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Mammals
Mammalian Sex Determination and
Dev. Of the Gonad
The gonads are also derived from the intermediate
mesoderm
Form testes or ovaries
But before they choose - come from a common
precursor structure = Bipotential Gonad!
In the chromosomes - how is sex determination
achieved in mammals?
Mammalian Sex Determination Overview
Bipotential gonad
starts as genital
ridge epithelium
in intermediate
mesoderm
Nephric duct
is the primitive
organizer:
Wolffian Duct
~ week 4 in humans
Differentiation of gonads in transverse section
Mullerian duct
develops parallel
to Wolffian duct
Differentiation of the male gonad
Cell Fates:
- Epithelium becomes Sertoli cells
- Mesenchyme becomes Leydig cells
- Germ cells will become spermatogonia
Ductal fates:
- Sertoli AMF degenerates Mullerian duct
- Leydig testosterone maintains Wolffian duct
- Testis cords will become seminiferous tubules
- Rete testis canals join testes cords to the
vas deferens and then epididymous.
Differentiation of the female gonad
Cell Fates:
- Epithelium becomes granulosa cells
- Mesenchyme becomes thecal cells
- Germ cells will become meiotic oogonia
- Together they form the follicles
Ductal fates:
- No AMF allows devo of Mullerian duct
- oviduct, uterus, cervix, upper vagina
- No testosterone degenerates Wolffian duct
Development of Male genital Ducts and Gland
Mesonephric ducts (Wolfian ducts) persist and are transformated into
the duct of epididymis.
Distal to epididymis mesonephric ducts acquires a thick investment of
smooth muscle and becomes to ductus deferens
Seminal Glands : Lateral outgrowths from caudal end of each
mesonephric duct. The part of mesonephric duct between the duct of
this gland and the urethra become ejaculation duct. (the secretion
nourish the sperms
Prostate : Multiple endodermal outgrowth arise from the prostatic
part of the urethra and grow into the surounding mesenchyme.
Secretions from the prostate contribute to the semen
Bulbourethral Glands : Pea sized structures developed from paired
outgrowths derived from the spongy part of urethra. Secretion
contribute to the semen
Development of Female Genital Ducts and Glands
The Paramesonephric ducts (Mullerian ducts) form
most of the female genital tract.
The uterine tubes developed from the unfused cranial
part of the paramesonephric ducts. The caudal, fused
portions of these ducts form the uterovaginal
primordium. Which gives rise to the uterus and the
superior portion of vagina
The Vaginal ephitelium is derived from the endoderm of
the urogenital sinus. The fibromuscullar wall of vagina
develops from surounding mesenchyme. Contact of the
ureterovaginal primordium with the urogenital sinus,
forming the sinus tubercle, induces the formation of
paired endodermal outhgrowths-sinovaginal bulbs. They
extend from the urogenital sinus to the caudal end of the
ureterovaginal primordium
Development of Female Genital Ducts and Glands
The sinovaginal bulbs fuse to form a vaginal plate. The
central cells of this plate break down, forming the lumen
of vagina.
Until late in fetal life, the lumen of the vagina is separated
from the cavity of the urogenital sinus by a membrane the
hymen formed by invagination of the posterior wall of the
urogenital sinus. The hymen rupturs during the perinatal
period
Female Auxiliary Genital Glands outgrowhts from the
urethra into surounding mesenchyme form the bilateral
mucus-secreting urethral glands and paraurethral
glands. Outgrowths from the urogenital sinus form the
greater vestibular glands in the lower one third of the
labia majora
Development of External Genitalia
Early in the fourth week, the proliferating mesenchyme
produces a genital tubercle primordium of penis or
clitoris.
In both sexes at the cranial end of the cloacal membrane.
Fgf 8 is involved in the signaling pathways in the early
development of the external genitalia
Labioscrotal swellings and urogenital folds soon
develop on each side of cloacal membrane. The genital
tubercle soon elongates to form primordial phallus
(penis or clitoris) . The urogenital membrane lies in
the floor of a median cleft, the urethral gr0ove, which
is bound by the urogenital folds.
In female fetuses, the urethra and vagina open into a
common cavity, the vestibule of vagina
Development of Male External Genitalia
Masculinization of the indifferent external genitalia is
induced by dihydrotestosteron from leydig cell
(testiscullar)
The primordial phallus enlarge and elongates to
become phenis, urogenital folds form the lateral wall of
the urethral groove on the ventral surface of the penis.
This groove is lined by a proliferation of endodermal cells
the urethral plate, which extends from the phallic portion
of urogenital sinus.
The urethral folds fuse with each other along the ventral
surface of penis to form spongy urethra.
The surface ectoderm fuses in the median plane of the
penis, forming the penile raphe and enclosing the spongy
uretra within the penis
At the tip of the glans penis, an ectodermal ingrowth,
extends toward the root of penis to meet spongy urethra
Development of Male External Genitalia
This cord canalizes and joint the previously formed
spongy urethra.
This juncture completes the terminal part of urethra
and moves the OUE to the tip of glans penis.
Corpora cavernosa and corpus spongiosum
developed from mesenchyme in the phallus.
The labioscrotal sweelings grow toward each other
and fuse to form scrotum.
Development of Female External Genitalia
Growth of the primordial phallus in the female fetus
gradually decreases as it becomes clitoris.
The clitoris develops in a similar way to the penis,
except that the urogenital folds do not fuse except
posteriorly, where they join to form the freenulum of
labia minora.
The unfused parts of the urogenital folds form the
labia minora .
The labioscrotal folds fuse posteriorly to form the
posterior labial commisure and anteriorly to form
the anterior labial commisure and mons pubis.
Most part of the labioscrotal folds remain unfused and
form two large folds of skin, the labia majora
Relocation of Testes and Ovaries
By 26 weeks, the testes have descend retroperitoneally from the
posterior abdominal wall to the deep inguinal rings. This change in
position occurs as the fetal pelvis enlarge and trunk of the embryo
elongates
Testiscular descent through ingunal canals and into the scrotum is
controled by testosteron. The gubernaculum guides testes through
inguinal canal and into the scrotum during the 26th weeks. When the
testes descend, they carry the ductus deferens decend. they are
ensheated by the fascial extensions of the abdominal wall :
The extension of the transversalis fascia becomes the internal
spermatic fascia
The extensions of the internal oblique muscle and fascia
become the cresmatic muscle and fascia
The extension of transversalis fascia becomes external
spermatic fascia
Within the scrotum, the testis projcts into the distal end of the
processus vaginalis tunica vaginalis (covers front & side testis)
Relocation of Testes and Ovaries
The ovaries also descend from the lumbar region of the
posterior abdominal wall and relocate to the pelvis;
however they do not pass from pelvis and enter the
inguinal canals.
The gubernaculum (fibrous cord) is attached to the
uterus near the attachment of the uterine tube.
The cranial part of the gubernaculum becomes the ovarian
ligament and the caudal parts forms the round ligament
of uterus.
The round ligament pass through inguinal canals and
terminate in the labia majora
The relatively small processus vaginalis in the female is
usually obliterated and it disappears long before birth, a
patent processus in a fetus is known as a vaginal process
of peritoneum (canal of Nuck)
Making decisions
Several human gene have been identified whose
function is necessary for normal sexual differentiation
The story starts in the genital ridge, which can become
either type of gonad. The genes encoding Wt 1, Sox9,
wnt 4,LHx9,Fgf9,GATA4 and Sf1 are expressed
Loss of function prevent the normal development
of gonad
Y chromosom (-) WNT4 , , Rspo Producing B
cathenin (+ feedback), Initiate the ovarian pathway
development, and prevent acumulation of Sox9 that
protein crucial for testis determination
Making decisions
If nucleus has a Y chromosome, the same set factor
activates the SRY gene. Sry protein binds to SOX9
promoter and elevates expression of this key gene in
the testis-determining pathway.
SOX 9 + FGF 9 block the ovary forming pathway,
blocking the function B-Catenin
BASICS!!! The male pathways : Make testes and dont
make ovaries.
The Female Pathways : Make ovaries and dont make
testes
The ovary pathway : Wnt4 and
R-spondin1
WNT 4 is expressed in genital ridges of both sexes,
later, undetectable in XY gonads . In transgenic xx
mouse lack the WNt4 gene, the ovary fails to form
properly.
Testosterone
True
Hermaphrodite
Weve also
seen
them in
worms...
Sex Determination in the Brain
Physical position
Bonellia
Larva drops to sea floor - female
Larva finds proboscis of adult female male
Population conditions
Goby fish
School is one male, many females
If male is lost, top females race to change
Most aggressive becomes dominant male
Figure 14.25 Sex determination in
Bonellia viridis
Note:
The mammalian X/Y system of sex determination
is not used in all species
For most species, other systems are used
In Drosophila a ratio system is used
Ratio between number of X chromosomes and number
of autosomes
if there is 1X, fly is male, if 2X fly is female
NOT dependent on Y
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