Animal Structure & Function

The digestive system | What is the rumen? | The

ruminant digestive system | Ruminant
ecophysiology | Mycotoxins and rumen function |
Monogastric digestion | Hindgut fermenters | Adaptations
to high-fibre diets | Human digestive system| References

The digestive system

The digestive system is one of the body's major organ
systems. All animals - with the exception of some
endoparasites such as tapeworms - have a digestive
system. In this section we're going to look at digestion in
ruminants, and compare their guts with those of other,
non-ruminant, mammals.
return to top

What is the rumen?

The rumen underpins much of our agricultural industry.

Without this stomach chamber, cows and other ruminants
would be much less efficient at turning grass into milk,
meat and wool. A cow's rumen has a capacity of up to 95
litres and contains billions of bacteria and other microbes.
These microbes produce the enzymes that digest
cellulose into sugars and fatty acids for their hosts to use.
A less desirable by-product is the potent greenhouse gas,
methane: a single cow can produce up to 280 litres of
methane a day.

The rumen is one of four stomach compartments found in

ruminants. Ruminants are animals such as cattle, sheep,
goats and deer. (In comparison, animals such as pigs,
dogs and horses have only a single stomach
compartment and are called nonruminants,
ormonogastric animals.) The rumen allows grazing
animals to digest cellulose, a very common carbohydrate
in plants.
 Cow's digestive tract, viewed from left side. Image
from Nickel et al. 1973

Three of the four ruminant stomach compartments make

up the forestomach. These three compartments the
rumen, reticulum, andomasum are an extension of the
lower oesophagus. The rumen, the first of the
forestomach chambers, stores and processes plant
material. It can be a very large structure indeed: in large
ruminants, the rumen may store up to 95 litres of
undigested food (Brooker et al. 2008). The rumen holds
plant material until it has been broken down,
releasing volatile fatty acids,
and fermentation of protein and carbohydrates has
begun.

Relative size of ruminant stomach chambers. Image

 from Nickel et al. 1973

Why is the rumen so big? Because most plants,

especially grasses, have a high cellulose content. A
cellulose molecule is a polymer, made up of a long chain
of subunits called simple sugars, or
monosaccharides.Vertebrate animals lack the enzyme,
called cellulase, needed to break down cellulose and
release these sugars. Instead, they use symbiotic
anaerobic bacteria that do possess the enzyme
cellulase. Huge numbers of these bacteria are present in
the rumen and the reticulum: with each gram of rumen
fluid contains 10 - 50 billion bacteria.

The ruminant digestive system is a very efficient

adaptation for extracting as much energy as possible
from a high cellulose diet. Because food is held in a
ruminants gut for a relatively long time, symbiotic
bacteria are able to grow and release nutrient sources
that are not otherwise available to vertebrates.

How does ruminant digestion work?

Ruminant digestion begins when a cow swallows a
mouthful of plants. The food is partially chewed and
mixed into a bolus with saliva, before being swallowed
and passing down the oesophagus into the rumen. When
ruminants are grazing they tend to swallow their food
quickly, with only minimal mastication. When the animal is
resting after grazing, it regurgitates this partially chewed
food, rechews it, and then swallows it again (This process
is know as chewing the cud, or rumination.) Depending
on the amount of fibre in their food, cattle may spend
between 3 6 hours per day chewing their cud
(Lofgreen et al 1957).

return to top

The ruminant digestive system

Oesophagus | Reticulorumen | Omasum |Abomasum |

Small intestine |Duodenum | Jejunum | Ileum | Caecum |
Large intestine

The ruminant digestive tract (University of Minnesota,

1996)

The Oesophagus
The oesophagus is a muscular tube that connects the
mouth with the forestomach. Food passes down the
oesophagus by contraction of the muscles in the walls
that push the food along in a series of waves called
peristalsis. The ruminant oesophagus is also capable of
reversed peristalsis or antiperistalsis. This allows food to
be easily regurgitated from the rumen and chewed.

The reticulorumen
The reticulorumen is composed of the rumen and the
reticulum. The reticulorumen is partially separated from
the rumen by the reticular fold, which allows mixing
between the two compartments. The contents of the
reticulorumen are mixed by contractions of the
reticulorumen wall. The mixing recirculates undigested
material preventing the rumen becoming clogged and
distributing symbiotic bacteria throughout the ingested
material. The reticulorumen becomes colonized by
symbiotic bacteria in the first week after birth. The
bacteria help to break down the food and release
nutrients by a fermentation process.

The omasum
When food has been broken down enough, it passes
from the reticulorumen through the reticulo-omasal
orifice to the omasum. The omasum wall is highly folded,
giving a large surface area which allows for the efficient
absorption of water and salts released from the partially
digested food. The omasum also acts as a type of pump,
moving the food from the reticulorumen to the true
stomach, the abomasum, where acid digestion takes
place.

Section through cow's stomach, viewed from the front.

Note the highly folded wall of the omasum. Image
from Nickel et al. 1973.

The abomasum
Unlike a ruminant's three forestomachs, the abomasum is
a 'secretory stomach'. This means that cells in the
abomasum wall produce enzymes and hydrochloric
acid which hydrolyse proteins in the food and also in the
microbes mixed in with the food. Hydrolysis breaks the
proteins into smaller sub-units (e.g. dipeptides and amino
acids), ready for further digestion and absorption in the
small intestine.
Because ruminants eat such large amounts of plant
material, there is an almost continuous flow of food
through the abomasum. In comparison, activity in the
stomach of monogastric animals generally has
a circadian rhythm associated with food intake (Djikstra,
2005).

Cow's stomach viewed from the right-hand side. Image

from Nickel et al. 1973

The small intestine

The small intestine is an elongated tube running from the
abomasum to the large intestine. In ruminants, the small
intestine is about 20 times longer than the length of the
animal - so a cow two metres in length would have a
small intestine 40 metres long!

A large proportion of the digestion and absorption of

nutrients and water occurs in the small intestine.
Enzymes in the small intestine break nutrient molecules
down into their building blocks. Carbohydrates are broken
down to simple sugars (monosaccharides), fats into fatty
acids andmonoglycerides, nucleic acids
into nucleotides and proteins into amino acids. Some of
these enzymes are on the surfaces of intestinal cells,
while others are secreted into the small intestine,
primarily from the liver and pancreas.
The small intestine has three regions: the duodenum, the
jejunum and the ileum. Partially digested food passes
from the duodenum along the small intestine by way of
peristaltic muscle contractions that start at the part where
the abomasum is joined to the duodenum.

Duodenum
The liver and pancreas both secrete materials
through ducts into the duodenum. The common bile duct
carries bile salts, a greenish fluid that is manufactured in
the liver, stored in the gall bladder (the ruminant gall
bladder does very little to concentrate the bile), and
released into the duodenum to digest fats. The main
pancreatic duct carries digestive secretions, which are
rich in enzymes and bicarbonate. The bicarbonate
neutralises acid from the stomach, which would otherwise
inactivate many of the duodenum's digestive enzymes.

Jejunum
The lining of the jejunum is specialised for the absorption
of carbohydrates and proteins. Its inner surface is
covered in finger-like projections called villi, which
increase the surface area available to absorb nutrients
from the gut contents. The villi in the jejunum are much
longer than in the duodenum or ileum. The epithelial cells
which line these villi possess even larger numbers of
microvilli, known collectively as the brush border. The
combination of villi and microvilli increases the surface
area of the small intestine, increasing the chance of a
food particle encountering a digestive enzyme and being
absorbed across the epithelium and into the blood
stream.

Nutrients can cross the intestine wall by either passive or

active transport. In passive transport molecules diffuse
into the intestinal cells down a concentration gradient (i.e.
they move from a region where they are in high
concentration to an area of low concentration.) The sugar
xylose enters the blood by passive transport. Active
transport requires energy. Amino acides, small peptides,
vitamins, and most glucose are moved across the
intestine lining by active transport. Once nutrients have
moved through the epithelial cells, they are taken up by
either capillaries or lacteals and then transported around
the body.

Ileum
The ileum's main function is absorption of vitamin B 12, bile
salts and whatever nutrients that were not absorbed by
the jejunum. At the point where the ileum joins the large
intestine there is a valve, called the ileocaecal valve,
which prevents materials flowing back into the small
intestine.

The caecum
The caecum is a pouch connected to the large intestine
and the ileum. It is separated from the ileum by the
ileocaecal valve, and is considered to be the beginning of
the large intestine. In herbivores the caecum is greatly
enlarged and serves as a storage organ that permits
bacteria and other microbes time to further digest
cellulose. Partially digested food enters the caecum
through the ileoacecal valve, which is normally closed.
The valve occasionally opens to allow food material in. As
there is only one opening to the caecum, digesta must
move in and out to the caecum through the same
opening.

The large intestine

In addition to the caecum the large intestine is made up
of the ascending colon, transverse colon, sigmoid colon,
rectum, and anus. Much of the large intestine comprises
the colon, which is shorter in length but larger in diameter
than the small intestine. The colon is involved in the
active transport of sodium, and absorption of water by
osmosis, from the digested material that it contains. It
also provides an environment for bacteria to grow and
reproduce. These symbiotic bacteria produce important
vitamins such as vitamin K, thiamine, and riboflavin,
required by the animal for proper growth and health.
Finally, the large intestine eliminates wastes. Undigested
and unabsorbed food, as well as other body wastes,
leave the intestine in the form of faeces, via the rectum &
anus.

return to top

Mycotoxins and rumen function

It's important for the rumen to function effectively as this

results in maximal digestion and absorption of nutrients
from the animal's feed, and this in turn maximises
production of milk, meat and wool.

Grass is often infected with a range of fungi

(called endophytesbecause they live within the plant's
tissues). Some of these fungi produce a range of toxic
chemicals, or mycotoxins, which can affect the animals
that eat the infected grass. Some of these chemicals
protect plants from insect attack and have little effect on
ruminants, while others have marked effects on
ruminants and other herbivores. A research group based
at the University of Waikato has studied the impact of
mycotoxins on rumen function.

The team looked specifically at the mycotoxins that

result in muscle tremors - such toxins are called
'tremorgens'. Tremorgens result in the condition called
'ryegrass staggers', because the increased excitability of
skeletal muscle causes animals to stagger when they
move. These toxins can both excite and inhibit smooth
muscle contractions. Because the wall of the rumen
contains smooth muscle tissue, tremorgens have the
potential to upset normal rumen function.

The rumen and the reticulum normally undergo regular

rhythmic contractions, which constantly mix and stir the
contents of both stomach chambers. These contractions
are controlled by the release of
aneurotransmitter chemical called acetylcholine. The
research team (McLeay et al. 1999) studied the impact
of several different tremorgens on acetylcholine release,
& thus on contraction of the muscles in the reticulum and
rumen.

Effect of ergovaline on reticulum contractions.

Image courtesy of Lance McLeay.

They found that at least some of the mycotoxins they

tested disrupted muscle contractions in the rumen and
reticulum. This disruption was sometimes severe and
lasted up to 12 hours. They concluded that "severe
disruption of digestion may occur in animals grazing
endophyte-infected pasture" (McLeay et al. 1999),
affecting both milk/meat production and the animals'
health. The endophytic fungi also produce another set of
compounts - the ergots (similar to adrenalin) that
increase body temperature, induce heat stress, and
cause marked effects on contractions of the rumen and
reticulum (as shown in the above image). This mycotoxin
may cause scours (diarrhoea) in affected animals.

return to top
Ruminant ecophysiology
Ruminant animals range in size from 10-600 kg i.e. they
are in the mid size-range of vertebrate animals. Why are
there no ruminants with mass <10kg? Smaller animals
have higher energy requirements per unit of body
weight. The small body size means that their small guts
would not be able to cope with the high retention times
and throughput of a ruminant digestive system. In
comparison, large nonruminants such as giraffes and
elephants have comparatively lower energy
requirements than ruminants per unit of body weight.
Therefore, they are able to extract enough energy from
plants without the need for rumination. These non-
ruminant herbivores have a somewhat different digestive
anatomy. The three forestomach chambers are absent,
replaced with a single secretory stomach, and plant
material is fermented in the caecum and large intestine.
This process is known as hindgut fermentation.

return to top

Monogastric digestion

Humans, horse, dogs and rabbits all have monogastric

digestion they have a single stomach chamber.
Animals with monogastric digestion are still able to
digest some of the cellulose in their diet, by way of
symbiotic gut bacteria. However, their ability to extract
energy from cellulose digestion is less efficient than in
ruminants. Monogastric animals have evolved a number
of behavioural and anatomical adaptations that improve
their ability to use plants as a food source.

Hindgut fermenters
Horses, rabbits, rodents and other herbivores use
cellulose and other fermentable plant material in much
the same way as ruminants. However, as they only have
a single stomach compartment, fermentation and
digestion of cellulose primarily occurs in the large
intestine and cecum.

Digestive tract anatomy of hindgut fermenters

The stomach and small intestine of hindgut fermenters

are similar in form and function to other non-ruminant
herbivores. Food passes down the oesophagus and into
the stomach (which does much the same job as the
ruminant abomasums). However, monogastric species
do not regurgitate their cud so all mastication occurs
when food is taken into the mouth. Acid and enzymatic
digestion begins in the stomach, and then partially
digested food moves from the stomach into the small
intestine where further breakdown and absorption of
nutrients occurs. Like the ruminant system, the small
intestine empties its contents into the caecum through
the ileocecal orifice.

However, unlike other monogastric herbivores, the

caecum and large intestine of hindgut fermenters is very
large and anatomically complex. The caecum and colon
in much the same way as the reticulorumen of ruminant
animals; slowing down the throughput of food and
allowing time for the fermentation of cellulose and other
plant material bysymbiotic bacteria. Food leaves the
caecum through an opening called the cecocolic
orifice and moves into the ascending colon. Here regular
muscle contractions in the colon wall efficiently mix the
digested food and allow absorption of water, salts and
the nutrients produced through fermentation. Food
moves only slowly through the colon in horses it takes
2-3 days to pass the length of the colon. The lower colon
absorbs water and concentrates waste material before
this is egested as faeces through the rectum.
Scientists believe that hind-gut fermentation is
comparatively less efficient than rumination when it
comes to extracting nutrients from cellulose-rich plant
material. This has driven the evolution of a number of
behavioural and physical adaptations in monogastric
animals that maximise the energy they extract from their
high-fibre diets.

return to top

Adaptations to high fibre diets

Horses
Horses lose a lot of the proteins produced by microbes
living in their colons. This is because there is no
opportunity for the animal to absorb many of the amino
acids that the microbes generate. To compensate for
this, horses consume comparatively more food than
ruminant animals (Duncan et al. 1990). This is helped by
the fact food travels faster through a horse's digestive
system than it does through a cow's gut..

Rabbits & Hares

Rabbits and hares (lagomorphs) are all small animals
and have a high metabolic rate. This means that they
need to obtain energy from their food quickly, eating little
and often. Rather than retaining food in the gut for long
periods of time, lagomorphs have evolved another
process to allow them to extract more nutrients from their
food. Plant material is digested in the caecum by
symbiotic bacteria. This produces volatile fatty acids,
which are absorbed through the lining of the caecum and
provide approximately 30% of the animal's energy
needs. The caecum contents then pass through the
large intestine and are egested as caecal pellets - which
are promptly eaten by the animal as they leave the anus!
(This process is called caecotrophy, and usually
happens while the animal is in its burrow.) The pellets
pass through the digestive system a second time,
allowing more nutrients to be extracted, and the
remaining undigested material leaves the body as
normal faecal pellets.

Rodents and omnivores

Many rodents are either partly or wholly herbivorous.
Their generally small size means that they have high
metabolic energy requirements but little physical
capacity for retentive digestion of vegetative matter.
Therefore, ingestion of plant material is generally
restricted to high energy sections of the plant such as
fruit, nectar and pollen or seeds; or to sections that are
more easily digestible such as growing tips, seedlings
and flowers. This selectivity is also practiced by
omnivorous animals such as bears, pigs, possums and
humans.

return to top

Human digestive system

An adult human's digestive tract is approximately 6.5

meters long and consists of the pharynx, oesophagus,
stomach, small intestine and large intestine. Digestion in
humans is similar to that of other monogastric animals.
However, unlike most herbivorous animals, humans
have a relatively small caecum with
a vermiform appendix. The appendix is a blind-ended
tube connected to the caecum near the point where the
small intestine joins the large intestine. The appendix
appears to be a vestigial structure, reduced in size and
function when compared to the same structure in other
animals. One explanation for this is that the human
appendix was once much larger and served a similar
function to the caecum of hind gut fermenters. Over
time, the diets of early humans changed to include more
meat and less high-fibre plant material. This meant that
there was no selective advantage in having a large
appendix (and in fact there would be an energy cost in
maintaining it), and individuals with a smaller appendix
became more common over time . Modern humans
would have difficulty extracting enough nutrients if they
were restricted to a diet similar to that of ruminant
animals. While we are encouraged to eat a diet high in
vegetables and fruit, that diet is generally restricted to
easy-to-digest material that is relatively low in
cellulose: fruit, flowers and new stems and leaves. In
other words, our diet is restricted by our inability to
extract sufficient nutrients from high-cellulose plant
material.

return to top

References

Brooker R.J., Widmaier E.P., Graham L.E. & Stiling P.D.

2008: Biology. McGraw-Hill. New York..

Nickel, R., Schummer, A., & Seiferle, E. (1973) The

Viscera of the Domestic Mammals. Verlag Paul Parey,
Berlin.

Dijkstra J. 2005: Quantitative Aspects of Ruminant

Digestion and Metabolism (2nd Edition). CABI Publishing.
Wallingford.

Duncan P., Foose T.J., Gordon I.J., Gakahu C.G. & Lloyd
M. 1990: Comparative nutrient extraction from forages
by grazing bovids and equids: a test of the nutritional
model of equid/bovid competition and
coexistence. Oecologia 84: 411-418.

Lofgreen G.P., Meyer J.H. & Hull J.L. 1957: Behavior

patterns of sheep and cattle being fed pasture or
soilage. Journal of Animal Science 16: 773-780.

L.M. McLeay, B.L. Smith & S.C. Munday-Finch (1999)

Tremorgenic mycotoxins paxilline, penitrem and lolitrem
B, the non-tremorgenic 31-epilolitrem B and
electromyographic activity of the reticulum and rumen of
sheep. Research in Veterinary Science 66: 119-127.

return to top

Monogastric Stomach - Anatomy & Physiology

Overview
Stomach Anatomy 1 - Copyright Nottingham 2008

The enlarged swelling of the gastrointestinal tract between the oesophagus and
the duodenum is called the stomach. It is a simple structure in carnivores and a
compound structure in ruminants.
The stomach functions as a reservoir of food where digestion occurs through chemical
and mechanical processes. This allows food to be broken down further and absorbed.
Development

The gut tube is formed from the folding of the splanchnopleure (mesoderm and
endoderm). The endoderm is the inner layer forming the epithelia and glands. The
layers around it are from the mesoderm forming the skeletal muscle,
(oesophagus and anus), smooth muscle (from lateral plate mesoderm) and connective
tissue.
The region enlarges and a swelling indicates where the stomach will form. The dorsal
surface becomes convex to form the greater curvature of the stomach and the ventral
surface becomes concave to form the lesser curvature. Two rotations of 90 degrees
occur along the longitudinal axis and then the dorso-ventral axis. The
dorsal mesogastrium becomes elongated (with the spleen) and expands into a large
fold along the ventral abdominal wall. This becomes the greater omentum which
covers all the abdominal organs. It is a superficial structure which is free to move.
The ventral mesogastrium becomes the lesser omentum. It is in between the
stomach and the liver. The rest of the ventral mesentry degenerates.
Stomach Anatomy 2 - Copyright University of Nottingham 2008

Structure and Function

The stomach is split into regions: cardia, fundic, body and pyloric. The entire stomach
is motile. It has a pH of 0.9 to 1.5.
The larger part of the stomach lies to the left of the midline, under cover from the
ribcage and in contact with the liver and diaphragm. The oesophagusopens into it at the
cardiac sphincter. The smaller part of the stomach has thicker walls and passes to the
right of the midline into the duodenum at the pyloric sphincter. The angular point
between the two parts of the stomach is called the angular notch (incisura).

Stomach Anatomy 3 - Copyright University of Nottingham 2008

The fundus is a blind dome rising above the cardia. The body extends from the cardia
ventrally and the pyloric part is on the right divided into a more muscular and a less
muscular half. The serosa (external peritoneum) covers the entire organ.
Contractions start near the cardia and spread distally, accelerating and becoming more
vigorous as they reach the pyloric region. The pyloric sphincter is open for 1/3 of the
time during contractions. The empty stomach lies completely within the rib cage and
does not contact the abdominal floor. Little secretion is produced and only small
peristaltic contractions occur. Once food is offered or anticipated, the secretions begin.
The stomach is supported by 4 folds of peritoneum:

Gastrophrenic ligament - from the greater curvature of the stomach to the crura
of the diaphragm
Lesser omentum- connecting the lesser curvature of the stomach and the initial
segment of duodenum to the liver in the region of the hepatic porta
Gastrosplenic ligament- connecting the greater curvature of the stomach to
the spleen by a double fold of peritoneum
Greater omentum- connecting the greater curvature of the stomach to
the duodenum and dorsal body wall
Vasculature

Vasculature of the stomach includes the coeliac artery (which is a

branch of the dorsal aorta). The coeliac artery splits into the hepatic
artery supplying the liver, pancreas and stomach (right gastric and
right gastro-epiploic arteries). The coeliac artery also splits into the
splenic artery which supplies the spleen and the stomach (left gastro-
epiploic artery), it also splits into the left gastric artery supplying the
stomach.
The gastro-epiploic arteries supply the greater curvature of the
stomach and the gastric arteries supply the lesser curvature of the
stomach. The numerous veins join the portal vein.
Innervation

Sympathetic fibres run with the arteries. Parasympathetic fibres

from the vagus nerve (CN X) are within the two vagal trunks. In
the proximal region of the stomach, vagal
stimulation suppresses muscular contraction (VIP) and in
the distal region, vagal stimulation increases muscular activity
(ACh).
Histology
 Glandular Stomach Histology Dog - Copyright RVC 2008

The monogastric stomach has a columnar epithelium. The folded

mucosa of the stomach forms longitudinal rugae. The folds form
invaginations calledgastric pits which are continuous with gastric
glands.
Layers of the stomach
The 4 layers of the stomach wall are:

1. Serosa/adventitia

2. Tunica muscularis

3. Submucosa

4. Mucosa
There are 3 layers of muscularis: the outer longitudinal, middle
circular and inner oblique. The pyloric sphincter is a thickened
tunica muscularis from the middle circular smooth muscle layer. In
the fundic region, the tunica muscularis is thinner, the glands are
straight and the gastric pits are shallow. There is also an abundance
of parietal and chief cells in the gland. In the pyloric region, the
tunica muscularis is thicker, the glands are coiled and the gastric pits
are deep. The cardia is a narrow muscle strip. Lymphatic vessels are
present in the submucosa.
Cell types
The stomach contains basophilic chief (zygomatic) cells that
secrete pepsinogen in response to vagus nerve stimulation and
gastrin release. Pepsinogen unfolds and cleaves itself (autocatalyses)
in response to hydrochloric acid, therefore only in acidic
environments.
 Fundic Region of Glandular Stomach (Parietal and Chief cells)-
Copyright RVC 2008

The stomach also contains goblet cells that secrete mucous. This
secretion protects against autodigestion.
There are also parietal (oxyntic) cells found in gastric pits that
secrete hydrochloric acid. This secretion aids digestion by activating
gastric enzymes, e.g. pepsinogen to pepsin. Hydrochloric acid kills
microorganisms and enzymes that enter with food. Hydrochloric acid
is secreted in response to vagus nerve stimulation or pepsin
secretion. Parietal cells are large and pyramid shaped, with a higher
abundance in the upper region of the glands.
Gastrin is released from pyloric G cells; Somatostatin is released
from pyloric D cells and histamine is released from Enterochromaffin-
like cells (ECL cells).

Lamina Muscularis 3 Regions - Copyright RVC 2008

Stomach glands are short, coiled, branched and tubular. They need to
be replaced due to wear and tear and are only found in the mucosal
layer.
Digestive Enzymes
The digestive enzymes include;
Proteases
They are secreted as an inactive zymogen, activated by hydrochloric
acid. Active pepsin is produced and completed near the brush border
to generate small peptides and individual amino acids. It starts in the
stomach and continues into the small intestine.
Carbohydrases
Carbohydrases, e.g. amylase, are within salivary and pancreatic
secretions and act to produce disaccharides. Disaccharides are
converted to monosaccharides near the brush border. Cellulases are
digested by symbiotic micro-organisms in the (ruminant stomachs).
Lipases
Lipases are assisted by bile salts which neutralise stomach acids and
emulsifies fats. The process generates free fatty acids,
monoglycerides and diglycerides.
Control of secretions

The release of gastric secretions is

under hormonal (gastrin), paracrine (histamine) and neural (ACh)
mediator control in the cephalic and gastric phases. Gastric
secretions are inhibited during the intestinal phase by CCK and
secretin.
For more information, see control of secretions.
Species Differences

The size of the non-glandular region in the simple stomach varies

between species. It is largest in the horse, pig and then smallest in
the dog.

Equine Stomch with Margo Plicatus - Copyright RVC 2008

Equine
A region called the margo plicatus is present which separates the
glandular and non-glandular parts of the equine stomach. The non-
glandular area is lined with squamous epithelium (not columnar). The
stomach is relatively small (10% GIT) with a strong cardiac
sphincter which prevents the animal from vomiting. The equine
stomach is rarely empty, retention time is short and expulsion into
the duodenum stops when feeding stops. A 500kg horse can produce
30l of gastric juice in 24 hours.
Canine
The stomach is of variable size ranging from 0.5 to 6l according to
breed. A full stomach can touch the bladder. The subglandular layer
of fibroblasts and collagen fibres are for protection, e.g. from
consuming bones. It is between the glands and the lamina
muscularis. A 25kg dog can produce 0.5 to 1l gastric juice in 24 hours.
Porcine
The cardia is thickened, taking up nearly half the area of the
stomach. The internal diverticulum is present, which can be seen
externally.

Links

Click here for Stomach Pathology.

Click here for Control of Feeding.