on the surface of the dermal core of the papillae is made up of dense,
tongue, the dorsal surface of the epiglottis, and white fibrous connective tissue irregularly ar
the surface of the soft palate. A substance to be ranged. It is not, however, as dense as the dermis
tasted must be in solution; therefore taste buds underlying the skin. Numerous myelinated nerve
are located only on moist mucous membranes. trunks enter the dermal cores of fungiform papil
There is a great deal of overlap of gustation and lae. These nerve fibers subserve the gustatory
olfaction, since the sense of smell combines with cells located in the taste buds found in the epi
the sense of taste in many taste sensations. The thelium of the fungiform papillae. They also
five different types of lingual papillae in the dog supply the perigemmal nerve endings outside
are: filiform, fungiform, vallate, foliate, and the taste bud. In the anterior two-thirds of the
conical. tongue these nerve fibers originate from the
lingual branch of the trigeminal nerve and from
FILIFORM PAPILLAE the chorda tympani nerve (a branch of the facial
nerve).
The term “filiform,” or thread-shaped, is
The taste buds are located on the dorsal sur
really a misnomer when considering the papilla
face of the fungiform papillae. They penetrate
as a whole. They are distributed on the anterior
the complete thickness of the epithelium and
two-thirds of the tongue. Figure 17-23 illustrates
rest on small, secondary dermal papillae. Not all
the papillae with the primary, secondary, and
fungiform papillae have taste buds associated
tertiary filiformes.
with them.
There is an average of four tertiary filiform
papillae on the anterior border of the filiform
papillae proper. The dermal cores do not extend FOLIATE PAPILLAE
into these keratinized papillae very far in the
dog. There are two secondary filiform papillae
on each side of the large primary filiform papilla. These leaflike papillae are described in most
The secondary filiform papillae have a well-
textbooks as being two in number; however,
developed dermal core supporting the keratin
each outpocketing of the epithelium and dermis
ized, stratified squamous epithelium. The pri
resembles the other papillae of the tongue.
mary filiform papilla is the largest and is single.
There are two groups of foliate papillae, with
It has a well-developed dermal core and a tough,
eight to twelve in each group (Figs. 17-27, 17-
cornified layer of epithelium covering its tip. The
28), located on the dorsolateral aspect of the
three types of filiform papillae are directed
tongue anterior to the anterior pillar of the
caudally and cover the normal tongue like rows
fauces.
of shingles. Their function is one of protection
Grossly, the papillae are arranged like the
and possibly the holding of food material. There
petals of a flower. There is a definite point
are no taste buds on the filiform papillae. The
toward which the basal portion of each group
stratified squamous epithelium is very thick.
of papillae converges.
Separating the papillae is a crypt (Fig. 17-29),
FUNGIFORM PAPILLAE which is deepest in the middle of the papilla and
very shallow dorsally and ventrally. The ducts
These mushroom-shaped papillae are located of subepithelial serous glands empty into the
on the anterior two-thirds of the tongue among bottom of these crypts between the foliate
the filiform papillae (Fig. 17-25). They may also papillae. Taste buds for the most part are located
The O rg an o f T a ste 869
S econdary filiform
papillae
Tertiary filiform
papillae
- Base of papilla
Vallate papillae
M e d ia n gro ove
A re a of fo lia t e papillae
Fungiform papilla
■Taste pores
Filiform papillae
Conical papillae
Tonsil
V allate papilla
Fungiform papilla
Filiform papillae
Fo liate papillae
F ig . 17-27. Base of tongue showing location of foliate, vallate, filiform, fungiform, and conical papillae, right lateral aspe
L y m p h a tic tissue
Serous glands
A
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W
** r.\ I J- t-/ i fr
' # /
O
f - U ,
iVx A
w,v m l> ?
■)'.« /.y«
* 4r-
,%x^
>*w
'4
>
X- M
'QV <*
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' c • ?; ^
r» *> * >' < ,
V' ‘t e V.fcT’ ■ 1 ■
fofV-* -
4 ,/
#4 **•£>
V
L'i
F ig . 17-29. Frontal section of tongue showing taste buds on foliate papillae.
S e c o n d a ry pap illa
Vallate papilla m o at
Vallate papilla
M o d ified conical
pap illae
Conical papillae
with entodermal epithelium. In the adult animal The vagus nerve supplies the fourth branchial
the junction between the ectodermal and ento arch, which forms most of the epiglottis and
dermal epithelium can be seen as an uneven line larynx. Bowne (1956) resected the vagus nerve
just in front of the rows of vallate papillae (Fig. just distal to the nodose ganglion and noted the
17-24). The epiglottis is derived from the third loss of taste buds on the affected side, while taste
and fourth branchial arches, while the soft palate buds were demonstrated on the unaffected side
is derived from the maxillary processes. in rare instances.
There are three cranial nerves directly associ The chorda tympani branch of the facial nerve
ated with the taste buds, namely, the chorda innervates the taste buds on the anterior two-
tympani branch of the facial nerve, the glosso thirds of the tongue of the dog. The sensory
pharyngeal nerve, and the vagus nerve. The tri branches of the glossopharyngeal nerve inner
geminal nerve also supplies sensory innervation vate the taste buds found in association with the
to the tongue, but according to Olmstead (1921, foliate and vallate papillae. It is thought that the
1922), it does not supply the nerve fibers to the vagus nerve supplies the taste buds on the epi
taste cells. glottis and larynx. The glossopharyngeal nerve
The facial nerve supplies the second branchial may also innervate the taste buds that are lo
arch. Fibers of the chorda tympani branch of cated on the soft palate.
the facial nerve have been traced from the According to Murray and Murray (1960),
geniculate ganglion, through the middle ear to nerves ending within the taste buds are widely
its junction with the lingual branch of the man distributed, but they seldom lie close to the taste
dibular nerve. Resection of the chorda tympani pit. The nerve fibers do not branch after they
nerve as it traverses the middle ear causes the have entered the taste bud. The fibers, after
taste buds, innervated by the sensory portion of losing their myelin sheath, appear to be en
the chorda tympani nerve, to degenerate. Taste closed within the cytoplasm of the gustatory
buds located on the fungiform papillae anterior cells. The cytoplasm of the taste cells frequently
to the vallate papillae disappear on the affected is folded over in a double layer, separating the
side after resection of the chorda tympani nerve. fiber from the intercellular space. (For additional
There is a crossing over of nerves, however, or a information on the tongue, see pages 654 and
dual innervation of the taste buds located on the 656.)
fungiform papillae close to the lingual groove.
The sensory portion of the glossopharyngeal BLOOD VESSELS AND NERVES
nerve arises from the superior and petrosal gan
glion in man (Arey 1954). The sensory rami pass The hypoglossal nerve, which is the sole motor
to the second and third branchial arches, and supply of the tongue, enters the muscle of the
eventually innervate most of the root of the tongue medial to the styloglossus muscle.
tongue and pharynx. Resection of the glosso The blood vessels pursue a tortuous course,
pharyngeal nerve as it leaves the petrosal gan and the veins are located in tissue spaces that
glion causes the taste buds located on the vallate permit venous engorgement. The lingual artery,
papillae, the trench wall surrounding the vallate one of the largest collateral branches of the ex
papillae, and those located on the foliate papil ternal carotid, passes anteroventrad in front of
lae on the denervated side to degenerate. There the hypoglossal nerve. The lingual artery enters
is no central vallate papilla in the dog as there is the tongue by passing medial to the hypoglossal
in the human being, and no dual innervation of muscle and lateral to the hyoid bone. The right
the right and left vallate papillae could be dem and left genioglossal muscles separate the lin
onstrated (Bowne 1956). The sensory portion of gual artery from its fellow of the opposite side.
the glossopharyngeal nerve innervates the soft Anastomoses exist between the lingual arteries
palate. Taste buds have been demonstrated his as well as between the sublingual and lingual
tologically in rare instances on the soft palate of arteries.
the dog.
The sensory innervation of the dog tongue,
particularly in reference to the development of
sensory fibers to the taste buds, has been exten BIBLIOGRAPHY
sively investigated (Okano 1953). The innerva
tion of taste buds in the larynx of the dog has Arey, L. B. 1954. Developmental Anatomy. 6th ed. Philadel
also been studied (Koizumi 1953). phia, W. B. Saunders.
T he Integum en t 875
Bowne, J. G. 1956. Macroscopic and microscopic structure taste buds of rhesus and cynoinalyus monkeys. Anat. Rec.
and age changes in the lingual papillae of the dog. M.S. 138: 211-233.
Thesis. Iowa State University, Ames, Iowa. Okano, S. 1953. Innervation, especially sensory innervation of
Koizumi, H. 1953. On innervation of taste buds in larynx in dog tongue. Tohoku J. exp. Med. 57: 169-179.
dog. Tohoku J. exp. Med. 58: 211-215. Olmstead, J. M. D. 1921. Effect of cutting the lingual nerve of
Kolmer, W. 1927. Geschmacksorgan. Hanb. d. Mikroshop. the dog. J. comp. Neurol. 33; 149-155.
Anat. d. Menschen, W. V. Mollendorf. 3: 154-191. —-— -------- 1922. Taste fibers and the chorda tympani nerve.
Murray, R. G , and A. Murray. I960. The fine structure of the J. comp. Neurol. 34: 337-341.
T H E IN TEG U M EN T
The skin is more than a covering envelope for specialized tactile or sinus hairs on the muzzle
the body. In addition to separating the fluid body as well as enlarged hairs (vibrissae) on the man
from its dry environment or hypotonic or hyper dible (submental) and above the eye (super
tonic fluid surroundings (it is nearly waterproof), ciliary). There are usually two tubercles (genal)
the integument protects against trauma, tem on each side of the face from which longer hairs
perature change, and entrance of disease-pro- grow. A tuft of hairs (interramal) stands out from
ducing organisms. The glandular elements of surrounding hair between the mandibles. The
the skin perform a secretory and excretory func specialized hairs of the eyelids (eyelashes) are
tion. This reaches its greatest development in the stiff and longer than other hairs. The ventral
mammary glands. The skin is the location of body surface is characterized by the median
vitamin D synthesis, and the subcutaneous tis raphe of the linea alba, the hairless umbilicus
sues serve as a reservoir for fat storage. As a and nipples, and the sparsely haired mammary
sensory organ the skin is important for the per glands, which are arranged in two rows with four
ception of touch, pressure, heat, cold, and pain. to six in each row. The skin is thickest over the
The skin of the dog serves a heat-regulating func neck and back (dorsum), where it is loosely at
tion, but not to the same degree as in some other tached.
animals, which are unable to bring about heat All skin areas are made up of epidermis and
loss by rapid respiration or panting. The skin, dermis. When the skin is removed during dissec
hair, and subcutis of the newborn puppy repre tion, subcutaneous tissue and cutaneous muscle
sent 24 per cent of the total body weight. This often remain with the skin, although they are
percentage is reduced to 12 per cent at maturity. not components of the skin. The thickness of
The integumentary system is involved in specific the various layers of the skin varies greatly from
dermatitis and also may reflect the general health one area to another. Epling (1953) and Webb
of a dog (icterus, cyanosis, dry scaly skin). Be and Calhoun (1954) have contributed much to
cause of variation in pigmentation, hair type, and the microscopic anatomy of canine skin.
skin thickness, there is a great range of color and
hair length in different breeds. NASAL SKIN
The skin is continuous at the natural body
openings with the mucous membrane of the di The nasal skin is usually heavily pigmented,
gestive, respiratory, and urogenital tracts, and tough, and moist. On close examination of the
with the conjunctiva of the eye. The hair is most surface of the planum nasale, shallow grooves
dense /on the dorsal and lateral portions of the are observed, which divide the surface into
body, while the abdomen, the inside of the polygonal, plaquelike areas, and give the nasal
flanks, the inside of the ears, and the under side skin an irregular appearance (Fig. 17-34, B).
of the tail are sparsely haired. Some areas, such Histologically, no glands can be demonstrated
as the nasal epidermis and carpal, metacarpal, in the epidermis or dermis (Fig. 17-34, C). The
tarsal, metatarsal, and digital foot pads, are dermis is composed of reticular, collagenous, and
hairless. These areas are modified and thickened elastic fibers, together with fibroblasts, blood
for specific functions. The claws or horny cover vessels, and nerves. The blood vessels and nerves
ing of the third phalanges of the digits are modi are larger in the deeper layers of the dermis than
fied for digging and defense. There are large in the more superficial layers. Directly under
876 C h apter 17. T he Sen se O rgans and In tegum en t
the epidermis the dermal papillae (Ham 1961) There are also secondary dermal papillae within
or papillary bodies (Trautmann and Fiebiger the conical structure.
1957) form an irregular line of attachment be The epidermis of the foot pad, which averages
tween the dermis and epidermis. The epidermal 1800 microns in thickness in the adult dog, is
surface is marked by deep grooves which divide composed of five layers: stratum cylindricum,
it into polygonal areas (Fig. 17-34, B). stratum spinosum, stratum granulosum, stratum
The epidermis of the nasal skin, which aver lucidum, and stratum corneum. The stratum
ages 630 microns in thickness in adult dogs, is cylindricum and stratum spinosum are fre-
composed of three layers: stratum cylindricum, quendy referred to collectively as the stratum
stratum spinosum, and stratum corneum. The germinativum. The stratum cylindricum is made
stratum cylindricum of the epidermis rests on up of a single layer of basal cells resting on the
the condensed, thickened superficial portion of connective tissue of the dermis. The stratum
the dermis and consists of one layer of cylindrical spinosum is composed of ten to fifteen layers of
cells. The stratum spinosum is made up of ten to diamond- or dome-shaped cells. In both the foot
twenty layers of diamond-shaped, dome-shaped, pads and the planum nasale, cell outlines and
or flattened polygonal cells which have a lighter- intercellular bridges may be observed on the
staining cytoplasm than the cylindrical cells. In spinous cells. The stratum granulosum is made
heavily pigmented nasal skin there are many up of four or five layers of flattened cells which
pigment granules in the cytoplasm. There is no contain basophilic keratohyalin granules in their
stratum granulosum or stratum lucidum in the cytoplasm. The stratum lucidum appears as a
epidermis of the nasal skin. The more peripheral shiny acidophilic layer of homogeneous sub
spinosum cells apparently do not undergo stance with refractile droplets called eleidin
keratinization as they do in other regions of (Bloom and Fawcett 1962). The stratum corne
epidermis. Their cytoplasm becomes weakly um of the foot pads consists of a thick layer of
acidophilic, and the nuclei become pyknotic, keratinized, non-nucleated material thicker than
the cells flattening out into a squamous type. As all the cellular layers combined (Fig. 17-35, C).
they approach the surface, they remain as a thin, The excretory ducts of the merocrine sweat
atypical nucleated stratum corneum, four to glands of the foot pad become continuous with
eight cell layers thick. The stratum corneum of the epidermis at the very depths of the epidermal
the nasal skin is surprisingly thin (Fig. 17-34, Q . pegs where their epithelium joins with the
stratum cylindricum of the epidermis. The
FOOT PADS lumen of the excretory duct then follows a
tortuous path through the epidermal cells to the
The skin of the foot pads is usually heavily surface where the glandular secretion is ex
pigmented and is the toughest region of canine pelled.
skin. The surface of the pads is rough, owing to
the presence of numerous conical papillae which HAIRY SKIN
are heavily keratinized and are readily seen with
the naked eye (Fig. 17-35, B). When dogs are The basic unit of hair production is the indi
kept on concrete or rough surfaces, the papillae vidual hair follicle. Each hair shaft is produced
sometimes become worn smooth so that they in a sleeve of epithelium which is continuous
are truncated or rounded instead of conical in with the surface epidermis. The follicle wall is
shape. divided into two layers, the outer and inner root
The digital cushion or base of the foot pad is sheaths. The follicle attains its greatest diameter
made up of subcutaneous adipose tissue which at the base, where it is dilated to form a bulb.
is partitioned by reticular, collagenous, and Invaginating the bulb is the dermal papilla.
elastic fibers (Fig. 17-35, C). Many elastic fibers The root of each hair extends down the center
are present in the deeper layers. Merocrine of the follicle to the bulb, where the germinative
sweat glands and lamellar corpuscles are em epithelial tissue produces the material which
bedded in the adipose tissue. The excretory ducts develops into the keratin shaft. A rich network
of the merocrine sweat glands traverse the of arterioles and capillaries supplies the germina
dermis on their way to the surface of the epi tive epithelium as long as the hair is growing.
dermis. Directly under the epidermis, the con There are periods during which the growth of
nective tissue is dense and papillate, forming the hair is arrested. At this time there is a regres
conical dermal cores for the epidermal papillae. sion of the hair root, and the dead hair is held in
T he In tegum en t 877
P la n u m n a s a le
S u rfa c e g ro o v e
S tra tu m
c o rn e u m
- E p id e r m is
E p id e r m a l peg
D e rm is
F ig . 17-34. Surface contour and histology of planum nasale. (After Lovell and Getty 1957.)
A. Gross appearance of nasal skin
B. Polygonal plaquelike areas
C. Histological section
C h ap ter 17. T he Sen se O rgans and In tegu m en t
*
I:/
■ M e ta c a rp a l pad
D ig ita l pad
Claw
-S tra tu m corneum
— C o n n e c tiv e tis s u e
c o re of p a p illa
' V ■ -' - v OA V ' ' i
* t „>A, C*' ... "r “ v E p id e rm is o f p a p illa
s&msaBSwSi D e rm is of fo o tp a d
- S w e a t g la n d s
F ig . 17-35. Surface contour and histology of foot pads. (After Lovell and Getty 1957.)
A. Gross appearance of pads
B. Conical papillae arranged on surface of digital pad
C. Histological section of foot pad
T he Integum en t 879
the follicle completely disconnected from the of a puppy during the first few days after birth
inactive germinal matrix. After a variable time that there is only a single hair emerging from
the dormant germinal cells become active and each external follicle orifice in the skin (Fig.
enter a period of organogenesis in which a new 17-36, A). The satellite or accessory hairs de
hair root is regenerated and production of hair is velop later. At the age of three months two to
resumed. At this time the old dead hair will be five accessory hairs accompany the main hair
shed and replaced by the new hair. Growing hair in the follicle (Fig. 17-36, B). At six months of
follicles are said to be in anagen, quiescent ones age the number in each bundle is five to fifteen,
in telogen, and the period of transition between which is typical of the compound follicle of the
the two, catagen. According to Comben (1951), dog (Fig. 17-36, C). The primary hairs have a
the hair follicle of the dog may produce up to better developed nerve and blood supply than
0.18 mm. of hair shaft per day during the active the satellite hairs. As a general rule the coarser
stage of the cycle. The hair shaft consists of a hairs appear earlier than the fine hairs. The indi
central medulla, a thick cortex, which forms the vidual follicle may be straight or curved. The
bulk of the hair, and a single-layered cuticle on short-haired breeds show straighter and longer
the outside. follicles, and the long-haired breeds more curved
follicles. Curling of the hair has been thought to
Embryology of Hair Follicle be related to the curve of the follicle into a saber
shape. There may be other factors involved,
The first hairs to appear in the fetal dog are such as unequal lateral growth of the fiber and
in the region of the eyebrows, upper lip, and difference in rates of growth in the various skin
chin. These develop into vibrissae, which are layers.
specialized sinus hairs or tactile hairs. The fol It is possible for a hair follicle to form one type
licles of the general hairy skin develop later. In of hair at one stage and another type at another
the general development of the pelage the hairs stage. In the development of the dog the nature
are farthest advanced cephalad, and the de of the puppy hair changes, and in the young
velopment spreads caudad. The primary hair adult the fine, fluffy hair of the pup is replaced by
germs form more or less simultaneously at fairly a coarser hair in the same follicle.
even distances. As the skin grows, new primary In a young adult dog the hair growth is pro
germs develop among the earlier ones. This re fuse and abundant. In old dogs the hair cover
sults in the two, three, or four groups of follicles ing is thinner, the hairs are not as long, and
being clustered together. The triad arrangement frequently the coloring changes to gray. Also,
is most frequent. Later, usually after birth, the
the hairs are more brittle and are accompanied
secondary germs develop close to the primary by loss of flexibility of the skin and subcutaneous
ones and form the compound follicle arrange
tissue.
ment.
The first evidence of the follicle in the embryo
is seen as a thickening of the epidermis (pre Compound Hair Follicle
germ stage) at regular intervals. The pre-germ
stage passes rapidly into the hair-germ stage as The hairy skin of an adult dog contains bun
the basal cells become higher and the entire dles of hairs which share common openings in
structure protrudes downward into the corium. the surface. These bundles are usually arranged
From its point of origin the hair germ grows in groups of three oriented into irregular rows.
obliquely downward into the mesenchyme in The typical bundle consists of a group of under
the form of a solid column. This is called the hairs and a single, longer and stiffer cover-hair.
hair-peg stage. The advancing border enlarges, The cover-hair of the central bundle of a three-
becomes bulbous, and envelops part of the bundle group is coarser than those in the lateral
mesenchymal material which was pushed down bundles.
ahead of the invading epidermal cells. Later, the The hair shafts that share a common opening
hair bulb and the dermal papilla become differ in the skin are enclosed in a common follicle
entiated into the productive hair follicle com down to the level of the sebaceous glands. Below
plete with glandular and muscular accessories. this point the hair shafts branch away from one
another into their own individual hair follicle and
Development of Compound Follicle bulb (Fig. 17-36). In this way as many as fifteen
after Birth hairs may share a single external follicle orifice.
The individual follicle and hair bulb of the cover-
It can be observed by examining the hair coat hair or guard-hair is larger and penetrates more
880 C h apter 17. T he Sen se O rgans and In tegum en t
deeply into the subcutaneous tissue than those of (1961) suggest that the study of skin gland se
the satellite or subsidiary hairs. There are breed cretions be labeled exocrinology as compared to
variations in the number of follicle bundle com endocrinology. Exocrinology is the study of sub
plexes per square centimeter and also in the stances of external secretion which arouse a
number of hairs in each bundle. Brunsch (1956) sexual response in other individuals. Parks and
found that smooth-haired dachshunds, smooth Bruce have made observations on the effects of
haired terriers, and toy poodles have 400 to 600 odors exciting neurohumoral responses affecting
bundles per square centimeter. German shep estrus, pseudopregnancy, and pregnancy in the
herds, Airedales, and rotweilers had only 100 to mouse. The secretions of the skin glands have
300 bundles per square centimeter. The other long been suspected of being related to the
breeds were somewhere in between these fig identification and attraction of male dogs to fe
ures. The number of hairs per bundle varied from males in estrus.
nine to fifteen in the rotweiler to two to five in
the dachshund. In general, the hair of those Hair Types in the Dog
breeds which have many bundles is finer than
in those breeds which have a smaller number of There is a great deal of variability in hair
hair bundles. Midget animals have a greater length, color, diameter and transverse contour
number of bundles with fewer and finer hairs as among the various breeds of dogs and between
compared to the larger animals of the same individuals of the same breed. Brunsch (1956)
breed. has classified canine hairs into six types:
The canine hair follicle could be defined as a 1. Straight hair (Linearhaar). This is a bristly
pilosebaceous-arrector muscle complex. The firm hair, often deeply pigmented. It is some
sebaceous glands of the individual hair follicles times called a protective hair, primary hair, or
bunch together in clusters and sometimes fuse. over-hair. This is the strongest hair and is the
The arrector pili muscles originate from the chief hair in the follicle bundle. It is usually the
outer root sheath of each hair follicle and then longest hair, and the shaft is either straight or
join together into a common muscle bundle bowed. It has a thick medulla and a thin cortex.
which is inserted in the dense dermal tissue be 2. Bristle hair (Grannenhaar). This is a bristle
neath the epidermis. When the arrector pili mus with a spinelike tip, but weaker and softer near
cle contracts, the entire complex of follicles is the base. The distal third is similar to type 1, but
elevated and the sebaceous gland material the proximal two-thirds may be slightly wavy.
empties into the upper common follicle struc In the hair coat it is difficult to distinguish from
ture, which is shared by all the hairs. A single ap type 1. The medulla is slightly smaller than that
ocrine gland is associated with each follicle of type 1. The bristle hair is shorter than the
complex. The coiled secretory tubule extends straight hair, but is regarded as an over-hair or
deep into the subcutaneous tissue. A direct ex protective hair. This type may be the chief hair
tension of this becomes the excretory duct for the in a bundle, but is usually a subsidiary hair to
apocrine secretion, which extends up along the type 1 .
follicle complex and empties into the common 3. Wavy bristle hair (Wellgrannenhaar).
part of the follicle above the opening of the This type is finer and shorter than type 2. It is
sebaceous glands. The apocrine glands are sweat wavy with a well-developed bristle. These are
glands, but do not play a large role in the heat- the larger subsidiary hairs, but are usually in
regulating mechanism of the dog. They are com cluded with the cover-hairs or protective hairs.
parable to the apocrine sweat glands associated The medulla and cortex are smaller than in type
with hair follicles of the axillary and pubic re 2 , but the cortex is relatively heavier.
gions of man. The oily secretion from the glands 4. Bristled wavy hair (Grannenwellhaar).
associated with the hair follicles tends to keep This is a long, soft hair which is shorter and finer
the skin soft and pliable and spreads out over the than type 3, with a poorly developed bristle and
hair shafts. This gives the coat a glossy sheen. a smaller medulla. It is wavy in the lower two-
During periods of sickness, malnutrition, or thirds of the shaft. This type represents the larg
parasitism, the hair coat frequently becomes est hairs of the undercoat.
dull and dry as a result of inadequate functioning 5. Large wavy hair (grobes Wellhaar). This
of the skin glands. type is shorter and finer than type 4, and the
It has been postulated that secretions of the shaft is very wavy with a small bristle on the tip.
skin glands may play an important role in the The medulla is very small and may be discon
reproductive cycle of mammals. Parks and Bruce tinuous. The cortex is relatively thick. This type
882 C h ap ter 17. T he S en se O rgans and In tegum en t
gives a furlike or wool-like feel to the undercoat. granules may remain between the cells, as is the
6. Fine wavy hair (feines Wellhaar). This case in the medulla, but most of them are en
type is shorter and finer than type 5 and is some gulfed by the cells. The amount of pigment and
times described as vellus hair, fuzz, down, or variations in location produce different optical
lanugo hair. The medulla is discontinuous or ab effects. The pigmentation may be uniform
sent. This type represents the finest and smallest through the entire length of the hair, or it may
hairs of the undercoat and is usually wavy with vary. In the agouti type of hair, which is found
a small and poorly developed bristle on the tip. in wild rabbits, wolves, and in some domesti
cated breeds of dog (German shepherd and Nor
Variability in Hair Coat of the Dog wegian elkhound), the tip of the hair is white,
and the thick part of the bristle is heavily pig
The formation of bristles at the tips of the hair mented (black or dark brown), the proximal two-
shafts of greater diameter than the remaining thirds of the hair having lighter pigmentation
parts of the hairs suggests that the early part of (yellow or red).
the hair growth cycle is the most productive. Despite the wide range of color hues that are
Bristle formation is greater in hair types 1, 2, and possible in the coat, microscopic examination
3 than in types 4, 5, and 6 . Brunsch (1956) ex has revealed only black, brown and yellow pig
plains the difference in growth intensity of dif ment granules. The black-brown pigment is des
ferent follicles on a physiological basis. Those ignated as “tyrosine-melanin,” since it is formed
follicles with a rich blood supply and source of by enzyme oxidation of tyrosine to melanin. The
metabolites will synthesize more hair shaft. Thus yellow-red pigment is designated as “pheo-
smaller follicles with less blood supply produce melanin.” Its origin is unknown. Da Fonsicaand
smaller hairs. Cabral (1945) have classified the dog’s coat ac
Gair (1928) classified the coat of the dog into cording to color and pattern. The studies of
three groups on the basis of hair length. The inheritance and genetic control of color and coat
normal coat, which resembles the hair covering patterns have been summarized by Winge
of wild canidae (wolf, jackal), is typified by the (1950) and Little (1957).
German shepherd. The short-hair type is repre
sented by the boxer. The long-hair type may be Implantation of Hairs in the Skin of the Dog
illustrated by the chow. There are many
variations among the long-haired types, such as Some of the differences that occur in the ap
wire hair, tight curly hair, and flat, long hair. pearance of the coat of various types of dogs are
The various coats observed in domestic breeds due to the variation in the implantation angle of
of dogs are made up of the six types of hair de the hair follicles. The chow, Airedale, and also
scribed by Brunsch (1956), with some excep the scotty have an implantation angle of 45 de
tions. The wire-haired breed, such as the grees. Other breeds, such as the long-haired
schnauzer, has a preponderance of bristle-type dachshund, cocker spaniel, and Irish setter,
hairs, with a seventh type not found in other have an implantation angle of less than 30 de
breeds. The cocker spaniel and the setter have grees. The majority of all breeds examined by
fine, long silky hairs with less obvious bristle de Brunsch (1956) had an angle between 30 and
velopment. The poodle has extremely long hair 40 degrees. There is a tendency for long-haired
which seems to grow continuously. Hair type 1, dogs to have a higher implantation angle. Gen
or straight hair, is absent, and all hairs resemble erally, the hairs slant in a caudal direction from
the wool hair type. The medullary canal of a the nose toward the tip of the tail.
poodle hair is greatly reduced or absent. The Niedoba (1917) has described the streams and
bristle formation of a poodle hair is character convergent and divergent whirls and the lines
ized by a rhythmic pattern of differences in the where streams of different direction join. The
thickness of the hair, thus suggesting continuous patterns are subject to great variation. Some of
growth with variations in growth intensity. the more obvious features that can be easily ob
served on short-haired dogs are the center of
Coat Color in the Dog nasal divergence, cheek whirls, ear center, ven
tral cervical stream, neck diverging line, diverg
The color of the hair shaft is produced by pig ing breast whirls, ventral center line (division of
ment cells in the bulb of the hair follicle. These hair cover on both sides of the body), thoracic
cells place granules of pigment in the cortical whirls from the ventral cervical stream, whirl in
and medullary cells during development. The the region of the elbow, and rump whirls.
T he In tegum en t 883
Seasonal Differences in the Dog’s cum, stratum spinosum, and stratum corneum.
Hair Coat and the Hair Cycle In a few areas the stratum granulosum and
stratum lucidum are evident, but these are in
The process of shedding is gradual, and the frequent and are in areas where keratinization
coat of one season gradually merges into that of is retarded (i.e. around hair follicle orifices). The
the next, so that the dog is never without a pro number of layers of cells varies between three
tective covering. Hairs mature and are shed and six. In regions where the stratum granu
according to the metabolic state of the skin. The losum and stratum lucidum are evident, there
rate of growth varies in different follicles and in are as many as eight layers of cells. Cells of the
different regions of the body. A hair which has stratum cylindricum usually have their nuclei
been shed naturally is differentiated from one oriented with the long axis perpendicular to the
which has been broken or shorn by the slightly surface of the skin. They have a smaller amount
bulbous proximal end, which is frayed out into of cytoplasm and stain more intensely than more
fibrillae. When club hairs are plucked during superficial cells. Fewer mitotic figures are pres
the resting phase (telogen), new hairs begin to ent in the stratum cylindricum of hairy skin than
grow at once, whereas new hair growth occurrs in the same layer of foot pads and planum nasale.
much later if the resting hair is allowed to shed The epidermal papillae are covered by a thick
naturally. This may influence the development ened epidermis which is usually six to twelve cell
of the coats of wire-haired fox terriers, which layers thick, about twice as thick as the surround
are customarily plucked when groomed for show ing epidermis (Fig. 17-37, B). The dermis under
purposes. When a growing hair is plucked dur the epidermal papillae is composed of very fine,
ing anagen, nearly all the lower half of the folli closely packed connective tissue fibers which ex
cle is pulled out with it. Clipping and shaving tend under the thickened epidermis to form the
have no effect on hair growth or the stage of the papilla. The dermal-epidermal junction of the
cycle. hairy skin is normally thrown up into folds which
The evidence presented by Comben (1951) parallel the surface. This is in contrast to the dis
indicates that there is little growth of hair in the tinct epidermal pegging present in the epidermis
dog in warm weather, whereas a period of active of the planum nasale, foot pads, and mucous
growth occurs when cool weather starts. The membranes.
dog sheds more cover-hair in the spring than in
the summer. The number of hairs in each bundle MUSCLES OF THE SKIN
increases in the winter. There is a great deal of
variation in the manner in which dogs shed their The erector pili muscles are best developed
hair, even among individuals of the same breed on the dorsal line of the neck, back, and tail.
kept under similar environmental conditions and Erector pili muscles (smooth muscle) are absent
diet. Endocrine secretions seem to have some from tactile hairs (which have striated fibers) and
influence on hair production. De Vita (1939) re from vibrissae.
ported alopecia over the dorsal sacral region in In various portions of the body striated muscle
bitches suffering from hyperplastic endometritis fibers and bundles radiate from the cutaneous
caused by excessive estrogen secretion. Gardner muscle into the corium and attach to the follicles
and de Vita (1940) showed that hair growth in of tactile hairs.
dogs was inhibited by the administration of es
trogens. GLANDS OF THE CANINE SKIN
Surface Contour of Hairy Skin Merocrine sweat glands are found only in the
and Histology of Epidermis foot pads (Nielsen 1953). They are placed deeply
in the fat and fibrous tissue under the thick foot
The skin surface is irregular because of scale pads. They are small, tightly coiled, tubular
like folds which form depressions into which the glands, with minute lumina which are lined with
hair follicles invaginate (Fig. 17-37). The pat cuboidal cells. They contain coarse granules
tern of skin folds is occasionally interrupted by scattered in the clear cytoplasm. Myoepithelial
the presence of knoblike enlargements, 0.33 to cells may be demonstrated peripheral to the
0.35 mm. in diameter, the epidermal papillae secretory tubules. The excretory ducts follow a
(Fig. 17-37, B). tortuous path through the dermis and epidermis
Histologically, the epidermis of the hairy skin, and empty in the crevices between the conical
varying in thickness from 30 to 40 microns, usu papillae of the foot pads. The merocrine secre
ally consists of three layers: stratum cylindri- tion is watery.
884 C h ap ter 17. T he Sen se O rgans and In tegum en t
C om p ound f o llic le
S c a le - lik e fo ld
| -----------T h re e g ro u p e d f o llic le s
E p id e rm a l p a p illa
E p id e rm is
H a ir fo llic le
A r r e c t o r pili
m u s c le
S e b a c e o u s glan d
D e rm is
A p ocrin e
s w e a t gland
F ig . 17-37. Surface contour, hair arrangement, and histology of the hairy skin. (After Lovell and Getty 1957.)
A. View of scalelike folds and arrangement of hair follicles
B. Histological section of hairy skin
T he In tegum en t 885
Apocrine sweat glands are found mainly in shafts are larger in diameter and differ in appear
connection with hair follicles (Speed 1941). The ance from the surrounding hair. The hairs of this
secretory parts of the glandular tubules are area emerge from the hair follicles, singly (Fig.
situated in the dermal and subcutaneous layers 17-38, B), whereas surrounding hair is of the
of the skin. The excretory duct passes up through complex follicle type, supporting six to eleven
the dermis and empties into the hair follicles hairs. The single hairs of the specialized area are
above the ducts of the sebaceous glands. The very stiff and coarse, and the surface of the skin
tubules and individual cells attain various sizes, has a yellow, waxy appearance probably due to
depending upon the secretory phase. In some an abundance of sebaceous secretion. The seba
sections, huge, dilated, cystlike tubules lined ceous glands and apocrine glands of the area are
with flattened elongated cells are found, while large, extending deep into the dermis and sub
in others the tubules are small with high cylindri cutaneous tissue (Lovell and Getty 1957). Hilde
cal epithelium. The highly developed mammary brand (1952) suggests that the tail gland area in
glands have a similar structure. wild canidae functions in recognition and identi
Sebaceous glands have the holocrine type of fication of the species.
secretion. Distributed over the integument in
connection with the hair follicles, they are larg BLOOD SUPPLY TO THE SKIN
est along the dorsal part of the neck, back, and
tail, particularly in the specialized tail gland The arteries to the skin include simple cutane
area. The Meibomian glands or tarsal glands of ous arteries, which reach the skin by running
the eyelids are also specialized sebaceous glands. between muscles while supplying small branches
The glands of the ear canal are apocrine and to the muscles, and mixed cutaneous arteries,
sebaceous. Cerumen is a product of both glandu which run through muscles, and supply large
lar types and appears as a fairly dry, dark brown muscular branches before terminating in the
ish substance. skin. Hughes and Dransfield (1959) have listed
The perianal or circumanal glands are most twenty-three mixed cutaneous arteries and six
numerous in the vicinity of the anal orifice (Parks teen simple cutaneous arteries. The vessels anas
1950). They consist of upper sebaceous portions tomose extensively with one another. The
with open ducts to hair follicles, and deeper, plexuses which are formed in the subcutaneous
nonsebaceous portions. The nonsebaceous lob fascia are especially evident over bony promi
ules are solid masses of large polygonal cells. It is nences and large muscle masses.
believed that the ducts to this portion of the Microscopically, the arterial supply to the skin
gland are solid and have no secretory activity. of the dog has been divided into three distinct
The anal sacs of the dog vary in size from a pea plexuses, all lying parallel to the surface. These
to a hazel nut. They are paired and lie on each are the deep or subcutaneous plexus, the middle
side of the anal canal between internal and ex or cutaneous plexus, and the superficial or sub-
ternal anal sphincter muscles. Each sac opens papillary plexus (Fig. 17-39).
onto the lateral margin of the anus by a single The subcutaneous plexus is made up of the
duct. The sacs are pockets of skin which form a terminal branches of the cutaneous arteries.
reservoir into which apocrine and sebaceous Branches from this plexus form the middle
glands open (Montagna and Parks 1948). They plexus, which is associated with the hair follicles
are lined by a thin, stratified squamous epitheli and glands.
um. Under the connective tissue supporting the The superficial plexus is formed by the union
epithelium there are many sebaceous and of small vessels arising from the middle plexus.
apocrine glands. The sebaceous glands tend to The papillary body contains numerous capillary
line the neck of the sac, while the apocrine glands loops which come from the superficial plexus.
are concentrated in the fundus. The combined In general, the veins and arteries parallel one
secretions of the tubules of the anal sac and the another. Arteriovenous anastomoses have been
sebaceous glands associated with its excretory observed in the deeper layers. Variations in the
duct forms a viscous, putrescent liquid. circulatory pattern have been noted in the vari
ous modified skin areas.
Tail Gland Area The lymphatics arise from capillary nets
which lie in the superficial part of the dermis or
An oval to diamond-shaped area, 1 to 2 inches surround the follicles and glands. The vessels
long, is located on the dorsal aspect of the tail, arising from these nets drain into a subcutaneous
not far from the sacrum (Fig. 17-38, A). The hair lymphatic plexus. Baum (1917) has described
886 C h apter 17. T he S en se O rgans and Integum en t
S im p le h a ir f o llic le
S c a le - l ik e fo ld
E p id e rm is
SSr— S im p le h a ir
f o llic le
D e rm is
— A rre c to r pili
m u s c le
L a rg e sebaceous
g la n d
L a rg e a p o c r in e
sw eat g la n d
) r1• •
F ig . 17-38. Surface contour, hair arrangement, and histology of tail gland area. (After Lovell and Getty 1957.)
Middle plexus
Deep plexus
Adipose tissue
F i g . 1 7 -3 9 . Schem atic section of th e skin of th e dog, showing epidermal papilla and blood vessels (veins in black).
T he In tegum en t 887
the lymph vessels and nodes associated with the maintains the pointed appearance of the claw.
skin of the dog. The coronary border of the claw fits into the
space under the ungual crest of the third pha
NERVE SUPPLY TO THE SKIN lanx. This relationship is hidden by the skin of
the claw fold. Dorsally, this fold is a modification
The small nerves of the skin lie within the sub of the hairy skin which is free from hair on one
cutis and are continued by a nerve plexus ex side and fused to the horn of the claw. As the
tending through the dermis to the epidermis. horny material is produced and grows out, it is
Branches arising from this plexus innervate the covered by a thin stratum tectorium which ad
epidermis, glands, muscles, and hair follicles. heres to the proximal part of the claw. A furrow
Some branches terminate in special nerve end along the volar or plantar surface of the claw
ings. separates it from the foot pad in a similar man
Every hair is equipped with a nerve ending. ner (Figs. 2-19, 2-20).
The sensory nerves penetrate the follicles im The periosteum of the third phalanx and the
mediately below the sebaceous gland ducts. dermis of the claw are continuous and fill the
There they divide and arrange themselves paral space between the bony and epidermal struc
lel to the longitudinal axis of the hair. In the folli tures. The vascularity of this tissue is well
cles of tactile hairs arborizations of nerve fibers demonstrated by the hemorrhage which follows
are opposed to the glossy membrane. The roots trimming the canine toenail into the dermal
of some are encircled by large circular sinuses corium. Microscopically, the corium of the cor
containing erectile tissue. When the pressure in onary and dorsal ridge areas has been described
the circular sinus is increased, the hair becomes as having a papillate structure (Trautmann and
a more efficient receptor. Fiebiger 1957).
Autonomic nerve fibers supply the glandular The stratum germinativum, which is the epi
tubules and the arrector pili muscles. dermal layer supported by the corium, is most
Lamellar corpuscles are easily demonstrated active in the coronary and dorsal ridge areas,
in the subcutaneous tissue of the foot pads. where most of the horny claw is formed. The in
ner surface of the claw wall bears small epider
SKIN GRAFTING mal lamellae. The epidermis of the claw is largely
composed of the horny stratum corneum, which
Autogenous skin grafting has been success consists of flat, cornified epidermal cells joined
ful in the dog (Jensen 1959). Four types have together into a horny mass. The epidermis of the
been used: delay transfer of a pedicle graft; free sole has a well-developed stratum granulosum
full-thickness graft; small deep graft; and split- and stratum lucidum.
thickness graft. The claw grows at a rapid rate, and if not worn
Histopathological studies of transplants indi off or trimmed will sometimes continue growing
cate that degenerative changes involve the epi in a circular fashion until the point of the claw
dermis and the upper layers of the dermis during approaches the region of the volar furrow be
the first eight to ten postoperative days, at which tween the base of the claw and the foot pad.
time regenerative processes equalize the degen
erative changes. The blood supply to the trans
plant is adequate by the twelfth day and
completely normal by twenty-four days. BIBLIOGRAPHY
Epling, G. P. 1953. The anatomy of the skin. In Canine Medi Montagna, W , and H. F. Parks. 1948. A histochemical study of
cine, Chapter 10. Evanston, 111., American Veterinary the glands of the anal sac of the dog. Anat. Rec, 100:297-
Publications, Inc. 318.
Gair, R. 1928. Die Wuchsformen des Haarkleides bei Haus- Niedoba, T. 1917. Untersuchungen iiber die Haarrichtung der
tieren nach Untersuchungen beim Hunde. Z. Tiersucht Haussaugetiere. Anat. Anz. 50: 178-192, 204-216.
u. Zuchtungsbiol 11(1)'. 57-88. Nielsen, S. W. 1953. Glands of canine skin; morphology and
Gardner, W. V , and J. de Vita. 1940. Inhibition of hair growth distribution. Am. J. vet. Res, 14: 448-454.
in dogs receiving oestrogens. Yale J. Biol. Med. 13: 213- Parks, A. S , and H. M. Bruce. 1961. Olfactory stimuli in mam
215. malian reproduction. Science 134: 1049-1054.
Ham, A. W. 1961. Histology. Philadelphia, J. B. Lippincott Parks, H. 1950. Morphological and cytochemical observations
Company. on the circumanal glands of the dog. Ph.D. thesis. Cornell
Hildebrand, M. 1952. The integument in canidae. J. Mammal. University, Ithaca, New York.
33: 419-428. Speed, J. G. 1941. Sweat glands of the dog. Vet. J , 9 7:252-256.
Hughes, H. V , and J. W. Dransfield. 1959. The blood supply Trautmann, A , and J. Fiebiger. 1957. Fundamentals of the
to the skin of the dog. Brit. vet. J. 115: 1-12. Histology of Domestic Animals. (Translated and revised
Jensen, E. C. 1959. Canine autogenous skin grafting. Amer. J. by R. E. Habel and E. L. Biberstein.) Ithaca, Comstock
vet. Res. 20: 898-908. Publ. Asso.
Little, C. C. 1957. The Inheritance of Coat Color in Dogs. Webb, A. J , and M. L. Calhoun. 1954. The microscopic anat
Ithaca, Comstock Publ. Asso. omy of the skin of mongrel dogs. Am. J. vet. Res, 15:
Lovell, J. E , and R. Getty. 1957. The hair follicle, epidermis, 274-280.
dermis, and skin glands of the dog. Amer. J. vet. Res. 18: Winge, O. 1950. Inheritance in Dogs. Ithaca, Comstock Publ.
873-885. Asso.
Index
Folio numbers in italics refer to illustrations.
889
890 In d ex
Arteries (Continued) Articular process ( Continued) Ascending branch of cranial fem oral artery.
palpebral, 301 o f coccygeal vertebrae, 58, 60 366
pancreaticoduodenal. See Pancreatico o f lum bar vertebrae, 56 Ascending cervical artery7, 320
duodenal artery. o f sacral vertebrae, 56 Ascending colon, 692
parotid, 300 of thoracic vertebrae, 54 Ascending m esocolon, 693
pedal, dorsal, 371 of vertebral arch, 51 Ascending pharyngeal artery, 293. 649
pericardiacophrenic, 321 Articular surface Ascending portion o f duodenum , 686
perineal, 379 for central tarsal, 89 Association fibers, 486
periosteal, 5 of arytenoid cartilage, 721 Association neurons, 466
peroneal, 367 of ribs, 61 A tlantal fossae, 52
pharyngeal, ascending, 293, 649 o f tibia, 85, 87 A tlantal ligam ent, transverse, 103
phrenic, 355 o f tibial tarsal bone, 89 A tlanto-axial articulation, 101-103
phrenicoabdominal, 355 Articularis genus muscle, 242 A tlanto-axial m em brane, dorsal, 101
plantar, 377 Articularis ulnaris radii proxim alis, 69 A tlanto-occipital articulation, 101
popliteal, 367-71 Articulatio angularis, 98 A tlanto-occipital ligam ent, lateral, 101
prostatic, 379 Articulatio antebrachiocarpea, 113 Atlanto-occipital m em brane, 101
pudendal. See Pudendal arteries. Articulatio atlanto-axialis, 101 A tlanto-occipital space, caudal aspect, 102
pulmonary, 287-8, 739 Articulatio atlanto-occipitalis, 101 Atlas, 51,52
radial. See Radial artery. Articulatio capitis costae, 106 caudal lateral aspect, 53
rectal. See Rectal artery. Articulatio com posita, 98 cranial lateral aspect, 53
renal, 351, 746 Articulatio condylaris, 98 ligaments, 102
sacral, median, 386 Articulatio costotransversaria, 106 A tria, 274-6
saphenous, 366 Articulatio coxae, 119 A trioventricular bundle, 283
Articulatio cricoarytenoidea, 721 A trioventricular fibrous rings, 276
satellite, o f ischiatic nerve, 386
Articulatio cubiti, 110 A trioventricular node, 283
scapular, circumflex, 325
septal, middle, 309 Articulatio ellipsoidea, 98 A trioventricular orifice, 274, 278
Articulatio fem oropatellaris, 122 A trioventricular ostium, right, 278
posterior, 304
Articulatio femorotibialis, 122 A trioventricular valves, 282
spermatic, internal, 354, 355
Articulatio genus, 122 A trium
sphenopalatine. See Sphenopalatine artery.
Articulatio hum eri, 108 left, 273, 276
splenic, 346
Articulatio hum eroradialis, 110 oblique vein of, 287
stylomastoid, 297
Articulatio hum eroulnaris, 110 o f middle m eatus, 716
subclavian, 316-24
Articulatio incudom allearis, 99 right, 273, 274
subcostal, 345
Articulatio incudostapedia, 99 right lateral aspect, 275
sublingual, 297
A trium dextrum, 273, 274
submental, 297 Articulatio intertarsea, 127
Articulatio intertarsea distalis, 127 A trium m eatus medii, 716
subscapular, 325
A trium sinistrum, 273, 276
superficial, o f right hindpaw , plantar as Articulatio intertarsea proxim alis, 127
Auditory artery, internal, 862
pect, 374 Articulatio m ediocarpea, 113
Auditory meatus, external, 851
suprarenal, 354 A rticulatio plana, 98
A uditory ossicles, 853-6
suprascapular, 320 Articulatio radioulnaris distalis, 113
joints, 99-100
supraspinous, 320 A rticulatio radioulnaris proxim alis, 110, 113
ligaments, 99, 100
systemic, 288-387 Articulatio sacroiliaca, 119
of right ear, 858
tarsal, 371 Articulatio sellaris, 98
A uditory tube, 719, 853
temporal. See Temporal artery. Articulatio simplex, 98
osseous, 19, 23, 42
testicular, 354 Articulatio spheroidea, 98
ostium, 853
thoracic. See Thoracic arteries. Articulatio talocruralis, 127
pharyngeal opening, 719
thoracodorsal, 325 Articulatio tem porom andibularis, 23, 99
A uerbach’s plexus, 633
thyroid, 289, 817 Articulatio tibiofibularis distalis, 127
Auricle (heart)
tibial, 371 Articulatio tibiofibularis proxim alis, 127
left, 276
tonsillar, 296 Articulatio trochoidea, 98 right, 274
transverse, o f neck, 317 Articulationes carpi, 113 Auricula (ear), 848
tympanic, 303 Articulationes carpom etacarpeae, 113 A uricula dextra, 274
ulnar. See Ulnar artery. Articulationes costochondrales, 108 A uricula sinistra, 276
umbilical, 378 Articulationes costovertebrales, 106 A uricular arteries, 859, 862
ureteral, caudal, 378 Articulationes intercarpeae, 113 great, 297, 859
urogenital, 378 Articulationes interm etacarpeae, 114 A uricular branch
uterine, 378, 379 Articulationes interphalangeae distales, 118 anterior, of superficial tem poral artery, 301
utero-ovarian, 355 Articulationes interphalangeae proximales, o f vagus nerve, 630
vaginal, 379 118 A uricular cartilage, 848, 850
vertebra], 316 Articulationes intratarseae, 127 A uricular nerves
vesical, 378 Articulationes m anus, 113-18 caudal, 559
zygomatic, 305 Articulationes m etacarpophalangeae, 114 great, 575
Arteriolae rectae spuriae, 746 Articulationes ossiculorum auditus, 99 internal, rostral, 556
Arthrology, 95-130 Articulationes pedis, 127-9 Auricular rami, 859
Articular angle o f scapula, 66, 67 Articulationes sternocostales, 108 of vagus nerve, 567
Articular branch o f caudal cutaneous sural Articulationes tarsi, 127 Auricular surface o f ilium, 81
nerve, 618 Articulationes tarsom etatarseae, 127 Auricular veins, 399, 862
Articular capsule Articulations, 95 A uricularis anterior superior muscle, 142
o f costovertebral joints, 106 atlanto-axial, 101-103 Auriculolabialis muscle, 139
of shoulder joint, 108 atlanto-occipital, 101 A uriculopalpebral nerve, 559
Articular cartilage, 96, 97 cricoarytenoid, 721 Auriculotem poral nerve, 556
Articular circumference o f radius, 69 Aryepiglottic fold, 724 Auris externa, 847
Articular facet o f rib, 61 Arytenoid cartilage, 721 Auris interna, 848, 856
Articular meniscus, 99 Arytenoideus transversus muscle, 159 Auris m edia, 847
Articular muscles, 133 action, 161 Autonomic end-form ation, 642
Articular process, 36 Ascending aorta, 288 A utonom ic ganglia, 626
894 I n dex
Canalis transversa, 12 Cardiovagal nerve ( Continued) C artilago parap atellare m ediale et laterale,
Canalis trigemini, 20 left, 637 123
Canalis vertebralis, 51 Carina, tracheal, 728 Cartilago parietalis dorsalis, 714
Canine teeth, 651 Carina tracheae, 728 Cartilago parietalis ventralis, 714
Canthi o f eyelids, 838 Carnassial teeth, 652 Cartilago scapulae, 66
Cap C arotid arteries Cartilago septi nasi, 714, 863
duodenal, 686 common, 289 Cartilago sesamoidea, 721
knee. See Patella. branches, 298 Cartilago thyroidea, 719
skull, 44 lateral aspect, 294 Cartilago vomeronasalis, 714, 866
Capillaries, lymphatic, 430 ventral aspect, 291 Cartilago xiphoidea, 64
C apitulum humeri, 69 external, 292 Caruncle
Capsula articularis, 96, 101 internal, 312 lacrimal, 840
o f shoulder joint, 108 Carotid body (glomus), 637 sublingual, 645
C apsula externa, 489 Carotid canal, 23 Caruncula lacrimalis, 840
Capsula extrema, 489 C arotid foram en Caruncula sublingualis, 645, 660
Capsula fibrosa o f kidneys, 741 external, 19, 23,42 C astration, effects, 831
Capsula fibrosa perivascularis, 703 internal, 23, 45 Catagen, 879
C apsula glomeruli, 746 posterior, 23 Cauda epididymidis, 757
C apsula interna, 488 C arotid sinus, 312, 637 C auda equina, 536, 572, 595
Capsula ossei labyrinthi, 20 ramus to, 567 C auda nuclei caudati, 488
Capsula prostatae, 762 Carpal bones, 73, 74 C auda pancreatis, 708
Capsular ligament, 96 left, palm ar aspect, 77 Caudal abdom inal artery, 359
C apsular synovial bursa, 251 Carpal canal, 114 Caudal angle o f scapula, 66
Capsularis coxae muscle, 242 Carpal fibrocartilage, palm ar, 114 Caudal antebrachial muscles, 217-22
Capsule Carpal joints, 113-14 Caudal articular processes
articular. See A rticular capsule. Carpal ligament, palm ar, transverse, 113 of coccygeal vertebrae, 58,60
Bow m an’s, 746 Carpal transverse ligament, palm ar, 231 o f sacral vertebrae, 56
external, 489 C arpom etacarpal joints, 113 Caudal articular surface of atlas, 52
extreme, 489 Carpus, 64, 73-4 Caudal auricular nerves, 559
fibrous, o f liver, 703 dorsal rete, 329 Caudal belly o f m. sartorius, 242
glomerular, 746 dorsal venous rete, 405 Caudal border
hypophyseal, 812 flexed, ligaments of, dorsal aspect, 116 of scapula, 66
internal, 488 left, dorsal aspect, 76 o f spleen, 458
joint. See Joint capsule, medial aspect, 76 Caudal brachial muscles, 209-10
o f left shoulder joint, 109 palm ar aspect, 77 Caudal cardiosym pathetic nerves, 636
o f prostate, 762 schematic section, 117 Caudal cardiovagal nerves, 632
otic, 19 superficial ligam ents, palm ar aspect, 115 Caudal circumflex hum eral artery, 325
T enon’s, 842 Cartilage Caudal circumflex vein of humerus, 403
C aput accessorium o f m. triceps, 210 accessory, o f nose, 714 Caudal clunial nerves, 611
C aput costae, 61 annular, 851 Caudal cornu o f thyroid cartilage, 719
C aput epididymidis, 757 articular, 96,97 Caudal cruciate ligament, 123
C aput femoris, 82 arytenoid, 721 Caudal cutaneous antebrachial nerve, 587
C aput fibulae, 88 auricular, 848, 850 Caudal cutaneous femoral nerve, 611
C aput hum erale, 221 costal, 61 Caudal cutaneous sural nerve, 615
C aput hum eri, 67 cricoid, 721 Caudal descending coronary artery, 285
C aput laterale o f m. triceps, 210 epiglottic, 719 Caudal epigastric artery, deep, 360
C aput longum o f m. triceps, 209 epiphyseal, 3 Caudal femoral artery, 367
C aput m ediale o f m. triceps, 210 interarytenoid, 724 Caudal femoral vein, 413
C aput nuclei caudati, 488 intersternebral, 64 Caudal flexure o f duodenum , 686
C aput pancreatis, 708 laryngeal. See Laryngeal cartilage. Caudal genicular arteries, 371
C ap u t rad iale o f m. flexor digitorum p ro nasal, 713, 715 Caudal gluteal artery, 386
fundus, 221 o f larynx, 719-24 Caudal gluteal nerve, 611
C aput radii, 69 o f nose, 713-16 Caudal gluteal vein, 414
C aput tali, 89 parietal, 714 Caudal iliohypogastric nerve, 599
C aput ulnare scapular, 66 Caudal incisure, 848
o f m. flexor carpi ulnaris, 220 scutiform, 851 Caudal internal auricular nerves, 559
ofm . flexor digitorum profundus, sesamoid, 721 Caudal labial branch o f perineal artery,
221 thyroid, 719 383
Cardia, 679 tracheal, 728 Caudal labial nerve, 614
Cardiac glands, 684 tym panohyoid, 38 Caudal laryngeal nerve, 570
Cardiac impression, 699 vom eronasal, 714,866 Caudal m argin o f lung, 735
o f lungs, 735 Cartilage bones, 4 Caudal maxillary alveolar nerve, 555
o f right lung, 738 C artilagejoints, hyaline, 96 Caudal m ediastinal cavity, 731
Cardiac lobe Cartilagines laryngis, 719 Caudal m ediastinal pleura, 733
o f left lung, 735 Cartilagines nasi, 713 Caudal m eningeal artery, 293
o f right lung, 738 Cartilagines parapatellares, 85 Caudal mesenteric artery, 351
Cardiac muscle, 131 Cartilagines tracheales, 728 Caudal mesenteric plexus, 640
Cardiac nerve, 636-7 Cartilaginous joint. 95-6 Caudal m esenteric vein, 407
Cardiac notch Cartilaginous nasal septum, 863 Caudal muscles
o f lung, 735, 738 Cartilago accessoria of nose. 714 o f crus, 252-62
o f stomach, 681 o f thigh, 236-40
Cartilago articularis, 96
Cardiac p ortion o f stomach, 681 Caudal nasal nerve, 555
Cartilago arytenoidea, 721
Cardiac skeleton, 276 Caudal pancreaticoduodenal artery, 350
Cartilago costalis, 61
Cardiac sphincter, 683 Caudal pancreaticoduodenal veins, 407,
Cardiac veins, 287 Cartilago cricoidea, 721 409
C ardiosym pathetic nerves, 636 Cartilago epiglottica, 719 Caudal pelvic aperture, 80
Cardiovagal nerve, 633 Cartilago epiphysialis, 3 Caudal portion o f duodenum , 686
caudal, 632 Cartilago interarytenoidea, 724 Caudal recess of om ental bursa, 678
cranial, 632 Cartilago intersternebralis, 64 Caudal rectal artery, 379
I n dex 897
Caudal rectal nerve, 611 Cavum vaginale, 754 Cervical lymph nodes. See Lym ph nodes,
Caudal rectal vein, 414 Cecal artery, accessory, 350 cervical.
Caudal scrotal branch o f perineal artery, 379 Cecal branches o f ileocecal artery, 350 Cervical nerves, 574-8
Caudal scrotal nerve, 614 Cecocolic orifice, 689 dorsal branches, dorsal aspect, 576
Caudal superficial epigastric artery, 363 Cecocolic sphincter, 689 schema, 573
Caudal surface Cecum, 689-92 Cervical p art o f spinal cord, 533
o f humerus, 69 muscle layers, dorsal aspect, 690 Cervical plexus, 575
of radius, 71 Celiac arteries Cervical region, upper, 629
of ulna, 72 mesenteric, ventral aspect, 356 Cervical rib, 51
Caudal thyroid artery, 289, 817 ventral aspect, 357 Cervical spinal cord, arteries, ventral aspect.
Caudal thyroid notch, 719, 721 Celiac trunk, 345 315
Caudal thyroid vein, 393, 819 Celiacomesenteric plexus, 638,640 Cervical vertebrae. See Vertebrae, cervical.
Caudal tibial artery, 371 Cells Cervical vertebral veins, right lateral aspect,
C audal tibial ligament, 123 chromaffin, 828 427
Caudal tibial vein, 413 glandular, o f stomach, 684 Cervicoauricularis medius muscle, 143
Caudal tibiofibular ligament, 127 glial, 469 C ervicoauricularis p rofundus anterior mus
Caudal ureteral artery, 378 olfactory, 867 cle, 143
Caudal ureteral vein, 416 Cem entum, 653 C ervicoauricularis profundus m ajor muscle,
Caudal uterine artery, 379 C entral branches o f m iddle cerebral artery, 143
Caudal vena cava, 405-407 313 Cervicoauricularis profundus m inor muscle,
Caudal vesical artery, 378 Central canal, 536 144
Caudal vesical vein, 416 Central gray m atter, 497 Cervicoauricularis superficialis muscle,
Caudate lobe o f liver, 703 C entral interm ediate substance, 536 143
Caudate nucleus, 488 Central tarsal bone, 89 C ervicoauriculo-occipitalis muscle, 142
Caudate process o f liver, 703 C entral veins o f liver, 704 Cervicointerscutularis muscle, 143
Cava medullaria, 4 Centrum o f vertebra, 5 1 Cervicoscutularis m edius muscle, 143
Cavernous portion o f male urethra, 111 Centrum tendineum perinei, 699 Cervicoscutularis muscle, 143
Cavernous sinus, 421 Cephalic vein, 401 Cervicothoracic ganglion, 635
Cavitas glenoidalis, 66 accessory, 403 Cervicouterine artery, 379
Cavity Cerebellar folia, 504 Cervix uteri, 786
abdominal. See Abdom inal cavity. Cerebellar fossa, 19, 45 Cervix vesicae, 748
cranial, 6,44-8 Cerebellar nuclei, 507 Cham bers of eye, 846
epidural, 541 Cerebellar peduncles, 503, 504, 507 Cheek teeth, 651
glenoid, 66 Cerebellar veins, 424 Cheeks, 646
infraglottic, 726 Cerebellomedullary cistern, 540 muscles, 134-9
joint, 96 Cerebellorubral tract, 507 Chemical regulation o f body, 807
mediastinal, 731,732 Cerebellothalamic tract, 507 Chem oreceptor afferents, 637
medullary, 4 Cerebellum, 480, 501, 504 Chiasm, optic, 496, 497
n a sa l See N asal cavity. arteries, 318 Chiasma opticum, 496
o f larynx, 724-6 dorsolateral view, 506 Chiasmatic cistern, 540
of nose, 6 Cerebral aqueduct, 495, 497, 542 C hief cells of stomach, 684
o f skull, 44-9 Cerebral arteries, 313 Choanae, 34,718, 863
oral, 645 dorsal aspect, 319 C hoanal border, 42
pelvic, 80,670 middle, lateral aspect, 318 C hoanal region, 42
pericardial, 267 Cerebral branch o f cerebrospinal artery, 317 C hondropharyngeus muscle, 154
peritoneal. See Peritoneal cavity. Cerebral cortex, 482 C horda inguinalis, 787
pleural, 732 Cerebral hemisphere Chorda obliqua, 110
pulp, 653 left, lateral view, 498, 499 C horda tympani, 853
subarachnoid, 540, 541 right, decorticated, 495 nerve, 559, 874
subdural, 541 medial surface, 487 C horda utero-ovarica, 780
thoracic. See Thoracic cavity. medial view, 498, 505 C hordae arteriae umbilicales, 749
tympanic, 20,23,851 Cerebral jugae, 16 Chordae tendineae, 278
vaginal, 754 Cerebral peduncles, 497, 500 Choriocapillaris, 844
Cavum abdominis, 667 Cerebral veins, 398, 422 Choroid, 843, 844
Cavum articulare, 96 Cerebrospinal artery, 317 Choroid arteries, 305, 313
Cavum conchae, 848 Cerebrospinal fluid, 542 Choroid plexus
Cavum cranii, 6,44 Cerebrum, 480, 482 o f fourth ventricle, 542
Cavum dentis, 653 arteries, 318 of lateral ventricle, 542
Cavum epidurale, 541 gyri, left dorsolateral view, 494 of third ventricle, 542
Cavum infraglotticum, 726 lobes, 482 o f ventricles, 540,541
Cavum laryngis, 724 sulci, left dorsolateral view, 494 C horoidal vein, 422
Cavum mediastinale caudale, 731 ventral view, 493 Chromaffin cell, 828
Cavum mediastinale craniale, 731 white m atter, 486 Chrom ophobes o f hypophysis, 813
Cavum m ediastinale medium, 731 Cerumen, 851, 885 Chyme, 681
Cavum m ediastinale ventrale, 732 Cervical artery Cilia, 838
Cavum nasi, 6, 48, 713, 716 ascending, 320 Ciliary arteries, 305
Cavum oris, 645 d e e p ,317 Ciliary body, 843, 844
Cavum oris proprium , 645 superficial, 320 Ciliary ganglion, 627
Cavum pelvis, 670 Cervical branch Ciliary nerve, long, 550
Cavum pericardii, 267 of occipital artery, 293 Ciliary processes, 844
Cavum peritonei, 667 of ventral buccal nerve, 563 Cingular gyrus, 485
Cavum pharyngis, 660 Cervical cardiac nerve, 636-7 Cingulate sulcus, 483
Cavum pleurae, 732 Cervical enlargem ent o f spinal cord seg Cingulum, 486
Cavum subarachnoideale, 540,541, 542 ments, 533 Cingulum m em bri pelvini, 78
Cavum subdurale, 541 Cervical fascia, 196 Cingulum m em bri thoracici, 64
Cavum thoracis, 728 Cervical ganglion, cranial, 635 Circle, arterial, o f brain, 313
Cavum tympani, 20, 23, 851 Cervical loop, 574 Circular m uscular c o at of stomach, 683
898 In d ex
Circulation, arterial, greater, 846 Cochlear p art o f vestibulocochlear nerve, 508 C ommon nasal m eatus, 718, 863
Circulus arteriosus cerebri, 310, 313 Colic arteries, 350, 351 Common palm ar digital nerves, 590
Circulus arteriosus m ajor, 846 Colic branch o f ileocecocolic artery, 350 Common peroneal (fibular) nerve, 618
Circulus articuli vasculosus, 96 Colic flexure, 692 Common plantar digital nerves, 622
Circum anal glands, 695, 885 Colic lym ph nodes, 448 Common vaginal tunic, 754, 761
Circumduction, 99, 133 Colic veins, 407, 409 Com m unicating branches of spinal nerves,
Circumferentia articularis, 69, 72 Collagen fibers, 471 471
Circumflex b ran ch o f left coronary artery, Collarette, 846 Communicating veins, 401
284 C ollateral branches o f intercostal arteries, C om partm ent
Circumflex vein o f scapula, 403 345 meniscomandibular, 99
Circumvallate papillae, 871 Collateral ganglia, 626 m eniscotemporal, 99
Cistern C ollateral ligaments. See Ligaments, collat Complexus muscle, 170
cerebellomedullary, 540 eral. Concha, nasal, 27, 863
chiasmatic, 540 Colliculus inferior, 500 Concha nasalis medialis, 863
interpeduncular, 540 Colliculus seminalis, 777 Concha nasalis ventralis, 863
lumbar, o f spinal subarachnoid cavity, Colliculus superior, 500 Conchae nasales, 863
542 Collum costae, 61 Conchohelicinus muscle, 142
o f lateral fossa, 540 Collum dentis, 649 Conduction, structures mediating, 471-9
subarachnoid, 540 Collum femoris, 82 Conduction system o f heart, 283
teat, 801 Collum humeri, 67 C ondylar joint, 98
C isterna cerebellomedullaris, 540 Collum radii, 71 Condyle
C isterna chiasmatis, 540 Collum tali, 89 humeral, 69
C isterna chyli, 431 Collum vesicae felleae, 705 occipital, 12, 42
C istem a fossae lateralis cerebri, 540 Colon, 692-3 o f femur, 84
C isterna interpeduncularis, 540 mesentery, 692-3 o f tibia, 85
C isterna magna, 540 muscle layers, dorsal aspect, 690 Condyli occipitales, 12
Cisternae subarachnoideales, 540
Colon ascendens, 692 Condyloid artery, 293
Clarke’s column, 536
Colon descendens, 692 Condyloid canal, 12,45
Claustrum, 489
Colon transversum, 692 Condyloid crest, 36
Clavicle, 64
Colonic nerve, lum bar, 641 Condyloid fossa
left, cranial aspect, 65
Color dorsal, 12
Clavicula, 64
o f coat, 882 ventral, 12,42
Clavicular tendon, 200
Claw, 887 vision, 846 Condyloid process, 36
Claw fold, 887 Colum na dorsalis, 536 Condyloid vein, 421
Cleft, hypophyseal, 814 C olum na lateralis, 536 Condylus humeri, 69
C leidobrachialis muscle, 201 C olum na ventralis, 536 Condylus lateralis et medialis, 85
Cleidocephalicus muscle, 201 Colum na vertebralis, 49 Condylus lateralis of femur, 84
Cleidocervicalis muscle, 201 Columnae anales, 694 Condylus medialis o f femur, 84
Cleidomastoideus muscle, 201 Columnae fornicis, 492 Cone, theca, 783
Clinoid process C olum nar zone o f anal canal, 694 Confluens sinuum, 419
anterior, 18 C olumns Conical papillae, 868, 871, 873
o f sella turcica, 810 anal, 694 Conjugal ligament, 106
posterior, 18, 45 C larke’s, 536 Conjugata of pelvis, 80
Clitoris, 791 dorsal, 536 Conjugate, pelvic, 80
artery, 383 lateral, 536 Conjunctiva, 837, 838
dorsal nerve, 614 o f fornix, 492 Conjunctival fornix, 838
vein, 414 ventral, 536 Conjunctival sac, 838
Clunial nerves, 610, 611 vertebral. See Vertebral column. Connecting peritoneum , 673
Coat(s) Com m issura alba, 539 Connective tissue o f skeletal muscles, 133-4
fibrous. See Fibrous coat. Commissura anterior, 488 Constrictor muscles o f female genitalia
hair. See H air coat. Commissura fornicis, 488 caudal aspect, 795
o f esophagus, 664-7 Commissura habenularum , 494 lateral aspect, 793
o f large intestine, 697-8 Commissura posterior, 493 Constrictor vestibuli muscle, 794
o f small intestine, 688-9 Commissurae labiorum , 646 Constrictor vulvae muscle, 794
o f stomach, 683-4 C ommissural fibers, 487 Contact surface o f tooth, 651
serous, o f liver, 703 Commissure, 539 Contrahentes digitorum muscles, 230
Coccygeal arteries, 386, 387 anterior, 488 Conus, ligament of, 276
Coccygeal fascia, superficial, 263 habenular, 494 Conus arteriosus, 278
Coccygeal nerves, 622-3 hippocam pal, 488 Conus medullaris, 533
Coccygeal part o f spinal cord, 533 o f lip, 646 Cor, 267
Coccygeal vein, lateral, superficial, 414 posterior, 493 Coracobrachialis muscle, 209
Coccygeal vertebrae, 49, 58-60 Com m on bile duct, 705, 706 Coracoid process, 67
Coccygeal vertebral veins, right lateral as C om m on carotid arteries, 289 Cord
pect, 428 C om m on colic artery, 350 spermatic, 758-62
Coccygeoanal muscle, 697 Common colic vein, 407 spinal. See Spinal cord.
Coccygeoanalis muscle, 194, 195 C om m on digital arteries Cornea, 843-4
Coccygeus muscle, 191 dorsal, 337 Corniculate process o f arytenoid cartilage,
Cochlea, 856, 857 palmar, 339 721
right, m edial view, 854 plantar, 377 Corniculate tubercle, 724
C ochlea tibiae, 87 C om m on digital nerves, dorsal, 619 C ornu cranialis et caudalis o f thyroid carti
Cochlear aqueduct, 22, 857 Common dorsal m esentery, 673 lage, 719
Cochlear canaliculus, 857 C om m on hepatic artery, 345 C ornu thyreoideum, 38
external opening, 20 C om m on iliac vein, 414, 416 Cornua
C ochlear duct, m em branous, 859 Common integum ent, 837 of antitragus, 848
Cochlear nerve, 563 C ommon interosseous artery, 329 of thyroid cartilage, 719
C ochlear nucleus, 508 Common interosseous vein, 403 Cornua uteri, 786
In d ex 899
Corona dentis, 649 C orticopontine fibers, 503 C ranial m esenteric artery, 349
Corona radiata o f ovary, 780, 832 C orticopontine tract, 500 branches, ventral aspect, 361
Coronal gyrus, 484 Corticospinal fibers, 503 C ranial m esenteric plexus, 640
Coronal sulcus, 483 Corticospinal tract, 500, 523 C ranial m esenteric vein, 407
Coronal suture, 14, 16 lateral, 508 C ranial muscles o f thigh, 240-42
Coronary arteries, 283-5 ventral, 508 C ranial nerves. See Nerves, cranial.
left, branches of, ventral aspect, 286 Costae, 61 C ranial pancreaticoduodenal artery, 346
Coronary groove, 273 C ostae fluctuantes, 61 C ranial pancreaticoduodenal vein, 409
Coronary ligament o f liver, 702 C ostae spuriae, 61 Cranial pelvic aperture, 80
Coronary plexuses, 637 C ostae verae, 61 Cranial pia mater, 540-41
C oronary sinus, 274, 287 Costal arch, 61 C ranial recess o f om ental bursa, 678
Coronoid crest, 36 Costal cartilage, 61 C ranial rectal artery, 351
Coronoid fossa, 69 Costal groove, 61 C ranial rectal vein, 407
Coronoid process, 36, 72 Costal pleura, 732 C ranial roots o f accessory nerve, 512
C orpora m ammillaria, 497 Costal surface C ranial scrotal branch o f external pudendal
Corpora quadrigemina, 497 o f lung, 734 artery, 363
Corpus adiposum auriculi, 851 o f scapula, 66 C ranial thyroid artery, 289, 817
Corpus adiposum infrapatellare, 122 Costocervical trunk, 317 Cranial thyroid notch, 719
Corpus adiposum orbitae, 837 Costocervical vein, 390 C ranial th y ro id vein, 390, 819
Corpus albicans, 783 Costocervical-vertebral venous trunk, 389 C ranial tibial artery, 371
Corpus callosum, 488 C ostochondral joints, 108 branches, cranial aspect, 373
vein of, 422 C ostodiaphragm atic recesses, 733 Cranial tibial ligament, 122, 123
Corpus cavernosum penis, 763, 764 Costom ediastinal recess, 732, 733 C ranial tibial vein, 413
Corpus cavernosum urethrae, 763, 764, 771 Costotransverse foram en, 61 C ranial tibiofibular ligament, 127
Corpus clitoridis, 791 C ostovertebraljoints, 106, 108 Cranial ureteric veins, 406
Corpus costae, 61 Costoxiphoid ligaments, 108 C ranial vault, 44
Corpus epididymidis, 757 Cover-hair, 879 Cranial vena cava, 389-401
Corpus femoris, 84 Cranial abdom inal artery, 359 C ranial venous sinuses
Corpus fibulae, 88 C ranial angle o f scapula, 66 dorsal aspect, 423
Corpus fornicis, 492 C ranial arachnoid, 540 lateral aspect, 420
Corpus geniculatum laterale, 496 C ranial arteries, m esenteric, ventral aspect, C ranial vertebral notch, 51
Corpus geniculatum mediale, 496 356 C ranial vesical arteries, 378
Corpus hemorrhagicum, 833 C ranial articular processes C raniolateral muscles of crus, 249-52
Corpus humeri, 67 o f coccygeal vertebrae, 58 C raniopharyngeal canal, 18
Corpus linguae, 654 o f sacral vertebrae, 56 Cranium. See also Skull.
Corpus luteum, 783, 833 Cranial articular surface o f atlas, 52 arteries, 314
Corpus m andibulae, 36 Cranial belly o fm . sartorius, 242 base, arteries of, dorsal aspect, 306
Corpus medullare, 504 Cranial border external surface, 14
Corpus nuclei caudati, 488 o f ilium, 80 nerves, 314
Corpus ossis hyoidei, 38 o f pubis, 80 Cremasteric artery, 360
Corpus ossis ilii, 80 o f scapula, 66 Crest
Corpus ossis incisivi, 27 o f spleen, 458 condyloid, 36
Corpus ossis ischii, 81 C ranial brachial muscles, 206-209 coronoid, 36
Corpus ossis pubis, 82 Cranial cardiosym pathetic nerves, 636 epicondyloid, lateral, 69
Corpus pancreatis, 708 C ranial cardiovagal nerves, 632 ethm oid, 34
Corpus penis, 763 C ranial cavity, 6,4 4 -8 ethm oidal, 30, 33
Corpus pineale, 494 C ranial cervical ganglion, 635 frontal, 16, 38
Corpus radii, 71 Cranial circumflex hum eral artery, 328 iliac, 80
Corpus restiforme, 504 Cranial cornu o f thyroid cartilage, 719 iliopectineal, 80
Corpus striatum, 488 Cranial cruciate ligament, 123 interosseous, 71, 72
Corpus tali, 89 Cranial cutaneous an teb rachial branches of intertrochanteric, 84
Corpus tibiae, 87 radial nerve, 586 m axilloturbinate, 30, 48
Corpus ulnae, 72 Cranial descending coronary artery, 285 median, o f cricoid cartilage, 721
Corpus uteri, 786 Cranial duodenal flexure, 686 nasoturbinate, 28
Corpus ventriculi, 681 Cranial dura m ater, 539-40 occipital, 12,44, 48
Corpus vertebrae, 51 venous sinuses, 419-22 o f lesser tubercle, 67
Corpus vesicae, 748 C ranial epigastric artery, 324 of ribs, 61
Corpus vesicae felleae, 705 Cranial femoral artery, 366 orbital. See Orbital crest.
Corpuscles Cranial femoral vein, 413 orbitosphenoidal, 16, 45
malpighian, 746 Cranial fossa palatine, 32
renal, 743, 746 anterior, 44, 45 petrosal, 19, 45
thymic, 462 caudal, 45,48 sacral, 56
Corpusculum renis, 743 middle, 45 sagittal. See Sagittal crest.
Corrugator supercilii muscle, 842 C ranial gluteal artery, 383 supraventricular, 278
Cortex C ranial gluteal nerve, 611 temporal, 23
cerebral, 482 C ranial gluteal vein, 416 tibial, 87
of kidneys, 743 C ranial hem orrhoidal artery, 351 transverse, 19
of lymph node, 434 C ranial iliohypogastric nerve, 599 ungual, 75
of ovary, 780 C ranial index, 8 urethral, 111
Cortex renis, 743 C ranial lab ial b ran ch o f external pudendal Cretinism, 821, 822
Corti, spiral organ of, 859 artery, 360 Cribriform foram ina, 25, 44, 45
Cortical branches o f m iddle cerebral artery, C ranial laryngeal nerve, 570, 630 Cribriform plate, 25, 27, 44
313 Cranial laryngeal vein, 398 Cribriform tract, spiral, 20
Corticocortical fibers, 486 C ranial m ediastinal cavity, 731 Cricoarytenoid articulation, 721
C orticonuclear fibers, 503 Cranial m ediastinal pleura, 732 Cricoarytenoideus dorsalis muscle, 159
Corticonuclear tract, 500 Cranial meninges, 539—41 Cricoarytenoideus lateralis muscle, 159
900 In d ex
Cricoesophageal muscle, 665 Cuneiform tubercle, 724 Deep dorsal m etatarsal veins, 417
Cricoesophageal tendon, 665 C upula, pleural, 733 Deep external fascia o f trunk, 196
Cricoid cartilage, 721 Cupula pleurae, 733 Deep facial vein, 394
Cricopharyngeus muscle, 156 C urvatura ventriculi m ajor, 679 Deep fascia
Cricothyroid branch o f cranial thyroid a r Curvatura ventriculi minor, 681 of head, 161
tery, 292 C urvatures o f stomach, 679-81 of neck, 175
Cricothyroid ligament, 721 Cuspis dorsalis, 282 o f pelvic limb, 263
C ricothyroideus muscle, 159 Cuspis ventralis, 282 of tail, 197
C rista capitis costae, 61 Cusps of thoracic limb, 230
C rista condyloidea, 36 o f atrioventricular valve, 282 Deep femoral artery, 360
C rista coronoidea, 36 semilunar, 282, 283 Deep femoral vein, 414
C rista epicondylica lateralis, 69 C utaneous an teb rachial branches, cranial, Deep inguinal lymph node, 446
C rista ethm oidalis, 33, 34 o f radial nerve, 586 D eep nasofrontal fascia, 161
C rista ethm oidalis dorsalis, 30 C utaneous antebrachial nerve D eep palm ar arch, 337, 339
C rista ethm oidalis ventralis, 30 caudal, 587 D eep palm ar m etacarpal arteries, 339
C rista frontalis externa, 16 lateral, 583 Deep palm ar m etacarpal nerves, 590
C rista frontalis interna, 16 medial, 583 Deep palm ar m etacarpal veins, 405
C rista galli, 25, 44 C utaneous brachial nerve, lateral, 582 Deep palm ar venous arch, 403
C rista iliaca, 80 Cutaneous branches D eep perineal fascia, 698
C rista iliopectinea, 80 dorsal, o f lum bar nerves, 599 Deep peroneal (fibular) nerve, 618
C rista interossea, 71 lateral, distal, o f intercostal nerve, 595 Deep plantar m etatarsal arteries, 377
C rista intertrochanterica, 84 o f third intercostal nerve, 594 Deep plantar m etatarsal nerves, 622
Crista m axilloturbinalis, 30 o f iliohypogastric nerve, 606 Deep plantar m etatarsal veins, 417
C rista m ediana o f cricoid cartilage, 721 o f thoracic nerves, 594 D eep temporal fascia, 161
C rista nasoturbinalis, 28 o f circumflex scapular artery, 325, 328 Deep temporal nerve, 556
C rista occipitalis externa, 12 o f coccygeal nerves, 623 Deep temporal vein, 398
C rista occipitalis interna, 12 of intercostal arteries, 342 Deep veins
C rista orbitalis, 34 o f saphenous nerve, 607 of pelvic limb, 413-16
C rista orbitalis ventralis, 39 o f subscapular vein, 403 of thoracic limb, 401-403
C rista orbitosphenoidalis, 18 ventral, o f intercostal nerve, 595 Deltoid tuberosity, 67
C rista palatina, 32 o f internal thoracic artery, 321 D eltoideus muscle, 204
C rista petrosa, 19 Cutaneous femoral nerve Demifacet, 54
C rista sacralis interm edia, 56 caudal, 611 D endrites, 464, 466
C rista sacralis lateralis, 56 lateral, 606 Dendritic zone o f neuron, 466
C rista sacralis m ediana, 56 Cutaneous helicine pouch, 848 Dens
C rista sagittalis externa, 12 C utaneous nerves o f trunk, lateral aspect, apical ligament, 101
C rista sagittalis interna, 12 601 of axis, 52
C rista supraventricularis, 278 C utaneous plexus, 885 D ental alveoli, 42
C rista temporalis, 23 C utaneous sural nerve, 615 D ental arches, 651
C rista terminalis, 274 C utaneous zone o f anal canal, 694 D ental formulae, 652
C rista tibiae, 87 Cutaneus trunci muscle, 182, 189 D entate gyrus, 485
C rista transversa, 19 Cystic artery, 346 D entate nucleus, 507
C rista tuberculi minoris, 67 Cystic duct, 705, 706 Dentes, 649
C rista unguicalaris, 75 Dentes canini, 651
C rista urethralis, 777 Dentes carnassei, 652
Cristae sagittales, 74 Dentes decidui, 649
Crown o f tooth, 649 Dentes incisivi, 651
Cruciate ligaments. See Ligaments, cruciate. D a r t o s , 754 Dentes molares, 652
Cruciate sulcus, 483 D eciduous dentition, 652 Dentes permanentes, 649
Crura Deciduous teeth, 649 Dentes premolares, 651
o f diaphragm , 180 Decussatio lemniscorum, 523 Denticulate ligament, 542
o f helix, 848 Decussatio nervorum trochlearium , 550 Dentin, 653
C rura clitoridis, 791 D ecussatio pedunculorum cerebellarium su- Dentinum , 653
C rural fascia, 196, 265 periorum , 500 D entition, 652
Crus, 78 Decussatio pyram idum , 508 D epressor auriculae muscle, 142
anterior, o f internal capsule, 488 Decussation Depressors of tail, 190
interosseous m em brane, 127 of medial lemniscus, 523 Descemet’s m embrane, 843
left, cross section, 256 o f pyramids, 508 D escending aorta, 288
muscles of, 257, 258, 259 o f superior cerebellar peduncles, 500 D escending branch
muscles, 249-62 tegmental, 501 o f cranial femoral artery, 366
posterior, o f internal capsule, 488 D eep artery o f penis, 383 of occipital artery, 293
Crus anterius capsulae internae, 488 Deep auricular artery, 300, 859 o f omocervical artery, 320
Crus cerebri, 500 D eep auricular vein, 399 Descending colon, 692
Crus helicis distale, 142 Deep brachial artery, 328 Descending genicular artery, 367
Crus interm edium o f diaphragm , 180 Deep brachial vein, 403 Descending mesocolon, 674, 693
Crus laterale D eep branch Descending portion o f duodenum, 686
of diaphragm atic crus, 180 o f deep circumflex iliac artery, 359 Dewclaw, 88, 92, 263
o f inguinal canal, 188 o f radial nerve, 583 Diabetes mellitus, 834
Crus m ediale D eep caudal epigastric artery, 360 D iagonal band, 490
of diaphragm , 180 Deep cervical lymph nodes, 439 Diameter
o f inguinal canal, 188 D eep circumflex iliac artery, 359 oblique, o f pelvis, 80
Crus posterius capsulae internae, 488 D eep circumflex iliac vein, 406,409 transverse, o f pelvis, 80
Cryptorchidism , 757 Deep cranial epigastric artery, 324 D iam eter obliqua, 80
C ubital vein, m edian, 401 D eep dorsal m etacarpal arteries, 337 D iam eter transversa, 80
C umulus oophorus, 780, 832 Deep dorsal m etacarpal veins, 403 Diaphragm , 178-81
C uneiform process o f ary ten o id cartilage, D eep dorsal m etatarsal arteries, 377 abdom inal surface, 179
721 D eep dorsal m etatarsal nerve, 619 action, 181
I n dex 901
Diaphragm (Continued) Distal lateral cutaneous branch (Continued) D orsal nerve ( Continued)
crura, 180 o f third intercostal nerve, 594 o f penis, 614
innervation, 181 Distal m uscular branch D orsal nuchal line, 12
lymph vessels, 441 o f intercostal nerve, 595 D orsal nucleus o f vagus nerve, 512
nerves, 578 o f musculocutaneous nerve, 583 D orsal palpebral artery, 301
pars costalis, 181 Distal palm ar venous arch, 405 D orsal pancreatic duct, 709
pars lumbalis, 180 D istal plantar venous arch, 417 D orsal papillary muscle, 281
pars sternalis, 181 Distal radioulnar jo in t, 113 D orsal parietal cartilage, 714
pelvic, 191 Distal tibiofibular joint, 127 D orsal p art o f liver, 699
thoracic surface, 179 D olichocephalic head, 8 D orsal petrosal sinus, 421
Diaphragma, 178 D orsal and ventral arch o f atlas, 51 D orsal portion o f pons, 503
Diaphragm a pelvis, 191 D orsal and ventral tubercle of atlas, 51 D orsal posterior alveolar artery, 312
D iaphragma sellae, 540, 812 D orsal artery o f penis, 383 D orsal proper digital arteries, 377
Diaphragmatic lobe D orsal atlanto-occipital m em brane, 101 lateral and medial, 337
of left lung, 735 D orsal branch D orsal proper digital nerves, 619
of right lung, 738 o f cervical nerves, 575 D orsal ramus
D iaphragmatic pleura, 733 o f coccygeal nerve, 622 o f accessory nerve, 570
Diaphragmatic surface o f first cervical nerve, 574 o f oculom otor nerve, 547
of heart, 274 o f intercostal arteries, 342 D orsal root ganglion, 471
of liver, 699 o f lateral saphenous vein, 409,417 Dorsal roots o f spinal nerves, 533, 536,572
of lung, 735 o f lum bar arteries, 355 dorsal aspect, 534, 535
Diaphysis o f long bones, 3 o f lum bar nerves, 595 D orsal sacral plexus, 610
Diarthrosis, 95 o f medial saphenous vein, 409, 417 D orsal sacral trunk, 610
Diencephalon, 480, 493-7 o f plantar proper digital nerve, 622 D orsal sagittal sinus, 419
mid-sagittal section, 811 o f radial artery, 337 D orsal spinocerebellar tract, 512
Digastric nerve, 559 o f sacral nerves, 610 D orsal sulcus, 28
Digastricus muscle, 144 o f saphenous artery, 367 D orsal surface
Digestive system, 645 o f second cervical nerve, 574 o f hindpaw , muscles, 262
and abdomen, 645-712 o f spinal nerve, 572 o f left lobe of pancreas, 708
Digit(s) o f ulnar nerve o f forepaw, 590 D orsal tegm ental decussation, 501
fifth, of forepaw, special muscles, 230 D orsal buccal gland, 840-42 D orsal thalam us, 494
first, arteries of, dorsal aspect, 376 D orsal buccal nerve, 563 D orsal thoracic nerve, 591
of forepaw, special muscles, 224 D orsal cerebellar veins, 424 D orsal vagal trunk, 632
special muscles, 263 D orsal cerebral vein, 398,422 D orsal venous rete o f carpus, 405
fourth, arteries of, medial aspect, 597 D orsal cervical cardiac nerve, 636-7 D orsal ventricular plexus, 637
medial aspect, 226 D orsal coccygeal trunk, 622, 623 D orsal wall o f stom ach, 679
nerves of, medial aspect, 597 D orsal cochlear nucleus, 508 D orsointerm ediate sulcus, 533
of right forepaw, arteries, medial as D orsal columns, 536 D orsolateral antebrachial muscles, 210-17
pect, 343 D orsal com m on digital arteries, 337,377 D orsolateral sulcus o f spinal cord, 533
second, of forepaw, special muscle, 230 D orsal com m on digital nerves, 619 D orsum linguae, 654
Digital arteries. See Arteries, digital. D orsal condyloid fossa, 12 D orsum o f tongue, 654
Digital impressions, 45 D orsal cutaneous branches D orsum penis, 763
Digital nerves o f intercostal arteries, 342 D orsum sellae, 18,45, 810
common, dorsal, 619 o f lum bar nerves, 599 Ductless glands, 807
dorsal, 587, 590 D orsal digital nerves, 587, 590 Ducts
palmar, common, 590 D orsal digital veins, 403,416 alveolar, 728
plantar, common, 622 D orsal esophageal trunk, 632 bile. See Bile ducts.
lateral, 622 Dorsal external arcuate fibers, 511 cochlear, m em branous, 859
medial, 619 D orsal external vertebral venous plexus, 428 cystic, 705, 706
proper, 622 Dorsal flexion, 98 hepatic. See Hepatic ducts.
proper, dorsal, 619 D orsal funiculi, 536, 539 interlobular, 705
palmar, 591 D orsal gray horn, 536 lacrimal, 838
Digital skeleton, 75 D orsal intercostal veins, 399 lymphatic, right, 435
Digital veins D orsal interoccipital sinus, 422 m andibular, 659
dorsal, 403, 416 D orsal interosseous artery, 333 mesonephric, 796
palmar, 405 Dorsal labial artery, 297 nasolacrimal, 716, 718, 838, 866
plantar, o f hindpaw, 417 D orsal labial nerves, 555 nasopalatine, 646, 866
Digital venous arches, 416 D orsal labial vein, 394 o f Bellini, 746
Digiti plantares communes, 622 D orsal m edian sulcus, 523 o f pancreas. 708-709
Dilator pupillae muscle, 846 o f spinal cord, 533 papillary. See Papillary ducts.
Diploe, 3,4 ,4 2 4 D orsal mesentery, com m on, 673 parotid, 658
Diploic veins, 424 D orsal m etacarpal arteries, 337 pronephric, 796
frontal, 393 D orsal m etacarpal nerves, 587 semicircular, m em branous, 859
Disc, intervertebral, 51, 103 D orsal m etacarpal veins, 403,404 sublingual, 660
Disci intervertebrales, 103 D orsal m etatarsal arteries, 371 thoracic, 431
Discus articularis, 97,99 D orsal m etatarsal nerves tracheal, 435
Discus intervertebralis, 51 deep, 619 Wolffian, 796
Dislocations, 98 superficial, 618 D uctuli alveolares, 728
Distal articular surface o f tibial tarsal bone, D orsal m uscular branch o f external ethm oi D uctuli biliferi, 705
89 dal artery, 305 D uctuli hepaticae, 705
Distal collateral radial artery, 329 D orsal nasal artery, 312 D uctuli interlobulares, 705
Distal communicating vein, 401 D orsal nasal concha, 27 D uctus choledochus, 706
Distal dorsal interosseous artery, 333 Dorsal nasal ligament, 716 D uctus cysticus, 705, 706
Distal interphalangeal joints, 118 D orsal nasal m eatus, 28, 716, 863 Ductus deferens, artery of, 378
Distal intertarsal joint, 127 D orsal nasal vein, 393 D uctus deferentes, 758
Distal lateral cutaneous branch D orsal nerve D uctus hepaticus communis, 705
of intercostal nerve, 595 o f clitoris, 614 D uctus lym phaticus dexter, 435
902 I n d ex
Hairy skin. See Skin, hairy. H epatic arteries, 345 H orm one (Continued)
Hallux, 88 proper, 704 lactogenic, 815
H am string muscles, 231 distribution of, 357 luteinizing, 783, 814
Hamulus o f spiral lam ina, 857 H epatic ducts, 705 m am m otrophic, 815
H am ulus pterygoideus, 34 visceral aspect, 707 metabolic, 808
H ard palate, 647 H epatic flexure, 692 morphogenetic, 808
H arder’s gland, 840 H epatic lobes, 702, 703 sex, female, 832
H assall’s bodies, 462 H epatic lobules, 704 male, 830
H ead H epatic lymph nodes, 447 som atotrophic, 814
arteries, 289-324 H epatic plexus, 640 thyrotrophic, 815
brachycephalic, 8 Hepatic veins, 407, 704 Horn, gray, 536
dolichocephalic, 8 H epatoduodenal ligam ent, 678 Horns, uterine, 786
fasciae, 161-2 H epatogastric ligament, 678 H orny claw, 887
fibular, ligament of, 127 H epatorenal ligament, 704 Humeral artery, 325, 328
frontal section, ventral aspect, 648 H erm aphroditism , 830 Humeral condyle, 69
humeral. See H um eral head. H ernia, perineal, 751 Humeral head
lymph nodes and vessels, 434-9 H eterotopic skeleton, 93 of m. flexor carpi ulnaris, 221
mesaticephalic, 8 H iatus aorticus, 180, 670 o f m. flexor digitorum profundus, 221
muscles, 134-62 Hiatus esophageus, 180, 670 Humeral ligament, transverse, 110
deep, dorsal aspect, 141 Hillock, seminal, 111 Humeroradial joint, 110
lateral aspect, 138 Hilus H um eroulnar joint, 110
lateral aspect, 167 o f lung, 735 Humerus, 64, 67-9
superficial, lateral aspect, 136, 137 o f lymph node, 434 body, 67, 69
nerves, 544-71 o f spleen, 458 caudal circumflex vein, 403
of caudate nucleus, 488 renal, 743 caudal surface, 69
of femur. See Femur, head. Hilus lienis, 458 collateral radial artery, 325
of fibula, 88 Hilus pulmonis, 735 cranial surface, 69
of hum erus, 67 Hilus renalis, 743 h e a d ,67
o f metacarpus, 74 H indbrain, 480 lateral surface, 67
o f m etatarsal bone, 92 Hindleg, right left, caudal aspect, 68
o f muscle, 132 superficial veins of, lateral aspect, 411 cranial lateral aspect, 68
o f radius, 69 medial aspect, 412 lateral aspect, 68, 203
of rib, 61 H indpaw, 78, 88-92 medial aspect, 203
of tibial tarsal bone, 89 arteries, 371 medial surface, 67
postganglionic parasym pathetic fibers in, left, extensor muscles, 260 neck, 67
629 muscles, 259, 261 nutrient artery, 325
preganglionic parasym pathetic fibers in, muscles, 262-5 Hyaline cartilage joints, 96
629 nerves, 619-22 H ydatides terminales, 786
radial, o f m. flexor digitorum profundus, phalanges, 92 H ydatids of M orgagni, 786
221 right, arteries of, dorsal aspect, 376, 621 H yoepiglotticus muscle, 161
superficial veins, lateral aspect, 396 p lan tar aspect, 624 Hyoglossus muscle, 154
sympathetic distribution o f autonomic deep arteries, p lan tar aspect, 375 Hyoid apparatus, 38, 100, 155
nervous system, 634-5 nerves of, dorsal aspect, 621 bones, 38
ulnar. See Ulnar head. p lan tar aspect, 624 dorsal aspect, 723
veins, ventral aspect, 397 superficial arteries, plan tar aspect, 374 lateral aspect, 40
Hearing, organ of, 837 veins of, 418 muscles, 148-50
Heart, 267-87 veins, 416-17 Hyoid bones, 38
and arteries, 267-388 Hinge joint, 98 anterior lateral aspect, 37
apex, 267, 274 H ip bone, 78 H yoid branches o f lingual artery, 296
base, 267, 274, 277 H ip joint, 119-22 Hyoid muscles
fibrous, 276 arteries, m edial aspect, 608 lateral aspect, 149
blood vessels, 283-7 muscles, 231-47 superficial, lateral aspect, 145
conduction system, 283 nerves, m edial aspect, 608 Hyoid venous arch, 394
dorsal aspect, 270 right, arteries of, dorsal aspect, 616 Hyopharyngeus muscle, 154
left lateral aspect, 271 lateral aspect, 609 action, 156
lymph vessels, 448 muscles of, lateral aspect, 609 innervation, 156
nerves to, 636-7 nerves of, dorsal aspect, 616 H yothyroid mem brane, 721
orientation, 273 lateral aspect, 609 H yperthyroidism, 822
right lateral aspect, 272 H ippocam pal commissure, 488 H ypertrophy o f prostate, 762
size, 273 H ippocam pal sulcus, 483 Hypochondriac regions, 672
surface topography, 212-4 H ippocam pus, 490 Hypogastric lymph node, 446
sym pathetic fibers to, routes, 637-8 fimbria of, 492 Hypogastric nerves, 638, 641
ventral aspect, 391 Hock, 88 Hypoglossal canal, 12, 45
weight, 273 Homologies o f genital organs in mammals, vein of, 421
H eat, 789 797 Hypoglossal foramen, 12. 42
Helicine pouch, cutaneous, 848 H orizontal fissure o f lungs, 738 Hypoglossal nerve, 512, 564, 571
H elicotrema, 857 H orizontal p art o f vomer, 34, 36 Hypoglossal nucleus, 512
Helix, 848 H orizontal ram us, posterior, o f splenial sul H ypophyseal arteries, 313
Hemal arches, 60 cus, 483 Hypophyseal capsule, 812
Hemal processes, 60 H ormone, 808 Hypophyseal cleft, 814
Hemiazygos vein, 399 adrenocorticotrophic, 815 Hypophyseal foramen, 812, 814
Hemispheres antidiuretic, 815 Hypophyseal fossa, 18, 45, 810
cerebral. See Cerebral hemisphere. catalytic action, 809 Hypophyseal stalk, 810
o f cerebellum, 504 follicle-stimulating, 783, 814 Hypophysis, 810-16
H em orrhoidal artery, cranial, 351 growth, 814 arterial supply, ventral aspect, 310
Hepar, 699 interstitial cell-stim ulating, 815 blood supply, 812
In d ex 909
Lateral internal auricular nerves, 559 Ligamenta flava, 106 Ligaments (Continued)
Lateral lemniscus, 500, 503 Ligam enta glenohum eralia m edialis et late o f head o f rib, 106
Lateral m am m ary branches o f intercostal ralis, 110 o f left forepaw, dorsal aspect, 116
nerve, 595 Ligamenta interdigitalia, 118 ofleft stifle joint, 124, 125
Lateral masses o f atlas, 51 Ligamenta interspinalia, 106 dorsal aspect, 126
Lateral meniscus, caudal tibial ligament, 123 Ligamenta intertransversaria, 106 ofleft tarsus, 128
Lateral nasal artery, 312 Ligamenta m etacarpea interossea, 114 o f neck o f rib, 106
Lateral nasal gland, 718 Ligamenta ossiculorum auditus, 99 o f nose, 716
Lateral nasal ligament, 716 Ligam enta palpebrales, 837 of occiput, 102
Lateral nasal nerves, 555 Ligamenta sesamoidea cruciata, 118 o f ossicles, 856
Lateral nasal vein, 393 Ligam enta sternocostalia radiata dorsalia et o f ovaries, 780
Lateral nuclear group, 496 ventralia, 108 o f patella, 123
Lateral olfactory gyrus, 490 Ligamenta umbilicales laterales, 749 o f pelvic limb, 119-29
Lateral olfactory stria, 490 Ligaments, 97, 98, 673 of pelvis. See Pelvis, ligaments.
Lateral palm ar digital nerve V, 590 acetabular, transverse, 119 o f ribs. See Ribs, ligaments.
Lateral palm ar p roper digital nerves, 591 annular. See Annular ligament. o f skull, 99-101
Lateral p art o f frontal sinus, 49 apical, o f dens, 101 o f stifle joint, 122-7
Lateral p lan tar digital nerve, 622 atlantal, transverse, 103 o f tarsus, 129
Lateral p lan tar nerve, 622 atlanto-occipital, lateral, 101 o f thoracic cage, lateral aspect, 107
Lateral regions, 672 broad, 779-80, 787 o f thoracic limb, 108-118
Lateral reticular nucleus, 523 o f uterus, 674 o f tubercle o f rib, 106
Lateral saphenous vein, 409 capsular, 96 o f uterus, 787
Lateral sigmoid gyrus, 484 carpal, palm ar, transverse, 113 ofvertebral colum n. See Vertebral column,
Lateral spinothalam ic tract, 523 caudal, o f auditory ossicles, 100 ligaments.
Lateral sulcus, 483 collateral, 97 o f xiphoid region, 108
Lateral surface o f body o f m andible, 36 ofinterphalangeal joints, 118 orbital, 32, 39, 843
Lateral tarsal vein, 417 o f m etacarpophalangeal joints, palpebral, 837, 842
Lateral thoracic artery, 324 114 patellar, straight, 85
Lateral thoracic nerve, 591 conjugal, 106 pulmonary, 734
Lateral umbilical ligaments, 749 coronary, o f liver, 703 radial collateral. See Radial collateral liga
Lateral ventricle, 482 costoxiphoid, 108 ment.
Lateral vestibular nucleus, 511 cricothyroid, 721 reflected, 182
Lateral vestibulospinal tract, 511 cruciate, 123 rostral, o f auditory ossicles, 100
Latissimus dorsi muscle, 201 o f left stifle joint, m edial aspect, 126 round, 122
Latus, 672 o f sesamoid bones, 118 fissure for, 702
Leaf, contrahentes, 230 o f stifle, 123 o f uterus, 787
Leg. See also Crus. deep, o f left forepaw, p alm ar aspect, 117 sacroiliac, 119
right, arteries of, lateral aspect, 617 denticulate, 542 sacrotuberous, 119
muscles of, lateral aspect, 617 dorsal, o f auditory ossicles, 99, 100 scrotal, 755
nerves of, lateral aspect, 617 ofinterphalangeal joints, 118 sesamoidean, 114, 118
true, muscles of, 249-62 duodenocolic, 686 short, o f vertebral column, 103-106
Lemniscus falciform, o f liver, 704 sternocostal radiate, 108
lateral, 500, 503 femoral, o f lateral meniscus, 123 sternopericardiac, 267
medial, 496, 501,503,523 femoropatellar, 123 superficial, o f left carpus, palm ar aspect,
decussation of, 523 femorotibial, 123 1/5
trigeminal, 496, 501, 503, 523 fibular collateral, 123, 129 supraspinous, 103
Lemniscus lateralis, 500 gastrosplenic, 458,675 tibial, o f meniscus, 122, 123
Lemniscus medialis, 496 glenohumeral, 110 tibial collateral, 123, 129
Lemniscus trigeminalis, 496, 523 hepatoduodenal, 678
Lens o f eye, 847 tibiofibular, 127
hepatogastric, 678 transverse, carpal, palm ar, 231
Lentiform nucleus, 488 hepatorenal, 704
Lesser cruciate sulcus, 483 of stifle joint, 123
humeral, transverse, 110 triangular, 703, 704
Lesser curvature o f stom ach, 681 inguinal, 183, 186
Lesser om entum , 675, 678-9, 704 ulnar collateral. See Ulnar collateral liga
interarcuate, 106 ment.
Lesser peritoneal cavity, 678
interdigital, 118 umbilical. See Umbilical ligaments.
Lesser petrosal nerve, 627 interosseous, o f antebrachium , 113 urogenital, o f male, ventral aspect, 759
Levator ani muscle, 191 intersesam oidean, 114 vocal, 726
action, 194 interspinous, 106 yellow, 106
innervation, 194 intertransverse, 106 Ligamentum, 97
Levator nasolabialis muscle, 140 lateral, o f auditory ossicles, 99 Ligamentum apicis dentis, 101
L evator palpebrae superioris muscle, 148, long, o f vertebral colum n, 103 Ligamentum annulare radii, 110
842 longitudinal, 103 Ligamentum arteriosum, 288
Levator veli palatini muscle, 156 m andibular, 99 Ligamentum atlanto-occipitalis lateralis,
Levatores costarum muscles, 176 meniscal. See Meniscal ligaments. 101
action, 178 m etacarpal, interosseous, 114 Ligamentum capitis femoris, 119, 122
innervation, 178 nasal, 716 Ligamentum capitis fibulare, 127
Levators, 143 nuchal, 103,104 Ligam entum capituli costae, 106
o f tail, 190 oblique, 110 Ligam entum carpi palm are transversum, 113
LH. See Luteinizing hormone. o f atlas, 102 Ligam entum collaterale fibulare, 123, 129
Lien. See Spleen. o f auditory ossicles, 99, 100 Ligamentum collaterale radiale, 110
Ligam enta alaria, 101 o f axis, 102 Ligamentum collaterale radiale breve, 114
Ligam enta annularia trachealia, 728 o f bladder, 749 Ligam entum collaterale tibiale, 123, 129
Ligam enta collateralia, 97,114 o f conus, 276 Ligamentum collaterale ulnare, 110
Ligamenta costoxiphoidea, 108 o f fibular head, 127 Ligamentum collaterale ulnare breve, 114
Ligamenta cruciata genus, 123 o f flexed carpus, dorsal aspect, 116 Ligamentum colli costae, 106
Ligam enta dorsalia ofinterphalangeal joints, o f forepaw, lateral aspect, 117 Ligamentum conjugate costarum , 106
118 o f head o f femur, 119, 122 Ligamentum coronarium hepatis, 703
In d ex 913
Ligamentum cricothyroideum, 721 Linea anorectalis, 694 Lobus occipitalis of cerebrum , 482
Ligamentum cruciatum caudale, 123 Linea mylohyoidea, 36 Lobus pancreatis dexter, 708
Ligamentum cruciatum craniale, 123 Linea nuchalis dorsalis, 12 Lobus pancreatis sinister, 708
Ligamentum denticulatum , 542 Linea nuchalis ventralis, 12 Lobus parietalis of cerebrum , 482
Ligamentum duodenocolicum, 686 Linea obliqua o f thyroid cartilage, 719 Lobus quadratus of liver, 702
Ligamentum falciforme hepatis, 704 Linea terminalis, 80 Lobus tem poralis o f cerebrum , 482
Ligamentum femorale menisci lateralis, 123 Linea transversa, 56, 84 Loin, muscles, 231-47
Ligam entum fem oropatellare m ediale et la- Lineae temporales, 14 Long ciliary nerve, 550
terale, 123 Lingua, 654 Long levator, 143
Ligamentum gastrolienale, 458, 675 Lingual artery, 296 Long thoracic nerve, 591
Ligamentum hepatoduodenale, 678 medial aspect, 299 Longissimus atlantis muscle, 168
Ligamentum hepatogastricum , 678 Lingual frenulum , 654 Longissimus capitis muscle, 168
Ligamentum hepatorenale, 704 Lingual nerve, 556, 564 Longissimus cervicis muscle, 168
Ligamentum inguinale, 183, 186 Lingual papillae, 868 Longissimus lum borum muscle, 166
Ligamentum interossei antebrachii, 113 Lingual ramus, 567 Longissimus muscle, 166
Ligamentum intersesamoideum, 114 Lingual surface Longissimus system, 164, 166-8
Ligamentum latum uteri, 674 o f body o f mandible, 36 Longissimus thoracis et lum borum muscle,
Ligamentum longitudinale dorsale, 103 o fto o th , 651 166
Ligamentum longitudinale ventrale, 103 Lingual tonsil, 662 Longissimus thoracis muscle, 166
Ligamentum m andibulae, 99 Lingual vein, 394 innervation, 168
Ligamentum nasale dorsum, 716 Lingula o f sphenoid, 19 L ongitudinal bundles. See Bundle, longitu
Ligamentum nasale laterale, 716 Lingula sphenoidalis, 19 dinal.
Ligamentum nuchae, 103 Lips, 646 Longitudinal fissure, 482
Ligamentum obliquum, 110 acetabular, 119 Longitudinal layer o f m uscular coat of stom
Ligamentum patellae, 123 commissures, 646 ach, 683
Ligamentum pulmonale, 734 glenoid, 98, 108 Longus capitis muscle, 173
Ligamentum reflexum, 182 muscles, 134-9 action, 175
Ligamentum sacroiliacum dorsale breve et Liquor folliculi, 780 innervation, 175
longum , 119 Liquor pericardii, 267 Longus colli muscle, 175
Ligamentum sacroiliacum ventrale, 119 Liver, 672, 699-706 Loop, cervical nerve, 574
Ligamentum sacrotuberale, 119 blood vessels, 704-705 Lower dental arch, 651
Ligamentum sesamoideum distale, 118 borders, 699-702 Lower lips, 646
Ligam entum sesamoideum laterale et m edi coronary ligament, 703 L um bar arteries, 345, 355
ale, 114, 118 diaphragm atic aspect, 700 L um bar cistern o f spinal subarachnoid cav
Ligamentum sternopericardiaca, 267 falciform ligament, 704 ity, 542
Ligamentum supraspinale, 103 fibrous appendix, 704 Lum bar colonic nerve, 641
Ligamentum tarsi transversi distalis, 129 fibrous capsule, 703 Lum bar enlargem ent of spinal cord seg
Ligamentum tarsi transversi proximalis, 129 lobes, 702-703 ments, 533
Ligamentum teres uteri, 787 lymph vessels, 453, 704-705 Lum bar lymph nodes, 446
Ligamentum tibiale caudalis menisci late nerves, 705 L um bar nerves, 595-610
ralis, 123 parts, 699 lateral aspect, 603
Ligam entum tibiale caudalis menisci m edi peritoneal attachm ents, 703-704 m edial view, 602
alis, 123 peritoneal fixation, 703-704 Lum bar p art o f spinal cord, 533
Ligamentum tibiale cranialis menisci late physical characteristics, 699 Lum bar plexus, 606
ralis, 123 porta, 702 L um bar region, 672
Ligam entum tibiale cranialis m enisci m edi processes, 702-703 cross section, 195
alis, 123 relations, 699-702 Lum bar splanchnic nerves, 638
Ligamentum tibiofibulare caudale, 127 serous coat, 703 Lum bar veins, 399, 406
Ligamentum tibiofibulare craniale, 127 sinusoids, 704 Lum bar vertebrae. See Vertebrae, lumbar.
Ligamentum transversum acetabuli, 119 structure, 704 Lum bar vertebral veins, right lateral aspect,
Ligamentum transversum atlantis, 103 surfaces, 699-702 428
Ligamentum transversum genus, 123 visceral aspect, 701 Lumbococcygeal region, muscles, 192
Ligamentum transversum humerale, 110 Lobar bronchi, 728 Lum bosacral plexus, 599, 606
Ligamentum triangulare dextrum, 703 Lobes lateral aspect, 604
Ligamentum triangulare sinistrum, 704 anterior, o f cerebellum, 504 Lum bosacral trunk, 610
Ligamentum tuberculi costae, 106 of hypophysis, 810 Lum bricales muscles, 224, 263
Ligamentum umbilicale laterale, 378 flocculonodular, 504 Lunate surface o f acetabulum , 78
Ligamentum umbilicale medianum , 749 o f cerebrum, 482 Lungs, 730, 734-40
Ligamentum venosum, 705 o f liver, 702-703 apex, 734
Ligamentum vocale, 726 o f lungs, shapes, 735-8 base, 734
Limbus, 844 o f pancreas, 708 blood vessels, 738-9
Limbus fossae ovalis, 274 o f thymus, 461 cardiac notch, 738
Line posterior, o f cerebellum, 504 diaphragm atic surface, 735
anconeal, 67 Lobules hilus, 735
anocutaneous, 694 hepatic, 704 horizontal fissure, 738
anorectal, 694 param edian, 507 interlobar fissures, 735-8
iliopectineal, 81 Lobuli hepatis, 704 interlobar surfaces, 735
mylohyoid, 36 Lobuli testis, 755 left, 736
nuchal, 12, 23, 38, 44 Lobus caudatus o f liver, 703 lobes, 735
oblique, o f thyroid cartilage, Lobus dexter et sinister o f thymus, 461 lobes, shapes of, 735-8
719 Lobus frontalis o f cerebrum , 482 lymph vessels, 445
temporal, 14, 38 Lobus hepatis dexter, 702 lymphatics, 739-40
terminal, 80 Lobus hepatis dexter lateralis, 703 margins, 735
transverse, 56, 84 Lobus hepatis dexter medialis, 703 mediastinal part, 734
Linea alba, 189 Lobus hepatis sinister, 702 relations to other organs, 738
Linea anconea, 67 Lobus hepatis sinister lateralis, 702 right, 737
Linea anocutanea, 694 Lobus hepatis sinister medialis, 702 cardiac impression, 738
914 In d ex
Masseter muscle, 150 M edial genicular artery, 367 M em brana interossea cruris, 127
action, 152 M edial genicular vein, 409 M em brana Shrapnelli, 853
innervation, 152 M edial geniculate body, 496 M em brana stapedius, 856
Masseteric artery, 300 M edial head o f m. gastrocnemius, 252 M embrana sterni, 108
M asseteric border o f zygom atic bone, 3 1 Medial hepatic lobes, 702, 703 M embrana synovialis, 96
Masseteric branch o f posterior deep tem po Medial iliac lymph node, 446 M em brana tym pani, 23
ral artery, 303 M edial lemniscus, 496, 501, 503, 523 M embrane
Masseteric fascia, deep, 161 decussation, 523 atlanto-axial, dorsal, 101
Masseteric fossa, 36 M edial longitudinal bundle, 501, 503 atlanto-occipital, 101
Masseteric nerve, 556 Medial m am m ary branches of intercostal Bow m an’s, 843
Masseteric vein, 398 nerve, 595 Descemet’s, 843
M astication, muscles of, 150-52,153 Medial meniscus, caudal tibial ligament, 123 fibrous, o f join t capsule, 96
lateral aspect, 151 M edial muscles o f thigh, 242-7 hyothyroid, 721
M asticatory surface o f tooth, 651 M edial (m uscular) branches o f cervical interosseous. See Interosseous membrane.
Mastitis, 803 nerves, 575 nictitating, 838
M astoid process, 22 M edial nuclear group, 496 o f eye, 843
Maxilla, 28-31 M edial olfactory gyrus, 490 serous, peritoneal, 672
external surface, 28, 29 M edial olfactory stria, 490 sternal, 108
medial aspect. 29 M edial palm ar proper digital nerves, 591 subsynaptic. 475
nasal surface, 30, 31 M edial p art o f frontal sinus, 49 synovial, 96
pterygoid process, 30 Medial plantar digital nerve, 619 tympanic. 23, 848, 851
sutures, 100-101 Medial p lan tar nerve, 619 M em brane bones, 4
Maxillary alveolar nerves, 555 M edial retropharyngeal lym ph node, 435, M embranous cochlear duct, 859
Maxillary artery, 301 438 M embranous labyrinth, 856. 859
external, 296 M edial saphenous vein, 409 M em branous nasal septum , 714
terminal branches, 298 M edial surface o f lung, 734 M em branous p art o f interventricular sep
lateral aspect, 302 M edial tarsal vein, 417 tum. 281
Maxillary border o f zygomatic bone, 31 M edial vestibular nucleus, 511 M embranous portion o f m ale urethra, 111
M axillary branch o f trigeminal nerve, lateral M edian aperture o f fo urth ventricle, 523, 542 M embranous sem icircular ducts, 859
aspect, 553 M edian artery, 328, 329 M em branous vestibule, 859
Maxillary foramen. 30 M edian coccygeal artery, 386 M em brum pelvinum, 78
M axillary fossa, 25, 33 M edian crest o f cricoid cartilage, 721 M em brum thoracicum , 64
Maxillary incisure, 16 M edian cubital vein, 401 M eningeal artery
Maxillary nerve, 554-5, 649 M edian fissure, ventral, 507, 533 anterior, 304
M axillary process, 32, 33 M edian groove o f tongue, 654 caudal. 293
M axillary recess, 30, 49 M edian nerve, 586 middle, 303
Maxillary sinus, 25, 30, 49 o f forepaw, 590 sulcus for, 14, 18
M axillary tuberosity, 30 M edian sacral artery, 386 Meningeal veins, 424
Maxillary vein M edian sacral vein. 416 Meninges, 539-42
external, 393 M edian sulcus, dorsal, 523 and spinal cord, 533-43
internal, 398 o f spinal cord, 533 cranial. 539-41
Maxillonasolabialis muscle, 135 M edian vein, 403 spinal, 541-2
M axilloturbinate, 27, 48 M ediastinal branches Meniscal ligaments, 122
cross section, 26 o f internal thoracic artery, 321 of left stifle joint, m edial aspect, 126
M axilloturbinate bone, 31 o f thoracic aorta, 342 M eniscom andibular com partm ent, 99
M axilloturbinate crest, 30, 48 M ediastinal cavity, 731, 732 M eniscotem poral com partm ent, 99
Meatus M ediastinal lymph nodes, 440 Meniscus. 97, 122
acoustic. See Acoustic meatus. M ediastinal part o f lung, 734 articular, 99
auditory, external, 851 M ediastinal pleura, 732, 733 lateral. 85
nasal. See Nasal meatus. M ediastinum, 181,731-2 ligaments of, 123
nasopharyngeal, 48, 718 lymph vessels. 441 medial, 85
temporal, 23 M ediastinum testis, 755 ligaments of, 122. 123
Meatus acusticus externus, 22 M edulla o fle ft stifle joint, dorsal aspect, 126
M eatus acusticus internus, 19 internal, o f kidney, 743 o f stifle joint, 122
M eatus nasi communis, 25, 3 1, 48, 718, 863 o f lymph node, 434 Meniscus articularis, 97
Meatus nasi dorsalis, 28, 48, 716, 863 o f ovary, 780 Meniscus lateralis, 122
M eatus nasi medius, 31, 48, 716, 866 M edulla oblongata, 507 Meniscus lateralis et medialis, 122
Meatus nasi ventralis, 31, 48, 718, 866 Medulla ossium flava, 4 M eniscus medialis, 122
Meatus nasopharyngeus, 48, 7 18 Medulla ossium rubra. 4 Meniscus medialis et lateralis. 85
Meatus temporalis, 23 M edulla renis, 743 M ental arteries, 303
Medial angle o f eyelid, 838 Medulla spinalis, 533 M ental foramen, 36
Medial auricular artery, 300 Medullary cavities. 4 M entalis muscle, 139
Medial auricular vein, 399 Medullary lamina, 488 M erocrine sweat glands. 883
Medial branches external, 496 Mesaticephalic head, 8
of cervical nerves III—VII, 575 Medullary veins, 424 M esencephalic tract, 501
o f iliohypogastric nerve, 606 M edullary velum, 504, 523 M esencephalon, 480, 497-501
o f lumbar nerves, 599 Megacephalic skull, 10 M esenteric arteries
o f proximal collateral radial artery, 329 M eibomian glands, 838 branches, ventral aspect, 362
of radial nerve, 583 Meissner’s plexus, 633 caudal, 351
o f superficial radial nerve, 587 M em brana atlanto-axialis dorsalis, 101 cranial, 349
of thoracic nerves, 594 M em brana atlanto-occipitalis dorsalis. 101 branches of, ventral aspect. 361
Medial circumflex femoral artery, 363 M em brana atlanto-occipitalis ventralis, 101 M esenteric lymph nodes, 447
Medial cutaneous antebrachial nerve, 583 M em brana fibrosa o f jo in t capsule. 96 Mesenteric plexus, 640
Medial dorsal digital nerves, 587, 590 M em brana granulosa o f ovary, 832 Mesenteric portion o f small intestine, 687
Medial dorsal proper digital nerves, 619 M em brana hyothyroidea, 721 M esenteric vein, 407
M edial femoral lymph node, 454 M em brana interossea antebrachii. 113 M esenterium, 673, 687
916 In dex
M esenterium dorsale com m une, 673 Middle ectosylvian sulcus, 483 Muscles fContinued)
M esentery, 673 M iddle m axillary alveolar nerves, 555 cardiac, 131
o f colon, 692-3 M iddle mediastinal cavity, 731 circumduction, 133
o f small intestine, 687-8 Middle m ediastinal pleura. 733 coccygeoanal, 697
root of. 688 Middle m em brane of eye, 843 constrictor. See Constrictor muscles.
M esethmoid, 24 Middle m eningeal vein, 424 constrictor vestibuli, 794
Mesocephalic skull, 10 M iddle m ental artery, 303 constrictor vulvae, 794
M esocolon, 673, 692 M iddle nasal meatus, 716, 866 corrugator supercilii, 842
ascending, 693 M iddle plexus, 885 cricoesophageal, 665
descending, 674, 693 M iddle rectal artery, 379 deep, of ear, dorsal aspect. 141
transverse. 693 M iddle suprasylvian gyrus, 484 lateral aspect, 138
Mesocolon ascendens, 693 M iddle suprasylvian sulcus, 483 o f face. 144-6
M esocolon descendens, 674, 693 M iddle tracheobronchial lymph node, 443 o f head, dorsal aspect, 141
M esocolon transversum , 693 Middle umbilical fold, 674 lateral aspect. 138
M esoduodenum , 673, 686 M iddle umbilical ligam ent, 749 deltoideus. See Deltoideus muscle.
M esogastrium, 673 M inor palatine artery, 308, 649 dilator pupillae. 846
M esojejunoileum, 673. 687 M inor palatine nerve, 554 epaxial. superficial, 165
Mesomere, 796 M inor petrosal nerve, 563 erector pili. 883
M esometrium, 779. 787 M itral area, 282 extensors, 133
M esonephric duct. 796 Mitral valve, 282 o f left hindpaw, 260
M esonephros, 796 M oderator band. 278 extraocular, 837. 840
M esorchium, 758, 761 M olar teeth. 652 extrinsic, of eyeball, 146-8
M esorectum, 673, 693 M onorchidism, 757 fiber, 131
Mesosalpinx, 779 Morgagni fixation. 133
M esotendon, 133 hydatids of, 786 flexors, 133
M esotympanicum. 851 lacunae of, 777 hamstring, 231
Mesovarium, 779 M orphogenesis head. 132
M etabolic horm ones, 808 relation o f hypophysis to, 815-16 hyoid. See Hyoid muscles.
M etacarpal arteries. 337. 339 relation o f thyroid gland to, 821-2 ischiocavernosus. See lschiocavernosus
M etacarpal bones, 74 M orphogenetic horm ones, 808 muscles.
left, dorsal aspect, 76 M otor end-plate, 479 ischiourethral, dorsal view. 768
palm ar aspect, 77 M otor neuron. See Neuron, motor. ischiourethralis. See Ischiourethralis mus
muscles on palm ar side of. 224 M otor units. 132 cles.
M etacarpal joints. 114-18 M outh. 645-6 laryngeal. See Laryngeal muscles.
M etacarpal ligaments, interosseous, 114 and associated structures, 645-60 layers, o f cecum, dorsal aspect. 690
M etacarpal nerves, 587, 590 angular artery, 297 of colon, dorsal aspect. 690
M etacarpal veins. 403. 404, 405 angular vein, 394 of ileum, dorsal aspect, 690
M etacarpals, 64 M ucoperiosteum, 5 leaf, orbitofrontoauricular, 139
M etacarpophalangeal jo in ts. 114 Mucosa levator palpebrae superioris, 842
M etacarpus. 74 laryngeal. 724-6 multipennate. 133
arteries, m edial aspect, 597 nasal. 718 ocular, extrinsic, arteries of. lateral as
base, 74 o f esophagus, m uscular layer, 665 pect. 306
body, 74 o f tongue, 654 o f abdom inal wall, 182-9
head, 74 septal, nerves of. 545 of anal region, 195
interosseous spaces, 114 ureteral, 748 o f arm. See Arm , muscles.
left, proxim al end. cross section. 115 M ucous coat o f basihyoid, dorsal aspect, 151
nerves, m edial aspect, 597 o f esophagus, 665 o f brachium, deep, 207
M etapophyses o f thoracic vertebrae, 54 o f large intestine, 697 o f cheek, 134-9
M etatarsal arteries. 371 o f small intestine, 688 o f crus, 249-62
M etatarsal bones. 92 of stomach, 684 o f dorsum o f nose, 139-40
left, 264 Mucous neck cells o f stomach, 685 o f esophagus, 666
dorsal aspect, 91 Multifidus cervicis muscle, 171 o f external ear, 140-44
plantar aspect, 90 Multifidus lum borum muscle. 171 dorsal aspect. 145
M etatarsal jo in ts o f pelvic limb, 129 Multifidus muscle. 170 o f eye, 147
M etatarsal nerves, 618, 619, 622 Multifidus thoracis muscle, 171 o f eyelids. 148, 842
M etatarsal veins o f hindpaw , 416. 417 M ultipennate muscles. 133 o f female perineum, caudolateral aspect.
M etatarsus. 92 Muscles 612
M etathalam ic nuclei, 496 abduction, 133 o f forehead, 139-40
M etencephalon, 480. 501-507 adduction. 133 o f forepaw. 222-30
M etestrus, 789, 802 agonists, 133 of gaskin. 249-62
M icrocephalic skull, 10 anal sphincter, internal, 698 o f gluteal region. 243
M idbrain, 480 antagonists, 133 of head. See Head, muscles.
M iddle articular surface o f tibial tarsal antebrachial. See Antebrachial muscles. o f hindpaw. 262-5
bone, 89 articular, 133 o f hip. 231-47
M iddle branch o f com m on hepatic artery. arytenoideus transversus, 159, 161 of hyoid apparatus. 148-50
346 axial, deep, 169 o f larynx, 159-61
Middle cardiac vein. 287 belly, 132 o f left crus, 257, 258, 259
M iddle cardiosym pathetic nerve, 636 between flexor tendons o f forepaw, 222-4 o f left hindpaw, 259, 261
Middle carpal joint. 113 biceps femoris. See Biceps fem oris muscle. o f leg, 249-62
M iddle cerebellar peduncle, 503, 504. 507 bipennate. 133 of lips, 134-9
M iddle cerebral artery, 313 brachial. 206-10 o f loin, 231-47
M iddle clunial nerves, 610 brachiocephalicus. See Brachiocephalicus of lumbococcygeal region, 192
M iddle colic artery. 350 muscle. o f male perineum , caudolateral aspect.
M iddle colic lymph node. 448 brachioradialis. See Brachioradialis mus 613
M iddle colic vein, 409 cle. of m andible, dorsal aspect, 15/
M iddle ear, 837, 847, 851-6 buccinatorius. See Buccinatorius muscle. o f mastication. See Mastication, muscles
Middle ectosylvian gyrus, 484 bulbocavernosus. 764 °f
In d ex 917
Nerves ( Continued) Nervi m etacarpales palm ares profundi, 590 Nervus hypoglossus, 512,571
peripheral, 473 N ervi m etacarpales palm ares superficiales, Nervus ilioinguinalis, 606
general structural and functional or 590 Nervus infraorbitalis, 555
ganization, 469-71 Nervi m etatarsei dorsales superficiales, 618 Nervus infratrochlearis, 554
peroneal (fibular), 618 Nervi m etatarsei plantares profundi, 622 Nervus intercostalis II, 594
petrosal. See Petrosal nerve. Nervi m etatarsei plantares superficiales, 619 Nervus intercostobrachialis, 594
phrenic, 578 Nervi nasales interni, 554 Nervus intermedius, 558
plantar, 619, 622 Nervi nasales laterales, 555 Nervus interosseus antebrachii anterior, 586
pterygoid, 555 Nervi olfactorii, 546 Nervus ischiadicus, 614
pterygopalatine, 554 Nervi perinei, 614 Nervus labialis caudalis. 614
pudendal, 611 Nervi pterygoidei, 555 Nervus lacrimalis, 550
radial, 583 Nervi pterygopalatini, 554 Nervus laryngeus caudalis, 570
rectal, caudal, 611 Nervi sacrales, 610 Nervus laryngeus cranialis, 570
rotator, 614 Nervi splanchnici pelvini, 615 Nervus laryngeus recurrens, 570, 630
sacral. See Sacral nerves. Nervi spinales, 572 Nervus laryngeus superior, 630
saphenous, 607 Nervi thoracales ventrales, 591 Nervus lingualis, 556
sciatic, 614 Nervi thoraci, 591 Nervus m andibularis, 555
scrotal, caudal, 614 Nervi vomeronasales, 546 Nervus massetericus, 556
spermatic, external, 607 Nervous system Nervus maxillaris, 554, 649
spinal. See Spinal nerves. autonomic, 626-44 Nervus m edianus, 586
splanchnic, 615, 638 parasym pathetic division, 626, 627-33 Nervus m edianus manus, 590
stylohyoid, 559 peripheral elements, 628 Nervus m etacarpalis dorsalis I, 587
subcostal, 594 sym pathetic distribution, in thoracic re Nervus m etatarsus dorsalis profundus, 619
sublingual, 556 gion, 635-6 N ervus m etatarsus plantaris superficialis,
suboccipital, 574 to head, 634-5 619
subscapular, 582 to neck, 634-5 Nervus m usculocutaneus, 582
superficial, o f neck, lateral aspect, 577 sym pathetic division, 626, 633-42 Nervus mylohyoideus, 556
supraorbital, 550 peripheral elements, 628 Nervus nasalis caudalis, 555
suprascapular, 582 central, 464, 480 Nervus nasalis externus, 554
sural, cutaneous, 615 veins of, 419-28 Nervus nasociliaris, 550
tem poral, deep. 556 introduction, 464-79 Nervus obturatorius, 610
terminal, 546 peripheral, 464 Nervus occipitalis major. 574
thoracic. See Thoracic nerves. Nervus abducens, 508, 558 Nervus oculomotorius, 547
thyroid, 635 Nervus accessorius, 511, 570 Nervus ophthalmicus, 550
tibial, 619 Nervus alveolaris caudalis maxillaris, 555 Nervus opticus. 547
to diaphragm , 578 Nervus alveolaris m andibularis, 556 Nervus palatinus accessorium, 555
to heart, 636-7 Nervus alveolaris medii m axillaris, 555 Nervus palatinus major, 554
to m. brachiocephalicus, 591 N ervus alveolaris rostralis maxillaris, 555 Nervus palatinus minor, 554
to pterygoid muscle, 555 Nervus auricularis m agnus, 575 Nervus peroneus (fibularis) communis, 618
to stapedial muscle, 559 N ervus auriculopalpebralis, 559 N ervus peroneus (fibularis) profundus, 618
to tensor tym pani muscle, 555 N ervus auriculotem poralis, 556 Nervus peroneus (fibularis) superficialis, 618
to tensor veli palatini muscle, 555 N ervus axillaris, 582 Nervus petrosus major, 558, 627
trigeminal. See Trigeminal nerve. Nervus brachiocephalicus, 591 Nervus petrosus m inor, 563, 627
trochlear. See Trochlear nerve. Nervus buccalis, 555 N ervus pharyngoesophageus, 570
tympanic, 563, 627 Nervus buccalis dorsalis, 563 Nervus phrenicus, 578
ulnar, 586 N ervus buccalis ventralis, 563 Nervus plantaris lateralis, 622
vagus, 567-70 Nervus canalis pterygoidei, 627 Nervus plantaris medialis, 619
vertebral, 635 N ervus cervicalis, 574 Nervus pterygoideus lateralis, 555
vestibular, 563 Nervus collector caudae dorsalis, 622 Nervus pterygoideus medialis, 555
vestibulocochlear, 563 N ervus collector caudae ventralis, 622 N ervus pudendus, 611
vomeronasal, 546 Nervus cutaneus antebrachii caudalis, 587 Nervus radialis, 583
zygomatic, 554 N ervus cutaneus antebrachii lateralis, 583 N ervus radialis m anus, 587
zygomaticofacial, 554 Nervus cutaneus antebrachii medialis, 583 Nervus rectalis caudalis, 611
zygomaticotemporal, 554 Nervus cutaneus brachii lateralis, 582 Nervus rotatorius, 614
Nervi auriculares caudales, 559 Nervus cutaneus femoris caudalis, 611 Nervus saphenus, 607
Nervi cervicales, 574 Nervus cutaneus femoris lateralis, 606 Nervus scrotalis caudalis, 614
Nervi clunii caudales, 611 Nervus cutaneus surae caudalis, 615 Nervus stapedius, 559
Nervi clunii medii, 610 Nervus cutaneus surae lateralis, 615 Nervus stylohyoideus, 559
Nervi clunium superiores, 599 Nervus cutaneus surae medialis, 615 Nervus subcostalis, 594
Nervi coccygei, 622 Nervus digastricus, 559 N ervus sublingualis, 556
Nervi digitales dorsales communes, 619 Nervus digitalis dorsalis I, 587 N ervus suboccipitalis, 574
Nervi digitales dorsales mediales et laterales Nervus digitalis dorsalis medialis V, 590 N ervus subscapularis, 582
II, III and IV, 587, 590 Nervus digitalis palm aris lateralis V, 590 Nervus suprascapularis, 582
Nervi digitales dorsales proprii mediales et Nervus digitalis plantaris lateralis, 622 Nervus suralis, 615
laterales, 619 Nervus digitalis plantaris medialis, 619 N ervus temporalis profundus, 556
N ervi digitales palm ares com m unes, 590, Nervus digitorum dorsalis lateralis V, 590 Nervus tensoris tympani, 555
591 Nervus dorsalis clitoridis, 614 Nervus tensoris veli palatini, 555
N ervi digitales p roprii palm ares laterales et Nervus dorsalis penis, 614 N ervus terminalis, 546
mediales, 591 Nervus erigens, 633 N ervus thoracicus longus, 591
Nervi digiti plantares proprii, 622 Nervus ethmoidalis, 554 Nervus thoracodorsalis, 591
Nervi iliohypogastrici craniales et caudales, Nervus facialis, 508, 558 Nervus thoracolateralis, 591
599 Nervus femoralis, 607 Nervus tibialis, 619
Nervi intercostales, 594 Nervus frontalis, 550 N ervus transversus colli, 575
Nervi labiales dorsales, 555 Nervus genitalis, 607 N ervus trigeminus, 503, 550, 649
Nervi lumbales, 595 Nervus glossopharyngeus, 511, 563, 649 N ervus trochlearis, 547, 550
N ervi m etacarp ales dorsales II, III an d IV, Nervus gluteus caudalis, 611 N ervus tympanicus, 563, 627
587 Nervus gluteus cranialis, 611 Nervus ulnaris, 586
In d ex 921
Ovaries (Continued) Palm ar carpal ligament, transverse, 113 Param edian lobule, 507
ligaments, 780 Palm ar carpal transverse ligament, 231 Paranasal sinuses, 49, 50, 866-7
nerves, 783 Palm ar com m on digital arteries, 339 Parapatellar fibrocartilages, 85, 123
structure, 780-83 Palm ar common digital veins, 405 caudal aspect, 126
Over-hair, 881 Palm ar digital nerves, 590 Paraphimosis, 778
Oviducts, 784-6 Palm ar digital veins, 405 Pararectal fossa, 693
anomalies and variations, 786 Palm ar interosseous artery, 333 Parasym pathetic division o f autonom ic ner
blood vessels, 786 Palm ar m etacarpal arteries, 337, 339 vous system, 626, 627-33
clinical considerations, 786 Palm ar m etacarpal nerves, 590 Parasym pathetic fibers in head region, 629
nerves, 786 Palm ar m etacarpal veins, deep, 405 Paraterm inal gyrus, 485, 492
structure, 784-6 Palm ar p ro p e r digital arteries, la te ra l and Parathorm one, 824
Ovulation, 833 medial, 339 P arathyroid III, 823
Oxytocin, 815 Palm ar proper digital nerves, 591 Parathyroid IV, 823, 824
Palm ar venous arch, 403, 405 Parathyroid bodies, 817
Palmaris brevis accessorius, 224 Parathyroid glands, 822-6
Palpebra dorsalis et ventralis, 838 development, 823-4
granulations, 540
P a c c h io n ia n Palpebra tertia, 838 gross anatom y, 822-3
Pads, foot, 876, 878 Palpebrae, 837 m icroscopic anatom y, 823
Palate, 647-9 Palpebral arteries, 301 pathology, 824-6
aponeurosis, 649 Palpebral branch o f auriculopalpebral nerve, physiology, 824
blood vessels, 649 559 ventral aspect, 818
bones, 27-38 Palpebral conjunctiva, 838 Parenchym a of m am m ary glands, 801
hard, 30, 32, 42, 647 Palpebral fascia, 842 Parenchym atous cells o f thyroid gland, 820
nerves of, 545 Palpebral fissure, 838 Parietal bone, 13-14
muscles, ventrolateral aspect, 158 Palpebral ligaments, 837, 842 digital impressions, 14
nerves, 649 Palpebral rim, 838 external surface, 14
soft, 647 Pam piniform plexus, 406 frontal border, 14
lymph vessels, 436 Pancreas, 706-10 interm ediate ridges, 14
muscles, 156 accessory, 708 internal surface, 14
structure, 647-8 blood vessels, 709-10 occipital border, 14
ventral aspect, 720 ducts, 708-709 sagittal border, 14
Palatine and sphenopalatine arteries, com endocrine function, 833-4 squam ous border, 14
mon trunk, 308 lobes, 708 sutures, 100
Palatine arteries, 308, 649 lymph nodes and vessels; 451, 709-10 ventral lateral aspect, 13
Palatine bone, 32-3, 101 nerves, 710 Parietal branches
dorsal medial aspect, 32 relations, 708 of abdom inal aorta, 355-9
Palatine branches o f ascending pharyngeal Pancreas accessorium, 708 of internal iliac artery, 383-6
artery, 296 Pancreatic angle, 708 o f thoracic aorta, 342—5
Palatine canal, 33 Pancreatic branches Parietal cartilage, 714
Palatine crest, 32 of caudal pancreaticoduodenal artery, 350 Parietal cells of stomach, 684
Palatine fissure, 28, 30, 42, 44 o f cranial pancreaticoduodenal artery, 346 Parietal diploic vein, 424
Palatine foramina, 30, 33, 39, 42 o f splenic artery, 349 Parietal layer o f serous pericardium , 267
Palatine glands, 647 Pancreaticjuice, 706 Parietal lobe of cerebrum , 482
Palatine muscles, 647 Pancreatic veins, 409 Parietal p art o f greater om entum , 675
Palatine nerve, 554, 555 Pancreaticoduodenal artery, 346, 350 Parietal peritoneum , 673
Palatine plexus, 649 Pancreaticoduodenal vein, 407, 409 Parietal plane, 39
venous, 398 Papilla incisiva, 646 Parietal pleura, 732
Palatine process, 28, 30 Papilla lacrimalis, 838 Parietal surface
Palatine sulcus, 30 Papilla parotidea, 645 of liver, 699
Palatine surface o f horizontal lamina, 32 Papillae o f spleen, 458
Palatine suture, 31, 33, 44 circumvallate, 871 Parieto-occipital suture, 14
Palatine tonsil, 662 conical, 868, 871, 873 Parietosphenoidal suture, 14, 19
Palatine veil, 647 duodenal, major, 706 Paroophoron, 786
Palatinus muscle, 156 epidermal, 883 Parotid artery, 300
Palatoethmoid suture, 27 filiform, 868, 869 Parotid duct, 658
Palatoethmoidal suture, 33 foliate, 868-71 Parotid gland, 657-8
Palatoglossal fold, 647 taste buds on, 872 Parotid lym ph node, 434
Palatolacrimal suture, 34 fungiform, 868, 869, 870 Parotid nerves, 556
Palatomaxillary suture, 31, 33 incisive, 646 Parotid papilla, 645
Palatopharyngeal arch, 647 lingual, 868 Parotideoauricularis muscle, 142
Palatopharyngeal muscle, 647 o f tongue, 654 Parotidom asseteric fascia, 161
Palatopharyngeus muscle, 156 optic, 846 Pars abdominalis
Palatum, 647 parotid, 645 o f esophagus, 664
Palatum durum, 647 renal, 743 of m. obliquus internus abdom inis, 183
Palatum molle, 647 sublingual, 660 Pars anterior o f hypophysis, 810
Palatum osseum, 30, 32 vallate, 868, 871,572 Pars ascendens o f duodenum , 686
Paleocortex, 482 Papillae linguales, 654 Pars basilaris, 12
Paleopallium, 482 Papillae m ammae, 799 Pars basilaris pontis, 501
Pallium, 482 Papillary duct, 801 Pars brachialis of brachiocephalicus, 201
Palmar antebrachial artery, 333 o f kidney, 746 Pars buccalis o f hypophysis, 814
Palmar antebrachial vein, 403 Papillary muscles, 278, 281 Pars bursalis o f greater om entum , 675
Palmar arch. See Arch, palmar. Papillary process o f liver, 699, 703 Pars cardiaca of stomach, 681
Palmar arterial arch, superficial, 337 Paraflocculus, 504 Pars cervicalis
Palmar branches Parafollicular cells o f thyroid gland, of esophagus, 664
of radial artery, 337 820 ofm . cleidocephalicus, 201
of ulnar nerve o f forepaw, 590 Parahippocam pal gyrus, 485 of spinal cord, 533
Palmar carpal fibrocartilage, 114 Parahypophysis, 812, 814 of thymus, 461
924 In d ex
Plexus venosus palatinus, 398 Postganglionic parasym pathetic fibers in Processes (Continued)
Plexus venosus rectalis, 414 head region, <529 pterygoid. See Pterygoid processes.
Plexus venosus urethralis, 414 Postlateral gyrus, 484 retroglenoid, 23, 42
Plexus venosus vaginalis, 416 Postlateral sulcus, 483 sphenoidal, 33
Plexus venosus vertebralis externus dorsa Postsphenoid, 16 spinous. See Spinous process.
lis, 428 Postsplenial gyrus, 484 styloid. See Styloid process.
Plexus venosus vertebralis externus ven Postsplenial sulcus, 483 supraorbital, 16
tralis, 428 Postzygapophysis, 51 temporal, 32
Plexus venosus vertebralis internus, 428 Potential transverse, of lum bar vertebrae, 56
Plexus vestibuli, 414 action, 466 of thoracic vertebrae, 54
Plica, fimbriated, 654 resting, 466 of vertebral arch, 51
Plica alaris, 716 Pouch uncinate, o f ethmoid, 25
Plica annularis, 662 helicine, cutaneous, 848 vaginal. See Vaginal process.
Plica aryepiglottica, 724 o f Rathke, 814 xiphoid, 64
Plica fimbriata, 654 Prussak’s, 853 zygomatic, 16, 23, 30
Plica gastropancreatica dextra, 679 rectogenital, 749 Processus accessorii, 54
Plica gastropancreatica sinistra, 679 rectovesical, 749 Processus alveolaris, 28
Plica ileocecalis, 687, 689 vesicogenital, 749 Processus anconeus, 69, 72
Plica semilunaris, 662 vesicopubic, 749 Processus angularis, 36
Plica synovialis, 96 Preauricular muscles, 142 Processus articularis caudalis, 51
Plica sublingualis, 645 Precava, 274, 389-401 Processus articularis cranialis, 51
Plica suspensoria ovarii, 780 Precommissural area, 485, 492 Processus caudatus o f liver, 703
Plica umbilicalis medialis, 674 Precruciate gyrus, 484 Processus clinoideus anterior, 18
Plica venae cavae, 734 Precruciate sulcus, 483 Processus clinoideus posterior, 18
Plica vestibularis o f larynx, 724 Prefrontal gyrus, 484 Processus condylaris s. articularis, 36
Plica vocalis, 726 Preganglionic parasym pathetic fibers in Processus coracoideus, 67
Plicae gastricae, 684 head region, <529 Processus corniculatus of arytenoid carti
Plicae latae uteri, 779 Pregnancy, false, 789 lage, 721
Plicae tubariae, 786 Premaxilla, 27 Processus coronoideus, 36, 72
Pogonion o f skull, 8 Prem olar teeth, 651 Processus cuneiformis of arytenoid carti
Point, fixed, o f cranial attachm ent o f esoph Prepubic tendon, 183 lage, 721
agus, 665 Prepuce, 778-9 Processus frontalis, 28, 30, 32, 34
Pons, 501 Preputial muscle, 778 Processus hemales, 60, 190
Pontine nuclei, 503 Preputialis muscle, 189 Processus interparietalis, 12
Pontine veins, 424 Preputium, 778 Processus jugularis, 12
Popliteal artery, 367-71 Presphenoid, 16 Processus lacrimalis, 31
Popliteal lymph node, 454 anterior lateral aspect, 17 Processus lateralis tali, 89
Popliteal notch, 87 dorsal aspect, 17 Processus lenticularis, 856
Popliteal region, arteries, m edial aspect, 372 Pressoreceptor afferents, 637 Processus mammillares, 54
Popliteal surface o f fem ur, 84 Presylvian sulcus, 482 Processus mastoideus, 22
Popliteal vein, 413 Pretracheal plexus, 637 Processus maxillaris, 32, 33
Popliteus muscle, 262 Prevertebral fascia, 176 Processus medialis et lateralis o f fibular tar
Pore, acoustic, internal, 45 Prevertebral ganglia, 626 sal bone, 89
Porta hepatis, 702 Prezygapophysis, 51 Processus muscularis of arytenoid cartilage,
Porta o f liver, 702 Processes 721
Portal lymph nodes, 447 accessory, See Accessory processes. Processus nasalis, 16, 28
Portal vein, 407-409, 704 acoustic, external, 39 Processus palatinus, 28, 30
ventral aspect, 410 alveolar. See Alveolar process. Processus papillaris, 699
Porus acusticus internus, 19 anconeal, 69, 72 of liver, 703
Postauricular muscles, 142 angular, 36 Processus pterygoidei, 19, 30
Postcava, 274, 405-407 articular. See Articular process. Processus retroglenoideus, 23
notch for, 738 caudal. See Caudal process. Processus sphenoidalis, 33
ventral aspect, 408 caudate, o f liver, 703 Processus spinosus, 51
Postcaval foramen, 670 ciliary, 844 Processus styloideus, 71, 72
Postcruciate gyrus, 484 clinoid. See Clinoid process. Processus temporalis, 32
Postcruciate sulcus, 483 condyloid, 36 Processus transversus, 51
Posterior cerebral vein, 422 coracoid, 67 Processus uncinatus, 25
Posterior cham ber o f eye, 846 coronoid, 36, 72 Processus vaginalis, 787
Posterior commissure, 493 cranial, 51 Processus vocalis of arytenoid cartilage, 721
Posterior crus o f internal capsule, 488 frontal, 28, 30, 32, 34 Processus xiphoideus, 64
Posterior ectosylvian gyrus, 484 hemal, 60 Processus zygomaticus, 16, 23, 30
Posterior ectosylvian sulcus, 483 interparietal, 12 Proestrus, 789
Posterior horizontal ram us o f splenial sul jugular, 12, 39, 42 Progesterone, 833
cus, 483 lacrimal, 31 Prognathism o f m andible, 8
Posterior hypophyseal artery, 313 lateral, o f fibular tarsal bone, 89 Projection fibers, 486, 488
Posterior intercarotid artery, 313 o f talus, 89 Prolactin, 815
Posterior lobe mammillary. See Mammillary processes. Prolapse o f vagina, 790, 791
o f cerebellum, 504 mastoid, 22 Prominence, laryngeal, 721
o f hypophysis, 810 maxillary, 32, 33 Prominentia laryngea, 721
Posterior m edullary velum, 504, 523 medial, o f fibular tarsal bone, 89 Prom ontorium , 20, 58
Posterior m ental artery, 303 muscular, o f arytenoid cartilage, 721 of middle ear, 853
Posterior rhinal sulcus, 482 nasal, 16, 28 Promontory
Posterior sigmoid gyrus, 484 odontoid, 52 of pyramid, 20
Posterior suprasylvian gyrus, 484 o f arytenoid cartilage, 721 o f sacrum, 58
Posterior suprasylvian sulcus, 483 o f liver, 702-703 Pronator quadratus muscle. 222
Posterior sylvian gyrus, 484 palatine, 28, 30 Pronator teres muscle, 217
Posterolateral fissure, 504 papillary, 699, 703 innervation, 220
In d ex 927
Pronephric duct, 796 Pudendoepigastric trunk, 360, 414 Radices craniales o f accessory nerve, 570
Pronephros, 796 Pudendum femininum, 791 Radices dorsales o f spinal nerves, 536
Proper digital arteries, palm ar, lateral and Pulmo, 734 Radices spinales
medial, 339 Pulmo dexter et craniales o f accessory nerve, 512
Proper hepatic arteries, 345, 704 lobus apicalis, 735 of accessory nerve, 570
Proper ligament o f ovary, 780 lobus cardiacus, 738 Radices ventrales o f spinal nerves, 536
Proper vaginal tunic, 755 lobus diaphragm aticus, 738 R adioulnar joints, 110, 113
Propria linguae muscle, 154 lobus interm edius, 738 Radius, 64, 69-71
Prorean gyrus, 484 Pulmo sinister annular ligament, 110
Prorean sulcus, 482 lobus apicalis, 735 body, 71
Prosencephalon, 480 lobus cardiacus, 735 caudal surface, 71
Prostata, 762 lobus diaphragm aticus, 735 cranial surface, 71
Prostate gland, 750, 762-3 Pulmonary. See also Lungs, distal extremity, 71
clinical considerations, 763 alveoli, 728 head, 69
Prostatic artery, 379 arteries, 287-8, 739 lateral aspect, 211
Prostatic portion o f male urethra, 777 fibrous ring, 276 lateral border, 71
Prostatic utricle, 777 ligament, 734 left, articulated, 70
Prosthion of skull, 8 ostium, 278 ulnar surface, 70
Protractor preputii, 778 pleura, 733 medial aspect, 211
Protuberance, occipital, 12, 44 plexuses, 637 m edial border, 71
Protuberantia occipitalis externa, 12 trunk, 287-8, 738 neck, 71
Protuberantia occipitalis interna, 12 valve, 283 nutrient artery, 333
Proximal articular surface o f tibial tarsal veins, 288, 739 Radix dentis, 649
bone, 89 Pulp Radix dorsalis o f spinal nerve, 572
Proximal collateral radial artery, 328 o f spleen, 460 Radix linguae, 654
Proximal collateral ulnar vein, 403 o f tooth, 653 Radix mesenterii, 688
Proximal communicating vein, 401 Pulp cavity, 653 Radix m otoria of trigem inal nerve, 550
Proximal interphalangeal joints, 118 Pulpa dentis, 653 Radix penis, 763
Proximal jntertarsal joint, 127 Pulpa lienalis, 460 Radix pulmonis, 735
Proximal m uscular branch Pulvinar, 496 R adix sensoria o f trigem inal nerve, 550
of intercostal nerve, 595 Puncta lacrimalia, 838 R adix ventralis of spinal nerve, 572
of m usculocutaneous nerve, 583 Punctum fixum o f cranial attachm ent of Rami alveolares maxillares m edia o f infra
Proximal palm ar venous arch, 405 esophagus, 665 orbital artery, 312
Proximal plantar venous arch, 417 Pupil, 846 Ram i bronchiales
Proximal radioulnar joint, 113 Putamen, 489 o f bronchoesophageal artery, 339
Proximal tibiofibular jo in t, 127 Pyloric antrum , 681 o f internal thoracic artery, 321
Prussak’s pouch, 853 Pyloric canal, 681 R am i cecales of ileocecal artery, 350
Pseudocyesis, 789 Pyloric glands, 684 Ram i centrales o f m iddle cerebral artery,
Pseudopregnancy, 833 Pyloric p art o f stomach, 681 313
Psoas major muscle, 232 Pyloric sphincter, 683 R am i c o lla te ra ls of intercostal arteries, 345
Psoas minor muscle, 231 Pylorus, 679 R am i com m unicantes o f sym pathetic trunk,
action, 232 Pyramid 634
innervation, 232 cerebellar surface, 19 Ram i corticales o f m iddle cerebral artery,
Pterygoid bone, 34 cerebral surface, 19 313
medial aspect, 35 decussation, 508 Rami cutanei
sutures, 101 o f medulla oblongata, 507 o f circumflex scapular artery, 328
Pterygoid branch o f m axillary artery, 305 o f temporal bone, 19 o f coccygeal nerves, 623
Pterygoid canal, 18, 34, 42 tympanic surface, 19, 20 o f subscapular vein, 403
nerve, 558, 627 ventral surface, 20 R am i cutanei antebrachiales craniales, 586
Pterygoid groove, 18, 42 Rami cutanei dorsales
Pterygoid muscle, nerves to, 555 of intercostal arteries, 342
Pterygoid nerves, 555 o f lum bar nerves, 599
Pterygoid portion o f m axillary artery, 304 Q u a d r a t e lobe o f liver, 702 R am i cutanei laterales
Pterygoid processes, 19 Q uadratus femoris muscle, 236 o f intercostal arteries, 342
o f maxilla, 30 Q uadratus lum borum muscle, 232 of thoracic nerves, 594
Pterygoideus lateralis muscle, 152 Q uadratus plantae muscle, 263 Ram i cutanei ventrales of internal thoracic
Pterygoideus medialis muscle, 152 Quadriceps femoris muscle, 240 artery, 321
Pterygopalatine fossa, 30, 33, 39 action, 242 Ram i dorsales
Pterygopalatine ganglion, 627 innervation, 242 nn. cervicales III—V II, 575
lateral aspect, 553 Q uadrigeminal bodies, 497 o f intercostal arteries, 342
Pterygopalatine nerves, 554 of lum bar nerves, 595
Pterygopalatine portion o f maxillary arteiy, of sacral nerves, 610
304 R am i duodenales o f cranial pancreatico
Pterygopalatine suture, 33, 34 R a d ia lartery, 328, 329, 337 duodenal artery, 346
Pterygopharyngeus muscle, 156 o f humerus, collateral, 325 Rami epiploici
Pterygosphenoid suture, 19, 34 R adial carpal bone, 73 o f right gastroepiploic artery, 346
Pubic region, 672 Radial collateral ligaments, 110, 114 of splenic artery, 349
Pubic symphysis, 119 Radial fossa, 69 Ram i episclerales o f ciliary arteries, 305
Pubic tubercle, 82 R adial head o f m. flexor digitorum pro R am i esophagei
Pubis, 78, 82 fundus, 221 o f bronchoesophageal artery, 339
body, 82 R adial nerve, 583 o f left gastric artery, 349
cranial border, 80 o f forepaw, 587 o f recurrent laryngeal nerve, 570
ramus, 82 superficial, 587 Ram i gastrici of right gastroepiploic artery,
Pubovesical excavation, 674 Radial notch, 72 346
Pudendal arteries, 360, 379, 765 Radial tuberosity, 71 R am i glandulares of great auricular artery,
Pudendal nerve, 611 Radial vein, 403 297
Pudendal veins, 414, 765 Radiation, thalam ic, 495 Rami hyoidei of lingual artery, 296
928 In d ex
Ram us scrotalis caudalis o f perineal artery, Recurrent laryngeal nerve, 570, 630, 633 Ribs (Continued)
379 Red bone marrow, 4 ligaments, 106-108
Ram us scrotalis cranialis o f external puden Red nucleus, 500 dorsal aspect, 105
dal artery, 363 Red pulp o f spleen, 460 ventral aspect, 105
Ram us septalis o f left coronary artery, 285 Reflex vein, 394 neck, 61
Ramus sinister o f common hepatic artery, Reflexes, visceral, 642 right lateral aspect, 63
346 Regio abdominis, 667 true, 61
Ram us sinus carotici, 567 Regio epigastrica, 672 tubercle, 61
Ramus spinalis Regio hypochondriaca dextra et sinistra, 672 ventral aspect, 62
of cerebrospinal artery, 317 Regio inguinalis dextra et sinistra, 672 Rim
of intercostal arteries, 342 Regio insularis, 484 o f vestibule o f larynx, 724
of lum bar arteries, 355 Regio lateralis dextra et sinistra, 672 palpebral, 838
Ramus superficialis Regio lumbalis, 672 R im a glottidis, 724
of cranial tibial artery, 371 Regio olfactoria o f nasal cavity, 863 R im a oris, 645
of deep circumflex iliac artery, 359 Regio pubica, 672 R im a palpebrarum , 838
of deep circumflex iliac vein, 409 Regio respiratoria o f nasal cavity, 863 R im a vestibuli o f larynx, 724
of radial nerve, 583 Regio sacralis, 672 Ring
R am us ventralis Regio umbilicalis, 672 femoral, 249
n. cervicalis I, 574 Region, abdom inal. See Abdomen. fibrous, aortic, 276
n. cervicalis II, 575 Relaxin, 833 atrioventricular, 276
o f spinal nerve, 572 Renal. See also Kidneys. o f base of heart, 276
Ramus visceralis arteries, 351,746 o f intervertebral disc, 103
of intercostal nerve, 594 corpuscle, 743, 746 of vertebra, 51
of internal iliac artery, 378 hilus, 743 pulm onary, 276
of lumbar nerve, 599 impression, 699 inguinal, 761
o f spinal nerve, 572 papilla, 743 tympanic, 23
Ranvier, nodes of, 469 pelvis, 743 vaginal, 754, 761
Raphe palati, 649 plexus, 640 R oof plates of ethm oid, 25
Rathke, pouch of, 814 sinus, 743 Root
Reception, structures mediating, 471-9 tubules, 743-6 dorsal, o f spinal nerve, 572
Receptor neurons, 466,468 veins, 406, 747 dorsal aspect, 534, 535
Recess Renes, 741 of accessory nerve, 512
caudal, of omental bursa, 678 Renin, 834 of lung, 735
costodiaphragmatic, 733 R eproduction, endocrine biology of, 830 o f mesentery, 688
costomediastinal, 732, 733 Reproductive organs, 751-98 of spinal nerves, 471, 533-6
cranial, of omental bursa, 678 Research, endocrine, im portance o f dog in, of tongue, 654
epitympanic, 20, 23, 851 810 o f tooth, 649
maxillary, 30,49 Respiratory bronchioles, 728 ventral, of spinal nerve, 572
piriform, 724 Respiratory region o f nasal cavity, 863 R oot canal, 653
Recessus caudalis omentalis, 678, 679 Respiratory system, 713-40 R oot filaments of spinal nerves, 572
Recessus costodiaphragm atici, 733 Response, structures m ediating, 471-9 Rostral internal auricular nerve, 556
Recessus costomediastinalis, 732, 733 Resting potential, 466 R ostral maxillary alveolar nerve, 555
Recessus cranialis omentalis, 678 Rete, dorsal, o f carpus, 329 R ostral salivatory nucleus, 558
Recessus epitympanicus, 20, 23 Rete carpi dorsale, 329 R ostrum o f sphenoid, 16
Recessus lienalis o f om ental bursa, 679 Rete testis, 755 R ostrum sphenoidale, 16
Recessus maxillaris, 3D Rete venosum dorsale carpus, 405 Rotation, 99, 133
Recessus phrenicocostalis, 181 R eticular form ation, 492 of alim entary tract, 674, 675
Recessus phrenicolumbalis, 181 R eticular nucleus, 496 R otator nerve, 614
Recessus piriformis, 724 lateral, 523 R otatores breves muscles, 171
Rectal arteries, 351, 379 Retina, 843, 846 R otatores longi muscles, 171
Rectal nerve, caudal, 611 central artery, 305 Rotatores muscles, 171
Rectal vein, 407, 414 R etractor anguli oculi muscle, 140, 842 Rotators, 143, 144
Rectal venous plexus, 414 R etractor bulbi muscle, 148, 840 Round foramen, 18
Recti muscles, 146 Retractor costae muscle, 178 Round ligament. See Ligaments, round.
action, 148 R etractor penis muscle, 194, 697, 764 Round window, 20, 853, 857
innervation, 148 Retroglenoid foramen, 23,42 R ubrospinal tract, 501, 503, 523
Rectococcygeus muscle, 194, 697 Retroglenoid process, 23, 42 Rugae o f vagina, 790
Rectogenital pouch, 749 R etroglenoid vein, 398, 421 Rump muscles, 231, 232-5
Rectouterine excavation, 674 R etropharyngeal lym ph node, m edial, 435,
Rectouterine space, 787 438
Rectovesical excavation, 674 Rhinal fissure, 482
Rectovesical pouch, 749 Rhinal sulcus, 482 Sac
Rectum, 691, 693-4 Rhinal veins, 419 alveolar, 728
muscles, special, 695-7 Rhinencephalon, 490 anal. See A nal sacs.
Rectus, sheath, 182 Rhom bencephalon, 480 conjunctival, 838
Rectus abdominis muscle, 186 Rhom boideus capitis muscle, 200 lacrimal. See Lacrimal sac.
action, 188 Rhomboideus cervicis muscle, 200 pericardial, ventral aspect, 269
innervation, 188 Rhomboideus muscle, 200 Sacci anales, 694
sheath, 187, 188 Rhom boideus thoracis muscle, 200 Saccule, 856
Rectus capitis dorsalis intermedius muscle, Ribs, 61-4 laryngeal, 726
172 body, 61 Sacculi alveolares, 728
Rectus capitis dorsalis m ajor muscle, 172 cervical, 51 Sacculus laryngis, 726
Rectus capitis dorsalis m inor muscle, 172 crest, 61 Saccus conjunctivae, 838
Rectus capitis lateralis muscle, 175 false, 61 Saccus lacrimalis, 838
Rectus capitis ventralis muscle, 175 floating, 61 Saccus medialis et lateralis, 122
Rectus femoris muscle, 240 head, 61 Sacral artery, median, 386
Rectus muscles o f eye, 840 joints, 106-108 Sacral canal, 58
930 In d ex
Spinalis et semispinalis thoracis muscle, 170 Stifle jo in t (Continued) Sublingual caruncle, 645
Spine. See also Vertebral column. left, cruciate ligam ents, m edial aspect, 126 Sublingual duct, 660
iliac, 80, 81 ligaments, 124, 125 Sublingual fold, 645
ischiatic, 80, 81 dorsal aspect, 126 Sublingual ganglion, 556
nasal, See N asal spine. meniscal ligaments, m edial aspect, 126 Sublingual gland, 659-60
o f scapula, 64 menisci, dorsal aspect, 126 Sublingual nerve, 556
o f vertebral arch, 51 ligaments, 122-7 Sublingual papilla, 660
Spinocerebellar tract, 512, 523 right, arteries of, 620 Sublum bar fossa, 672
Spinothalam ic tract, 523 nerves of, 620 Sublum bar muscles, 231-2
Spinotransversalis fascia, 196 sesamoid bones, 85 ventral aspect, 233, 234
Spinotransverse fascia, 176 Stomach, 672, 679-85 Subm andibular fascia, superficial, 161
Spinous process blood vessels, 685 Subm andibular ganglion, 627
o f axis, 52 capacity, 681-3 Submental artery, 297
o f lum bar vertebrae, 56 coats, 683-4 Submucous coat
o f thoracic vertebrae, 54 curvatures, 679-81 of esophagus, 665
o f vertebral arch, 51 fundus, 681 o f large intestine, 698
Spiral lam ina, osseous, 857 glands, 684-5 o f small intestine, 688
Spiral organ o f Corti, 859 glandular cells, 684 of stomach, 683
Splanchnic nerves, 615, 638 longitudinal section, 680 Submucous plexus, 633
Spleen, 458-61, 672 lymph nodes and vessels, 451 Suboccipital nerve, 574
blood supply, 358 nerves, 685 Subpapillary plexus, 885
functions, 461 position, 681-3 Subscapular artery, 325
internal structure, 458, 460,461 regions. 681 Subscapular fossa, 66
lymph nodes and vessels, 451 shape. 681-3 Subscapular nerve, 582
Splenial gyrus, 484 walls, 679 Subscapular vein, 403
Splenial sulcus, 483 Stop, 39 Subscapularis muscle, 206
Splenic artery, 346 Straight sinus, 419 Subsplenial flexure o f dentate gyrus, 485
splenic branches, 349, 460 Stratum circulare Substance
Splenic branches o f splenic artery, 349, 460 of large intestine, 698 androgenic, 830
Splenic flexure, 692 o f m uscular coat of stomach, 683 intermediate, 536
Splenic lymph nodes, 447 Stratum corneum Nissl, 464
Splenic plexus, 640 o f claw, 887 perforated, anterior, 490
Splenic portion o f greater om entum , 675 o f foot pad, 876 Substantia alba o f spinal cord, 536-9
Splenic recess o f om ental bursa, 679 o f nasal skin, 876 Substantia compacta, 4
Splenic vein, 409 Stratum cylindricum, 876 Substantia corticalis, 4
Splenium corporis callosi, 488 Stratum germ inativum , 876 Substantia gelatinosa, 536
Splenium o f corpus callosum, 488 o f claw, 887 Substantia grisea centralis, 497
Splenius muscle, 162,164 Stratum granulosum o f foot pad, 876 Substantia grisea o f spinal cord, 536
action, 164 Stratum longitudinale Substantia interm edia centralis, 536
innervation, 164 o f large intestine, 698 Substantia interm edia lateralis, 536
Squam a frontalis, 16 o f m uscular coat o f stom ach, 683 Substantia nigra, 500
Squamosum, 23 Stratum lucidum of foot pad, 876 Substantia perforata anterior, 490
Squamous suture, 14, 23, 95 Stratum spinosum, 876 Substantia spongiosa, 4
Stalk Stratum vasculare of uterus, 787 Subsynaptic m em brane, 475
hypophyseal, 810 Stria malleolaris, 851 Subthalamic nucleus, 497
o f epiglottis, 719 Stria m edullaris thalam i, 494 Subthalamus, 497
Stapedial muscle, nerve to, 559 Stria olfactoria lateralis et medialis, 490 Sulci, 482
Stapedius muscle, 144, 856 Stria terminalis, 490 o f cerebrum, left dorsolateral view, 494
fossa for, 22 Striae, olfactory, 490 Sulcus
Stapes, 853, 856 Stroma ansate, 483
Stellate cardiac nerves, 636 corneal, 843 anthelicine, 848
Stellate ganglion, 635 o f m am m ary gland, 801 calcanean, 89
Stellate veins o f kidneys, 747 Styloglossus muscle, 154 callosal, 483
Sternal branches o f in tern al thoracic artery, Stylohyoid bone, 38 callosomarginal, 483
321 Stylohyoid nerve, 559 cingulate, 483
Sternal lymph node, 440 Stylohyoideum s. pars stylohyoidea, 38 coronal, 483
Sternal m em brane, 108 Stylohyoideus muscle, 146 cruciate, 483
Sternebrae, 64 Styloid process, 71, 72 dorsal, 28
Sternocephalicus muscle, 200 o f antitragus, 848 dorsointermediate, 533
Sternocostal joints, 108 Stylomastoid artery, 297 dorsolateral, o f spinal cord, 533
Sternocostal radiate ligaments, 108 Stylomastoid foramen, 22, 44 ectogenual, 483
Sternocostal surface o f heart, 274 Stylopharyngeus muscle, 156 ectolateral, 483
Sternohyoideus m uscle, 148, 173 Subarachnoid cavity, 540, 541 ectomarginal, 483
action, 150 Subarachnoid cisterns, 540 ectosplenial, 483
innervation, 150 Subcallosal area, 485, 492 ectosylvian, 483
Sternom astoideus muscle, 200 Subcallosal bundle, 486 entolateral, 483
Sterno-occipitalis muscle, 200 Subcallosal gyrus, 485 for middle meningeal artery, 14, 18
Sternopericardiac ligament, 267 Subclavian artery, 316-24 for transverse sinus, 12
Sternothyroideus muscle, 173 Subcortical fibers, 486 genual, 483
Sternum , 64 Subcostal artery, 345 hippocampal, 483
right lateral aspect, 63 Subcostal nerve, 594 lateral, 483
ventral aspect, 62 Subcostal veins, 399 median, dorsal, 523, 533
STH. See Somatotrophic hormone. Subcutaneous plexus, 885 occipitotemporal, 483
Stifle joint. 78, 122-7 Subdural cavity, 541 olfactory, 482
fascia, 265 Sublingual artery, 297 palatine, 30
left, capsule, 124,125 medial aspect, 299 postcruciate, 483
In d ex 933
Tract (Continued) Triceps brachii muscle (Continued) Tuba auditiva ossea, 19, 23
m am m illothalam ic, 496 innervation, 210 Tubae uterinae, 784
mesencephalic, 501 Tricuspid area, 282 Tube
olfactory, 490 Tricuspid valve, 282 auditory. See A uditory tube.
olivocerebellar, 512 Trigeminal ganglion, semilunar, 550 eustachian, 719, 853
olivospinal, 523 Trigeminal lemniscus, 496, 501, 503, 523 fallopian, 784
rubrospinal, 501, 503, 523 Trigeminal nerve, 503, 550-58, 649 Tuber, omental, 702
spinal, See Spinal tract. canal for, 20,45 Tuber calcanei, 89
spinocerebellar, 512, 523 dorsal aspect, 548 Tuber cinereum, 497
spinothalam ic, 523 lateral aspect, 551, 560 T uber coxae, 81
tectobulbar, 503 m axillary branch, lateral aspect, 553 T uber ischiadicum, 81
tectonuclear, 501 nucleus of, 501, 511 Tuber maxillae, 30
tectospinal, 501, 503, 523 spinal tract, 503 Tuber omentale, 702
uveal, 844 Trigone Tuber sacrale, 80
vestibulospinal. See Vestibulospinal tract. fibrous, 276 T uber scapulae, 66
Tractus cerebellorubralis, 507 olfactory, 490 Tuberal portion of hypothalam us, 496
Tractus cerebellothalam icus, 507 vesical, 749 Tubercle
Tractus corticonuclearis, 500 Trigonum femorale, 249, 363 anterior, of thalamus, 496
Tractus corticopontinus, 500 Trigonum fibrosum dextrum , 276 corniculate, 724
Tractus corticospinales ventrales, 508 Trigonum fibrosum sinistrum, 276 cuneiform, 724
Tractus corticospinalis, 500 Trigonum lum bocostale, 180 dorsal and ventral, of atlas, 51
Tractus corticospinalis lateralis, 508 Trigonum olfactorium, 490 greater, 67
Tractus fastigiobulbaris, 507 Trigonum vesicae, 749 intercondyloid, 87
Tractus m am m illothalam icus, 496 Trochanter intervenous, 274
T ractus m esencephalicus nervi trigemini, greater, 82 lesser, 67
501 lesser, 84 muscular, 13,42
Tractus olfactorius, 490 third, 84 nuchal, 13
Tractus olivocerebellaris, 512 T rochanter m ajor, 82 of dentate gyrus, 485
Tractus rubrospinalis, 501 T rochanter m inor, 84 of rib, 61
Tractus solitarius, 512 T rochanter tertius, 84 ligament of, 106
Tractus spinalis nervi trigemini, 503 Trochanteric fossa, 84 o f teeth, 649
Tractus spinocerebellaris dorsalis, 512 Trochanteric surface o f femur, 84 olfactory, 490
Tractus spinocerebellaris ventralis, 523 Trochlea, 146 pharyngeal, 13
Tractus spiralis foram inosus, 20 femoral, 84 pubic, 82
Tractus tectonuclearis, 501 proximal, 89 urethral, 791
Tractus tectospinalis, 501 Trochlea femoris, 84 Tuberculae dentis, 649
Tractus vestibulospinalis lateralis, 511 Trochlea humeri, 69 Tuberculae musculares, 13
Tractus vestibulospinalis ventralis, 511 Trochlea tali proximalis, 89 Tuberculae nuchales, 13
Tragicus lateralis muscle, 144 Trochlear nerve, 547-50 Tuberculum anterius thalami, 496
Tragohelicinus muscle, 142 dorsal aspect, 548 Tuberculum corniculatum, 724
Tragotubohelicinus muscle, 142 nucleus of, 501 Tuberculum costae, 61
Tragus, 848 Trochlear notch, 72 Tuberculum cuneiforme, 724
Transversalis fascia, 670 Trochlearis, 146 Tuberculum dorsale et ventrale, 51
Transverse artery o f neck, 317 T rochoidjoint, 98 Tuberculum gyri dentati, 485
Transverse canal, 12,45 Truncus bicaroticus, 289 Tuberculum infraglenoidale, 66
Transverse cervical nerve, 575 Truncus brachiocephalicus, 289 Tuberculum intercondylare mediale et lat
Transverse colon, 692 Truncus celiacus, 345 erale, 87
Transverse crest, 19 Truncus coccygei dorsalis, 622 Tuberculum intervenosum, 274
Transverse diam eter o f pelvis, 80 Truncus coccygeus ventralis, 623 Tuberculum majus, 67
Transverse facial artery, 300 Truncus corporis callosi, 488 Tuberculum minus, 67
Transverse fibers o f pons, 503 Truncus costocervicalis, 317 Tuberculum olfactorium, 490
Transverse foramen, 51, 52 Truncus lumbosacralis, 610 Tuberculum pharyngeum, 13
Transverse groove, 45 Truncus pudendoepigastricus, 360 Tuberculum pubicum, 82
T ransverse ligam ents. See Ligaments, trans Truncus pulmonalis, 287, 738 Tuberculum sellae, 18,45
verse. T ru n cu s venosus p ro fu n d u s penis et bulbi, Tuberculum supraglenoidale, 66
Transverse lines, 56, 84 414 Tuberositas deltoides, 67
Transverse mesocolon, 693 T runcus venosus pudendoepigastricus, 414 Tuberositas iliaca, 81
Transverse m etatarsal artery, 371 Trunk Tuberositas ossis tarsalis quarti, 92
Transverse m etatarsal vein, 417 arteries, superficial, 348 Tuberositas plantaris, 89
Transverse processes. See Processes, trans bicarotid, 289 Tuberositas radii, 71
verse. brachiocephalic, 289-316 Tuberositas supracondylaris medialis et lat
Transverse sinus. See Sinuses, transverse. celiac, 345 eralis, 85
Transverse sulcus, 14 coccygeal, 622, 623 Tuberositas teres, 67
Transversospinalis system, 164, 168-73 costocervical, 317 Tuberositas tibiae, 87
Transversus abdom inis muscle, 183 esophageal, 623 Tuberositas ulnae, 72
action, 186 fasciae, 194-7 Tuberosity
innervation, 186 lumbosacral, 610 deltoid, 67
Transversus costarum muscle, 178 muscles, 162-89 iliac, 81
Transversus thoracis muscle, 178 deep, ventral aspect, 185 infraglenoid, 66
Trapezius cervicis m uscle, 197 superficial, ventral aspect, 184 ischiatic, 81
Trapezius muscle, 197 pudendoepigastric, 414 maxillary, 30
Trapezius thoracis muscle, 197 pulm onary, 287-8, 738 of fourth tarsal bone, 92
Trapezoid body, 508 sacral, dorsal, 610 radial, 71
Triangle, femoral. See Femoral triangle. vagal, 632 scapular, 66, 67
Triangular ligament, 703, 704 venous, costocervical-vertebral, 389 supracondylar, 85
Triceps brachii muscle, 209 TSH. See Thyrotrophic hormone. supraglenoid, 66, 67
action, 210 Tuba auditiva, 719, 853 teres, 67
In d ex 937
Tuberosity (Continued) Tym panohyoideum s. pars tym panohyoidea, U rinary bladder (Continued)
tibial, 87 38 clinical considerations, 751
ulnar, 72 Tym pano-occipital jo in t, 13, 23 fixation, 7 4 9
Tubular glands o f external auditory meatus, Tyrosine-m elanin, 882 ligaments, 7 4 9
851 nerves, 7 4 9 -5 1
Tubules structure, 7 4 8 - 9
renal. 7 4 3 -6 U rinary organs, 7 4 1 -5 1
seminiferous, 755 U l n a , 64, 7 1 - 3 lymph vessels, 457
straight, of testes, 755 body, 72 Urocyst, 74 8
Tubuli renales, 7 4 3 -6 caudal surface, 72 Urogenital apparatus, 741
Tubuli renales contorti, 746 cranial surface, 72 Urogenital artery, 37 8
Tubuli seminiferi, 755 distal extremity, 72 Urogenital ligam ents of male, ventral as
Tubulus renalis rectus, 743 lateral aspect, 211 pect, 759
Tumors of m ammary glands, 803 lateral border, 72 Urogenital system, 741
Tunic left, articulated, 70 and m am m ary glands, 7 4 1 - 8 0 6
fibrous, of spleen, 4 6 0 radial surface, 70 embryology, 7 9 6 - 8
vaginal. See Vaginal tunic. medial aspect, 211 female, lateral aspect, 781
Tunica adventitia o f esophagus, 664 medial border, 72 ventral aspect, 742
Tunica albuginea, 755 nutrient artery, 333 U rogenital vein, 4 1 4
of ovary, 780 proxim al extremity, 72 U terine artery, 3 7 8 , 3 7 9
Tunica conjunctiva, 837 U lnar artery, 333 Uterine milk, 833
Tunica conjunctiva bulbi, 838 collateral, 32 8 U terine ostium, 7 8 4
Tunica conjunctiva palpebrarum , 83 8 recurrent, 3 2 9 U terine vein, 4 1 6
Tunica externa o f eye, 843 U lnar carpal bone, 73 U tero-ovarian artery, 3 5 5
Tunica fibrosa U lnar collateral ligament, 110, 114 U tero-ovarian plexus, 64 0
of eye, 843 U lnar head Uterus, 6 7 2 , 7 8 6 - 9
of spleen, 460 o f m. flexor carpi ulnaris, 220 anomalies and variations, 7 8 9
Tunica interna o f eye, 843 o f m. flexor digitorum profundus, 221 blood vessels, 7 8 8 - 9
Tunica intima, 276 U lnar nerve, 58 6 body, 7 8 6
Tunica media o f eye, 843 o f forepaw, 5 90 broad ligament, 67 4
Tunica mucosa, 645 U lnar notch, 71 cervix, 786
o f esophagus, 665 U lnar tuberosity, 7 2 clinical considerations, 78 9
of large intestine, 697 U lnar vein, 40 3 horns, 78 6
of oviduct, 786 Umbilical aperture, 6 7 0 ligaments, 78 7
of small intestine, 688 Umbilical artery, 3 7 8 nerves, 7 8 8 - 9
of stomach, 684 Umbilical fold, middle, 6 7 4 relations, 7 8 7
of ureter, 748 Umbilical ligaments, 3 7 8 , 7 4 9 structure, 7 8 7 - 8
of uterus, 788 Umbilical region, 67 2 tunics, 78 7 , 7 8 8
o f vagina, 790 Umbilicus, 189 Uterus didelphys, 7 8 9
Tunica m ucosa linguae, 654 Umbo m em branae tym pani, 851 U terus m asculinus, 7 7 7
Tunica muscularis U ncinate bundle, 4 8 6 U terus unicornis, 7 8 9
of esophagus, 665 Uncinate notch, 25 Utricle
of large intestine, 6 9 8 Uncinate process o f ethm oid, 25 o f ear, 856
of oviduct, 784 Ungual crest, 75 prostatic, 7 7 7
of small intestine, 688 U nipennate muscle. 132 U triculus prostaticus, 77 7
of stomach, 683 U pper dental arch, 651 Uvea, 844
of ureter, 748 U pper lips, 64 6 Uveal tract, 84 4
of uterus, 787 U reteral artery, caudal, 37 8 Uvula, muscle, 156
of vagina, 790 U reteral mucosa, 7 4 8
Tunica nervea oculi, 843, 846 U reteral vein, caudal, 4 1 6
Tunica serosa Ureteric vein, cranial, 4 0 6
of large intestine, 698 Ureters, 7 4 7 - 8 Va g a l trunk, 63 2
of liver, 703 anomalies, 7 4 8 Vagina, 7 9 0 -9 1
of oviduct, 784 blood vessels, 74 8 vestibule, 791
of small intestine, 688 clinical considerations, 74 8 Vaginal artery, 37 9
of stomach, 683 nerves, 74 8 Vaginal cavity, 75 4
of uterus, 787 structure, 7 4 8 Vaginal process, 7 7 9
peritoneal, 672 U rethra, 751 o f male, sagittal section, 759
Tunica vaginalis communis, 754 bulb, 763 Vaginal ring, 7 5 4 , 761
Tunica vaginalis parietalis, 7 5 4 , 761 female, 7 9 4 - 6 Vaginal tunic, 7 5 4 , 7 5 5 , 761
Tunica vaginalis propria, 7 5 4 , 755 anomalies and variations, 7 96 Vaginal vein, 4 1 6
Tunica vaginalis visceralis, 7 5 4 , 755 clinical considerations, 7 9 6 Vaginal venous plexus, 4 1 6
Tunica vasculosa oculi, 844 male, 7 7 7 - 8 Vagus nerve, 51 1 , 5 1 2 , 5 6 7 - 7 0 , 6 3 0 - 3 3 , 874
Tunica vasculosa o f eye, 843 clinical considerations, 7 78 Vallate papillae, 86 8 , 8 7 1 , 872
Turbinates, 717 U rethra feminina, 79 4 Vallecula, 71 9
nasal, 2 7 , 3 1 ,8 6 3 U rethra masculina, 7 77 Valva aortae, 28 2
Tympanic artery, 303 U rethral branches Valva atrioventricularis dextra, 2 82
Tympanic bullae, 42 o f prostatic artery, 3 7 9 Valva atrioventricularis sinistra, 28 2
Tympanic cavity, 20. 2 3 , 851 o f vaginal artery, 3 7 9 Valva mitralis, 2 8 2
Tympanic incisure, 851 U rethral crest, 777 Valva trunci pulm onalis, 283
Tympanic membrane, 2 3 , 848, 851 U rethral groove, 93 Valvae atrioventriculares, 2 8 2
Tympanic nerve, 563, 627, 853 U rethral tubercle, 791 Valve
Tympanic plexus, 563, 627 U rethral venous plexus, 4 1 4 aortic, 2 8 2
Tympanic ring, 23 Urinary bladder, 6 7 2 , 7 4 8 -5 1 atrioventricular, 28 2
Tympanicum, 22 anomalies, 751 mitral, 282
Tym panohyoid cartilage, 38 blood vessels, 7 4 9 -5 1 o f foramen ovale, 2 7 6
938 In d ex
Vena cordis media, 287 Vena pudenda interna, 414 V enae m etatarseae dorsales profundae, 417
Vena cordis parva, 287 Vena radialis, 403 Venae m etatarsea plantares profundae, 417
Vena corporis callosi, 422 Vena rectalis caudalis, 414 Venae m etatarseae plantares superficiales,
Vena costocervicalis, 390 Vena rectalis cranialis, 407 417
Vena diploica frontalis, 393, 424 Vena reflexa, 394 Venae pancreaticae, 409
Vena diploica occipitalis, 424 Vena renis, 747 Venae phrenicae, 407
Vena diploica parietalis, 424 Vena retroglenoidalis, 398, 421 Venae phrenicoabdom inales, 406
Vena dorsalis penis, 414 Vena sacralis media, 416 Venae pulm onales, 288, 739
Vena dorsalis ventriculi sinistri, 287 Vena saphena lateralis, 409 Venae pulm onales dextrae, 288
Vena ductus deferens, 414 Vena saphena medialis, 409 Venae pulm onales sinistrae, 288
Vena emissaria occipitalis, 419 Vena sublingualis, 394 Venae renales, 406
Vena ethmoidalis externa, 393 Vena submentalis, 394, 398 Venae rhinales, 419
Vena facialis, 393 Vena subscapularis, 403 Venae spinales, 426
Vena faciei profunda, 394 Vena tarsea lateralis, 417 Venae subcostales, 399
Vena femoralis, 413 Vena tarsea medialis, 417 Venae suprarenales, 406
Vena femoralis caudalis, 413 Vena tem poralis profunda, 398 Venereal sarcoma, 791
Vena femoralis cranialis, 413 Vena tem poralis superficialis, 398 Venous arch
Vena femoralis profunda, 414 Vena testicularis dextra, 406 digital, 416
Vena frontalis, 393 Vena testicularis sinistra, 406 hyoid, 394
Vena gastrica dextra, 409 Vena thalam ostriata, 422 palm ar, 403, 405
Vena gastrica sinistra, 409 Vena thoracica interna, 390 plantar, 417
Vena gastroduodenalis, 409 Vena thoracodorsalis, 403 Venous palatine plexus, 398
Vena gastroepiploica dextra, 409 Vena thyroidea caudalis, 393 Venous pathw ays in bulbus glandis, 775
Vena gastroepiploica sinistra, 409 Vena thyroidea cranialis, 390 Venous plexus
Vena gastrosplenica, 409 Vena thyroidea ima, 390 rectal, 414
Vena glutea caudalis, 414 Vena tibialis caudalis, 413 urethral, 414
Vena glutea cranialis, 416 Vena tibialis cranialis, 413 vaginal, 416
Vena hemiazygos, 399 Vena transversa facei, 398 vertebral, 428
Vena ileocecocolica, 407 Vena truncus costocervicalis-vertebralis, Venous rete, dorsal, o f carpus, 405
Vena iliaca communis, 414, 416 389 Venous sinuses
Vena iliaca externa, 413 Vena ulnaris, 403 cranial, dorsal aspect, 423
Vena iliaca interna, 414 Vena ureterica caudalis, 414, 416 lateral aspect, 420
Vena iliolumbalis, 416 Vena ureterica cranialis dextra, 406 of cranial dura m ater, 419-22
Vena infraorbitalis, 393 Vena ureterica cranialis sinistra, 406 o f spleen, 460
Vena interossea communis, 403 Vena urethralis, 414 vertebral, 426
Vena jugularis externa, 393 Vena urogenitalis, 414 Venous system, 389-429
Vena jugularis interna, 390 Vena uterina, 416 Venous trunk, costocervical-vertebral, 389
Vena labialis dorsalis, 394 Vena vaginae, 416 Ventral atlanto-occipital m em brane, 101
Vena labialis ventralis, 394 Vena vertebralis, 390 V entral branches
Vena laryngea cranialis, 398 Vena vesicae caudalis, 414, 416 o f cervical nerves, 575
Vena laryngea impar, 398 Venae arcuatae, 426 o f first cervical nerve, 574
Vena lienalis, 409 o f kidneys, 747 o f lum bar nerves, 599
Vena lingualis, 394 Venae basivertebrales, 426 o f sacral nerves, 610
Vena malaris, 394 Venae bronchoesophageae, 399 of second cervical nerve, 575
Vena mandibularis, 393 Venae centrales o f liver, 704 o f spinal nerve, 572
Vena masseterica, 398 Venae cerebelli dorsales, 424 o f thoracic nerves, 594
Vena maxillaris externa, 393 Venae cerebelli ventrales, 424 Ventral buccal nerve, 563
Vena maxillaris interna, 398 Venae cerebri, 422 Ventral cardiac veins, 287
Vena m ediana, 403 Venae cerebri dorsales, 422 Ventral cerebellar veins, 424
Vena mediana cubiti, 401 Venae cerebri internae, 422 Ventral coccygeal artery, 387
Vena meningea media, 398 Venae comitantes, 389 Ventral coccygeal trunk, 622, 623
Vena mesenterica caudalis, 407 Venae cordis, 287 Ventral cochlear nucleus, 508
Vena mesenterica cranialis, 407 Venae cordis minimae, 287 Ventral columns, 536
Vena m etatarsea dorsalis superficialis, 416, Venae cordis ventrales, 287 Ventral condyloid fossa, 12
'417 Venae digitales com m unes palm ares, 405 Ventral corticospinal tract, 508
Vena m etatarsea dorsalis superficialis proxi- Venae digitales dorsales pedis, 416 Ventral cutaneous branch o f intercostal
malis, 417 Venae digitales palm ares, 405 nerve, 595
Vena m etatarsea p lantaris superficialis Venae digitales plantares, 417 Ventral esophageal trunk, 632
proximalis, 417 Venae digiti dorsales, 403 Ventral external arcuate fibers, 511
Vena m etatarsea transversa, 417 Venae diploicae, 424 Ventral external vertebral venous plexus,
Vena nasalis dorsalis, 393 Venae esophageae, 399 428
Vena nasalis lateralis, 393 Venae hepaticae, 407, 704 Ventral extrem ity o f spleen, 458
Vena obliqua atrii sinistri, 287 Venae ilei, 407 Ventral funiculi, 536, 539
Vena occipitalis, 390 Venae intercostales dorsales, 399 Ventral gray horn, 536
Vena omocervicalis, 393 Venae interlobares o f kidneys, 747 Ventral interoccipital sinus, 422
Vena ophthalmica, 393 Venae interlobulares Ventral intraoccipital synchondroses, 13
Vena ovarica dextra, 406 o f kidneys, 747 Ventral labial artery, 297
Vena ovarica sinistra, 406 o f liver, 704 Ventral labial vein, 394
Vena pancreaticoduodenalis caudalis, 407, Venae intervertebrales, 426 Ventral margin o f lung, 735
409 Venae jejunales, 407 Ventral m edian fissure, 507, 533
Vena pancreaticoduodenalis cranialis, 409 Venae lumbales, 399, 406 Ventral m ediastinal cavity, 732
Vena perinealis, 414 Venae meningeae, 424 Ventral m ediastinal pleura, 732
Vena pharyngea, 398 Venae m etacarpeae dorsales profundae, Ventral muscles of vertebral column, 174
Vena poplitea, 413 403 Ventral m uscular branch o f external eth
Vena portae, 407, 704 Venae m etacarpeae dorsales superficiales, moidal artery, 304
Vena profunda brachii, 403 403 Ventral nasal concha, 27, 863
Vena prostatica, 414 Venae m etacarpeae palm ares, 405 Ventral nasal meatus, 718, 866
940 In d ex