Development of the Female Reproductive System

Gonadal Development

Indifferent Stage
The primordial germ cells will migrate from the yolk sac
to the gonadal ridge during the indifferent stage. These
cells pass through the early primitive streak and
become located as a small cluster of cells in the
extraembryonic mesoderm. These cells become
incorporated into the endoderm of the posterior wall of
the yolk sac, where they are dislocated from the
embryonic disc. Later on, the PGCs shift to a site in the
mesoderm along the yolk sac and allantois stalks.

From there they apparently migrate in the wall of the hindgut and through the dorsal mesentery
until they reach the newly formed genital ridge. PGCs divide during migration to the gonadal
ridge. PGCs undergo mitotic arrest. The site of development for gonads is an elongated region
of steroidogenic mesoderm along the ventromedial border of the mesonephros.
When the PGCs arrive in the gonadal ridge, the
resident mesenchymal cells and the coelomic
epithelium proliferate and, consequently, the
developing gonadal ridges moves towards the
coelomic cavity. Cords of epithelial cells from the
mesonephric tubules and regressing glomerular
capsules goes into the mesenchyme of the
genital ridge and form irregularly shaped cords,
the primitive sex cords or gonadal cords, that
incorporate PGCs. In both sexes these cords are
temporarily connected to the surface epithelium.

Differentiation
Gonadal cords
In female animals, the expression of Dax-1, with the absence of SRY gene, may result to the
suppression of the formation of a testis and allows the indifferent gonads to develop into
ovaries.
The sex cords derived from the coelomic
epithelium leak into the mesenchyme and
grow towards the mesogenital region and
combines with mesonephric tubules.
These primary cords, called medullary
cords, contains some primitive germinal
cells, but most frequently remain less well
developed and decay. The next set of sex
cords is named cortical cords, which
reaches only the cortical region of the
future ovary. After following breakdown of
cortical cords, germ cells undergo a
period of enhanced mitotic activity, which ceases shortly after birth or before having birth.
Oogonia is obtained during this period of multiplication hundreds of thousands of primordial
germinal cells. High percentage of oogonia later undergo degenerative change called atresia.

As the period of the mitotic activity is completed, individual oogonia becomes surrounded by a
layer of squamous cells, called follicular cells. There are three sites of origin of follicular cells
have been proposed: cells of the coelomic epithelium, primitive sex cords of mesonephric
origin, and a combination of both.
One oogonia surrounded by follicular cells constitutes a
primordial follicle. The follicular cells induce the enclosed
oogonia to enter the prophase of first meiotic division, which
results in the formation of a primary oocyte. Period of growth is
the phase when the germ cells develop. The primary oocytes do
not progress to the tertiary stage of development, period of
maturation, until stimulated by gonadotrophic hormones at the
onset of puberty. At the end of the period of growth in domestic
animals, except horses. Rete ovarii consists of peripheral cortical
area which contains the primordial follicles, and central
medullary area composed of degenerating intra-gonadal tubules.
The superficial germinal epithelium of the ovary doesn´t revert to
mesothelium, therefore the ovary is not covered with the
peritoneum in the adult. The connective tissue of the ovary
develops from the mesenchyme. Below the germinal epithelium
becomes denser, forming fibrous web called tunica albuginea

Descent of the ovaries

The ovaries migrates caudally similar to the testes from their origin on the poster abdominal wall. The
ovaries moves to their final position within the true pelvis.

Later on the gubernaculum becomes the round ligament and ovarian ligament.
Sexual Duct System Development
Undifferentiated

The undifferentiated sexual duct system consists of

paired mesonephric ducts (Wolffian ducts) and
paired paramesonephric ducts (Müllerian ducts). The
paramesonephric, Müllerian ducts is formed
bilaterally in both male and female embryos on the
lateral side of the mesonephros, close to the
mesonephric duct. A longitudinal paramesonephric
invagination develops in the coelomic epithelium of
the mesonephric ridge. Afterwards, the coelomic
epithelium invaginates into the underlying
mesenchyme forming a longitudinal groove. Margins
of this groove attach and grow together. The
peritoneal lining separates from the duct. At the
cranial end of the duct forms a funnel, which opens
into the body cavity. The Mullerian duct is located laterally and in parallel with Wolffian duct. Its
caudal end elongates caudally and then turns medially. Whereby the Wolffian duct crosses
ventrally, approaches to the opposite end and both open into the posterior wall of the urogenital
sinus at the Müllerian prominence

Differentiated
The absence of anti-Müllerian hormone enables the
Müllerian ducts to develop into the female
reproductive tract, which comprises the oviductus
(Fallopian tube), uterus, cervix and vagina. The
mesonephric ducts regresses, but occasionally a
vestigial structure occurs as a small duct lying
parallel to the uterine tube, extending from the
epoophoron through the broad ligament to the
vagina. Gartner´s duct or canal may form a cyst in
the vaginal wall. Epoophoron is the vestige of the
mesonephric tubules localized near the developing
ovary. The paroophoron is from the mesonephric
tubules located caudally to the developing gonad.
The anterior parts of Müllerian ducts open into the coelomic cavity by the extended funnel,
which develops into the infundibulum of oviductus. Towards the posterior end, the Müllerian
ducts turn medially and their transverse portions cross mesonephric ducts ventrally. From there
the pars uterina of oviductus, and isthmus develop. The epithelial lining of the oviductus rises
from the coelomic epithelium of the mesonephric ridge. The next layers of the wall have the
mesenchymal origin. After Müllerian ducts reach the midline, the posterior ends fuse, giving rise
to uterovaginal canal (canalis uterovaginalis), from which the vagina and uterus develop. The
uterovaginal canal will grow toward the urogenital sinus. The Mullerian tubercle is formed at the
point of contact, the uterovaginal canal protruding into the urogenital sinus. After merging the
paramesonephric ducts in the midline, a broad traverse pelvic fold is established. This fold
extends from the lateral sides of the fused ducts toward the wall of the pelvis, is the broad
ligament of the uterus.

4 main forms of mammalian uterus

Uterus duplex – marsupials, rodents and lagomorphs

there are two fully separated uteri and both cervices open separately
into the common vagina

Uterus simplex- higher primates

the entire uterus is fused to a single organ, thus the oviducts open
into one common cavity.

Uterus bipartitus- ruminants, hyraxes, cats, and horses.

the cranial portions of the uterus remain separate, giving rise to

cornua uteri separated by an intercornual septum. The caudal
potions fuse, forming the uterine body, which remains divided into
two cavities by the partition, but enters the vagina by a single cervix.
Uterus bicornis- dogs, pigs, elephants, whales, dolphins, and ungulates

The horns are distinct for less than half their

length; the lower part of the uterus is a common
chamber, the body.

The vagina rises from combined posterior ends of the Müllerian ducts. After they join the
urogenital sinus, the blind ends of Müllerian ducts fuse with the lining of the urogenital sinus to
form the epithelial vaginal plate. Subsequently, cannulation of these fused structures occurs,
forming the lumen of the vagina. The lumen is separated from the urogenital sinus by a thin
membrane and the hymen which consists of the epithelial lining of the sinus and a thin layer of
vaginal cells. In animals, the hymen subsequently breaks down, the remnants persist only
rarely. The caudal portion of the urogenital sinus forms the vestibulum vaginae.

External Genitalia Development

Undifferentiated
The external genitalia are derived from three complexes of mesodermal tissue located around
the cloaca. At the anteroventral end of the cloacal membrane, the genital tubercle is formed.
Lateral to the cloacal membrane, the cloacal (urogenital) folds is extending most of its length.
These are subdivided into the anal folds anteriorly and posteriorly urethral folds. Distance
between the anus and the base of the genital tubercle are commonly used to differentiate male
from female fetuses; a disparity is first detected at Day 30 in canine and Day 42 in bovine
embryos. After that time, the anogenital distance remains constant in the female and increases
in the male. Peripheral to the urogenital folds and posterolateral to the cloacal membrane, are
the genital and labioscrotal swellings. These are present and similar in both sexes prior to sex
differentiation. Shortly after the appearance of the genital tubercle, and prior to the completion of
the development of the urorectal septum, the epithelial lining of the floor of the urogenital sinus
expands anteroventrally along the ventral margin of the elongating genital tubercle. These
endodermal cells form the urethral plate, a solid cord of cells extending inward from the ventral
surface of the tubercle. The urethral plate hollows to form a canal and the urogenital folds
enlarge by proliferation of mesenchymal cells on both sides of the urethral plate. A median
urethral groove becomes established on the ventral surface of the genital tubercle.
Differentiated

In the female, the urogenital folds that

border the urogenital orifice fuse only at
their dorsal and ventral ends and form
the labia minora. They overgrow the
genital tubercle, which becomes
internalized in the floor of the vestibule.
Another consequence of the failure of
the urogenital folds to fuse is that the
urogenital sinus opening does not
become incorporated into the phallus
and the genital tubercle does not enlarge into a penile structure, as in the male. Instead, it gives
rise to the clitoris, which appears vestigial in most domestic animal species. However, the
genital tubercle does form a glans, and therefore the small clitoris has both a body and a glans.
The genital swellings shift cranially to the genital tubercles and disappear during fetal
development. Therefore no labia majora are formed in domestic animals as they are in humans.
The urogenital sinus remains as the vestibule, with openings from both the vagina and urethra.
The female urethra, which develops from the more cranial part of the urogenital sinus is
homologous to the prostatic urethra of the male.