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In males: Differentiation of the

Quiz
accessory sex glands Quiz 29

The accessory sexual glands originate from two epithelial tissues. They either come
from the epithelial mesodermal origin of the mesonephric duct (Wolff) or from
theepithelial endodermal origin of the urogenital sinus.

The seminal vesicle differentiates itself Fig. 37 - Differentiation of the Legend


during the 12th week from a protrusion internal sex canals in the 4th month
on the deferent duct, near where it Fig. 37
opens at the back wall of the urogenital Median sagittal section
sinus (future prostatic part of the urethra). at the level of the
prostate shows the
They thus have a mesodermal origin. development and
These paired glands produce a viscous, differentiation of the
fructose-rich secretion that serves as seminal vesicle and the
asource of energy for the sperm cells. prostate
The portion of the mesonephric duct that
lies between the junction of the seminal
vesicle and the prostatic part of the
urethra is called the ejaculatory duct.
Between the two ejaculatory duct
junctions the prostatic utricle (remainder
of the paramesonephric duct) has its
opening. This location is called
theseminal colliculus.
1 Ureter
2a Deferent duct
2b Ampulla of the deferent duct
3 Seminal vesicle
4 Ejaculatory duct
5 Prostatic utricle
6 Prostate
7a Prostatic part of the urethra
7b Diaphragmatic part of the urethra
8 Ureter opening in the urinary
bladder
9 Urinary bladder

The prostate develops from a protrusion Fig. 38 - Male urethra with Legend
on the dorsal wall of the prostatic part of accessory glands
the urethra during the 12th week. Fig. 38
Theprostate's glandular Schematic front section
epitheliumdevelops, therefore, from cells through the male sex
canals shows the
that have their origin in the endoderm, differentiated
while the stroma and smooth muscle accessory glands.
develops from cells with a mesodermal
origin - under the inducing influence of
DHT (see: hormonal factors in sex
differentiation). The glands become active
after the 15th week and surround the two
ejaculatory ducts and the prostatic utricle
as well as the prostatic part of the urethra.

Over the course of the 12th week and


parallel to the development of the
prostate, bulbourethral (Cowper's) and
urethral (Littre's) glands form, originating
in pairs of endodermal protrusions of
the spongy part of the urethra, which
follow from the prostatic membranous
parts. Finally, the seminal fluid is
augmented by secretions from the
seminal vesicle, the prostate, the
bulbourethral and urethral glands.

1 Deferent duct
2 Ampulla of the deferent duct
3 Seminal vesicle
4 Ejaculatory duct
5 Prostate
6 Outflow canals of the prostate
7 Bulbourethral gland (Cowper's)
8 Urethral gland (Littre's)
9a Prostatic part of the urethra
9b Membranous part of the urethra
9c Spongy part of the urethra

In males: Differentiation of the


Quiz
urogenital sinus Quiz 12

In studies of the urinary system we have seen that the cloaca is subdivided by the uro-
rectal septum into the rectum (dorsal) and the primary urogenital sinus (ventral).

The primitive urogenital sinus is in contact with the allantois. The upper part forms
the future urinary bladder, while the middle and lower part forms the definitive
urogenital sinus and, from it, the pelvic and phallic parts of the urethra (stage 17).

Fig. 39 - Development of the urogenital Fig. 40 - Development of the urogenital Legend


sinus, stage 13 (ca. 32nd day) sinus, stage 23 (ca. 56th day)
Fig. 39
The cloaca subdivides.
Observe that the
metanephros in this
stage still lies quite
caudally.

Fig. 40
The uro-rectal septum
has subdivided the
cloaca into the primary
urogenital sinus
(ventrally) and the
1 Allantois 4 Phallic part of the definitive urogenital
rectum (dorsally).
2 Cloaca sinus
Out of the primary
3 Urinary bladder 5 Rectum
urogenital sinus arise
6 Pelvic part of the definitive urogenital
the urinary bladder
sinus
and the definitive
urogenital sinus.

The pelvic part of the definitive urogenital Fig. 41 - The development o fhe urethra Legend
sinus forms the prostatic and ca. 12th week
membranous parts of the urethra, Fig. 41
thephallic part of the definitive The spongy part of the
urogenital sinus, the spongy part of the urethra comes to a
dead end in the outer
urethraand the urethral vestibule. section of the penis.
Around the 4th month an epithelial Only in the 4th month
cordgrows on the glans penis - into the does an epithelial
depths - and binds it with the spongy part lamella form from the
of the urethra. tip of the penis to the
blind end of the
urethra. From this
arises the urethra
within the glans.
1 Urachus
2 Rectum
3 Urinary bladder
4 Ureter
5 Urethra, prostatic part
(orange zone)
6 Membranous part of the urethra
7 Spongy part of the urethra
8 Seminal vesicle
9 Bulbourethral gland (Cowper's)
10 Deferent duct

Summary:
From the mesonephric duct (Wolff) arise:
 Epididymis
 Deferent duct
 Seminal vesicle
 Ejaculatory duct

From the endoderm of the urogenital sinus arise:

 Urinary bladder
 Out of the pelvic part of the definitive urogenital sinus: the prostatic and
membranous parts of the urethra, the prostate and the bulbourethral gland
(Cowper's).

 Out of the phallic part of the definitive urogenital sinus: the spongy part of the
urethra and the urethral glands (Littre's)

In females: Differentiation of the


Quiz
canal system in the genital organ Quiz 13

During the 7th week the canal system of the female sex organs differentiates. Quiz
Themesonephric duct and its tubules atrophy and out of the paramesonephric Quiz 22
duct(Müller) arises the future fallopian tube, the uterus and the upper part of the
vagina.
Reminder
Sometimes, a few embryonic remnants of the mesonephric duct remain in the form of Scheme showing the
the epoöphoron, the paroöphoron at the level of the mesovarium, and a row of molecular factors
involved in the
smallcysts of Gartner. development of the
sexual apparatus
Out of the upper, non-fused portion of the paramesonephric duct (Müller) arises the
fallopian tube and its ampulla. The lower section fuses after it crosses medially on both
sides of the inferior ovarian gubernaculum and forms the utero-vaginal canal. The
medial septum in between disappears at the end of the 3rd month.

Fig. 42 - Differentiated female sex organs, ca. 4th month Legend


Fig. 42
Shown is the atrophied
mesonephric duct
(Wolff) that, however,
leaves certain
embryonic remnants
behind. Out of the
paramesonephric duct
(Müller) arise on both
sides the fallopian
tubes and through
fusion of both sides the
uterus and the upper
part of the vagina
(blue). The lower part
of the vagina (yellow)
comes from the
urogenital sinus
(endoderm). To be
noted is also the
development of the
ligaments and the
hymen, the middle part
1 Epoöphoron of which usually
2 Paroöphoron disintegrates at around
3 Ovarian ligament the time of birth.
4 Atrophied mesonephric duct (Wolff)
5 Cysts of Gartner
6 Hymen
7 Suspensory ligament of ovary
8 Fallopian tube (ampulla)
9 Vesicular appendage (Morgani)
10 Uterus
11 Round ligament of uterus
12 Vagina
13 Insertion of the round ligament of uterus at the genital swelling

Overview
Table for comparing the differentiation of the internal sex organs in both sexes
To be observed is the development of the Fig. 43 - Formation of the uterus, Legend
ligaments. The ovarian gubernaculum 7th – 8th weeks
gets attached on the developing utero- Fig. 43
vaginal canal there where it goes over Up to the 7th week two
into the fallopian tube. Above it forms the canal systems on each
side exist in both
ovarian ligament and below the round sexes. In the 8th week
ligament of uterus, which goes through the paramesonephric
the inguinal canal and inserts in the ducts (Müller) fuse in
female genital swelling (labia majora). the lower portion after
If the separating wall beyond the fusion they have crossed
location of the two paramesonephric medially on both sides
ducts is not resorbed, various utero- of the mesonephric
duct (Wolff).
vaginal abnormalities result.

Fig. 44 - Formation of the uterus Fig. 45 - Formation of the uterus Legend


after 8 weeks ca. 3rd month
Fig. 44
Formation of the utero-
vaginal canal through
fusion of the lower
section of the two
paramesonephric ducts
(Müller). From the
upper section - on both
sides - arise the
fallopian tubes with
their ampullae.

Fig. 45
At the end of the 3rd
month the separating
wall dissolves in the
uterus and the vagina.
1a Paramesonephric duct (Müller) 1b Fallopian tube The uterus lengthens
2a Mesonephric duct (Wolff) 2b Atrophied mesonephric duct (Wolff) in that the solid lower
3a Lower gubernaculum 3b Ovarian ligament end of the
4a Utero-vaginal canal 3c Round ligament of uterus paramesonephric duct
5a Urogenital sinus 4b Uterus stretches in a
5b Vagina downward direction
and is subsequently
canalized. Out of the
lower section arises the
upper part of the
vagina. It joins with
the vaginal lamina,
which arises from the
urogenital sinus and
forms the lower portion
of the vagina.

The blind end of the utero-vaginal canal forms the sinu-vaginal eminence 22 and Quiz
ends at the back wall of the urogenital sinus (SUG). Quiz 15
The sinu-vaginal eminence becomes thicker due to epithelial proliferation and retracts,
while the wall of the SUG also thickens there. These epithelial layers, which form at the
lower end of the utero-vaginal canal, are known as the vaginal plate. At their cranial
end they form a circular protrusion, the location of the future vaginal fornix.
Fig. 46 - Female sex organs Fig. 47 - Female sex organs Legend
ca. 7th week ca. 12th week
Fig. 46
The utero-vaginal
canal comes up against
the urogenital sinus
and forms the sinu-
vaginal eminence.

Fig. 47
This sinu-vaginal
eminence becomes
thicker due to
epithelial proliferation.
This also leads to a
epithelial proliferation
in the SUG epithelium.
Together they form the
vaginal plate.

1 Genital tubercle 4 Perineum


2 Vestibule 5 Rectum
2a Urovaginal sinus: pelvic part 6 Utero-vaginal canal
2b Urovaginal sinus: phallic part 7 Urinary bladder
3 Vaginal plate 8 Urethra

Through canalization of the vaginal plate Fig. 48 - Female sex organs Legend
the utero-vaginal canal opens itself ca. 3rd month
towards the outside. The upper 3/4 of the Fig. 48
vagina comes from the mesoderm and The canalization of the
the lower fourth from the endoderm. vaginal plate begins in
the 3rd month.
The fibromuscular walling forms from
the neighboring mesenchyma. The
vagina is separated from the SUG by the
hymen. Its origin is not entirely clear.
Discussed is a passive invagination of the
back wall of the SUG.

Fig. 49 - Female sex organs Fig. 50 - Female sex organs Legend


ca. 5th month ca. 9th month
Fig. 49
In the 5th month the
vaginal canal is
completely canalized,
but the lumen is
separated from the
SUG by the hymen.

Fig. 50
Normally, the hymen
tears open at the time
of birth. The uterus
and the vagina then
have a connection to
the vaginal vestibule.
2 Vaginal vestibule
3a Uterine cavity
3b Uterine cervix (neck) Animation
6a Vagina: The lower fourth out of
endoderm Differentiation of the
6b Vagina: The upper 3/4 out of female sex organs
mesoderm
9 Hymen

In females: Differentiation of the


accessory sex glands
The accessory glands arise from the endoderm of the SUG:

 The greater vestibular glands (Bartholini) are paired glands that form in the
course of the 12th week from the endoderm of the SUG. The outflow canals
empty sideways in the vaginal vestibule. They correspond to the bulbourethral
gland (Cowper) in males.

 The lesser vestibular glands (Skene) or paraurethral glands also form from
epithelial buds (endoderm) of the SUG and grow into the neighboring
mesenchyma. They are distributed over the whole vaginal vestibule and - in
males - correspond to the prostate.

In females: Differentiation of the


Quiz
urogenital sinus Quiz 14

In females the cranial portion of the pelvic part of the definitive SUG remain narrow
and forms the female urethra, which is very short.
With the progressing development the caudal portion of the pelvic part shrinks and
becomes included in the phallic part (vaginal vestibule). Therein discharge the urethra
and the vagina.
The phallic part of the definitive SUG enlarges to become the vaginal vestibule that
is caudally closed off externally by the urogenital membrane. It then tears in the course
of the 7th week 19 .

Fig. 51 - Urogenital sinus (SUG) Fig. 52 - Vaginal vestibule of the uterus Legend
ca. 12th week and vagina, ca. 9th month
Fig. 51
In females, the
development of the
SUG begins in the 3rd
month, at the same
time as the formation
of the vagina.

Fig. 52
The pelvic part of the
SUG has shrunk and
will be retracted into
the phallic part in order
to form the definitive
vaginal vestibule.

1 Genital tubercle 2 Vaginal vestibule


2 Vestibule 3a Uterine cavity
2a SUG: Phallic part 3b Uterine cervix
2b SUG: Lower part of the pelvic part 6a Vagina: Lower fourth out of endoderm
3 Vaginal plate Vagina: Upper 3/4 out of mesoderm
4 Perineum 6b Hymen
5 Rectum
6 Utero-vaginal canal 9
7 Urinary bladder
8 Urethra: From the upper portion of the
pelvic part of the definitive SUG

Summary: Overview
From the paramesonephric duct (Müller) arises: Formation of the
 Uterus ovaries and the
female genitalia
 Fallopian tube
 Vagina (3/4)
Overview
The embryonic
From the SUG endoderm arises: structures and their
derivatives in both
sexes
 Vagina (1/4)
 Urethra
 Vaginal vestibule

 Urethral glands, para-urethral and vestibular

Summary: Development of the


internal genital organs
In males the internal sex organs come from the mesonephric duct (Wolff) that
differentiates itself into the epididymis, deferent duct, seminal vesicle and the
ejaculatory duct. The paramesonephric duct (Müller) atrophies. It leaves behind
embryonic remnants such as the testicular appendage (hydatid) and parts of the
prostatic utricule.

In females the paramesonephric duct (Müller) remains in existence and


differentiates itself into the fallopian tube with its ampullae and, following its fusion at
the caudal end, into the uterus and the upper part of the vagina. The mesonephric
duct (Wolff) with its tubules atrophies and leaves embryonic remnants such as the
ductus longitudinalis epoöphori (Gartner), epoöphoron and paroöphoron.

21.5 The external genitalia


 Indifferent stage
 Differentiated stage of the male genitalia

 Differentiated stage of the female genitalia

Indifferent stage Quiz

Quiz 27

During the third week the cloacal membrane is formed and lies below the umbilical
cord.
In the fourth week, as the lower abdominal wall is being formed, the cloacal
membrane is shifted caudally by immigrating mesenchyma coming from various
sources.
It is now delimited in front by a prominent mesenchymatous eminence with an epithelial
covering, the anlage of the future genital tubercle.

end of the 5th week severalcloacalFig. 54 - Indifferent external genitalia, Legend


form on both sides of the cloacal membrane. At ca. 6th week
their anterior end they join, delimited by the genital
Fig. 54
In front, the cloacal
membrane is delimited
by a swelling, the
future genital tubercle,
that at the rear
continues in the two
cloacal folds.

Genital tubercle
Cloacal folds
Cloacal membrane

During the 7th week the urorectal septum 16 divides the cloacal membrane into
aurogenital membrane (ventrally) and an anal membrane (dorsally). From this
moment on the ventral section of the cloacal swellings around the urogenital sinus are
called urethral folds and in the dorsal section around the anus, anal folds. Lateral to
the urethral folds a further pair of tori form, the genital swelling.

As we have seen, the urogenital membrane dissolves during the 7th week 19 , so
that the urogenital sinus communicates freely with the amniotic cavity.

Fig. 55 - Indifferent external genitalia, Fig. 56 - Indifferent external genitalia, Legend


ca. 7th week ca. 9th week

Fig. 55
During the 7th week
the urorectal septum
(8) subdivides the
cloacal membrane, so
it is separated into
urogenital and anal
membranes.

Fig. 56
From the cloacal folds
around the urogenital
orifice arise the
urethral folds and, from
the one around the
anal orifice, the anal
Genital tubercle folds. Outside the
Urethral Anal
folds orifice urethral folds a further
Anal Genital
folds swelling prominence arises on
Urogenital orifice both sides, the genital
swelling.

It is important to remember that the morphology of the external genitalia in both sexes is very
similar up to the 9th week.

Differentiated stage of the male


genitalia
Under the effects of androgen the differentiation and development of the male sexual
organs become visible after the 3rd month.

The genital tubercle becomes longer and out of it forms the penis. The urethral
foldsalso lengthen ventrally. Between these extends the urogenital sinus and forms
theurethral groove, which is lined with endoderm. The floor of this sulcus thickens
through epithelial proliferation and forms the urethral plate that temporarily fills it out.
Later a groove forms again and the two urethral folds fuse on the underside. This
section will become the spongy part of the urethra, which for now terminates in a dead Reminder
end in the anterior part of the penis. Diagrams ofindifferent
In the rear part, the genital swellings transform themselves into the scrotum. stages

From the fused urethral folds an erectile Fig. 57 - Differentiated external male Legend
mesenchymatous tissue, the genitalia, ca. 10th week
penilespongy body, arises in the penis. Fig. 57
At the distal penis section a ring-shaped Lengthening of the
furrow delimits the glans. Above the genital tubercule and
formation of the penile
spongy body arise the two cavernous anlage. Proliferation on
bodies (corpora cavernosa) and thus the floor of the urethral
complete the penile erectile system. groove that is bounded
by the urogenital folds.
The two genital swellings also fuse in
the middle and form the scrotum. The
line along which they fuse on the penis
and scrotum is called the raphe mediana.

Fig. 58 - Differentiated external male Fig. 59 - Differentiated external male Legend


genitalia, ca. 12th week genitalia, ca. 14th week
Fig. 58
Increasing fusion of
the urehtral folds at
the ventral edge of the
penis, back to front.
This fusion forms the
definitive spongy part
of the urethra, which
ends blindly shortly
before the end of the
penis.

Fig. 59
The two genital folds
also fuse in the middle
and form the scrotum.
The line along the
scrotum and penis
2 Urethral groove 9 Raphe mediana (ano-genital) where they fuse is
4 Fused urethral folds 10 Urinary meatus called the raphe
5 Penis 11 Perineum mediana.
7 Genital swelling 12 Anus
8 Scrotum

During the 4th month two ectodermal invaginations arise on the tip of the penis.
First, a solid epithelial cord forms from the penis tip and binds itself with the dead-
ended spongy part of the urethra at the level of the ring-shaped furrow. As soon as
thisepithelial cord has been canalized, one speaks in this section of the glandar
urethrawith the urinary meatus.
The two circular epithelial ingrowths, glandar lamella, form the prepuce that at the time
of birth is still stuck to the glans but, during childhood, comes away from it.

Fig. 60 - Penis, prepuce, spongy part Fig. 61 - Penis, prepuce, spongy part Legend
of the urethra, ca. 12th week of the urethra, ca. 14th week
Fig. 60
In the spongy part of
the urethra forms
erectile tissue, the
penile spongy body,
which extends into the
glans penis. Above it
the two cavernous
bodies complete the
penile erectile system.

Fig. 61
From the 14th week
the two epithelial
hollows on the glans
penis open and create
a connection within the
glans between the
1 Epithelial hollow of the glans penis 1 Glandar part of the urethra with spongy and glandar
2 Prepuce epithelial depression the navicular fossa part of the urethra
3 Prepuce 2 Resorbed prepuce epithelial
4 Cavernous body ingrowth
5 Spongy body
6 Spongy part of the urethra

Overview table Animation


Overview table for comparing the differentiation of the external genitalia in males and Development of the
females. penis
Formation of the testes and the external genitalia

Differentiated stage of the female


genitalia
The mechanisms that are responsible for the development of the external female
genitalia are still not well understood. It is known, though, that the placenta produces
estrogen and the ovaries also appear to contribute some.

The genital tubercle lengthens only a little, retracts again and, after 14 weeks,
theclitoris is recognizable.
The genital swellings do not fuse and, for this reason, the urgenital sinus remains Reminder
wide open with the urethra in the anterior part and the vagina in the posterior Diagrams ofindifferent
part(vaginal vestibule). stages

The urethral folds also do not fuse. Out of Fig. 62 - Differentiated female genitalia Legend
them arise the labia minora. ca. 10th week
From the genital swellings arise the labia Fig. 62
majora. They fuse only in the rear portion In the female, the
and form the posterior commissure of genital tubercle
lengthens only a little
the labia. In the front they form the mons and shrinks again while
pubis. forming the clitoris.
The urethral folds do
Towards the rear, the posterior labial not fuse and the
commissure is continued by urogenital sinus
theperineum. remains wide open.

Fig. 63 - Differentiated female genitalia Fig. 64 - Differentiated female genitalia Legend


ca. 12th week fetus, from the 14th week
Fig. 63, Fig. 64
The urehtral folds do
not fuse – the labia
minora arise from
them. From the genital
swellings arise the
labia majora. They
fuse only in the rear
part and form the
posterior labial
commissure that is
continued towards the
rear by the perineum.

1 Genital tubercle 7 Perineum


2 Genital swelling 8 Posterior labial commissure
3 Clitoris 9 Anus
4 Labia minora 10 Hymen
5 Labia majora 11 External urethral orifice
6 Mons pubis

Overview table
Overview table comparing the differentiation of the external genitalia in males and in
females.
Formation of the ovaries and the external genitalia