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20.

1 Formation of the urinary organs


 Development of the urinary organs (3rd - 4th week)

Development of the urinary organs (3rd - 4th Quiz


week) Quiz 07

The development of the urinary system is closely related to that of the genital systemwhich will be described, Quiz
though, in a separate module. The urinary organs consist of the kidneys (which produce urine, among other
Quiz 16
things), the ureter (transport of the urine from the kidneys to the bladder), the bladder (temporary storage for
the urine) and the urethra (transport of the urine from the bladder to the external world).

In all vertebrates the kidneys and ureters develop out of the intermediate mesoderm, whereas bladder and
urethra derive from the urogenital sinus.

Fig. 1 - Transverse section and dorsal view of an embryo (trilaminar) Legenda


(ca. 21 days)
Fig. 1
Schematic diagram:
Transverse section (in the
midcephalic region) with a
dorsal view of the three-
layered embryo towards the
end of the 3rd week of
development. The
intermediate mesoderm lies
between the somites
(paraxial mesoderm) and the
lateral mesoderm (out of
which the coelom arises).

1 Paraxial mesoderm
2 Intermediate mesoderm
3 Lateral mesoderm
4 Notochord
5 Amnion
6 Intraembryonic coelom
7 Endoderm
8 Ectoderm
9 Somatopleural (mesoderm and ectoderm)
10 Splanchnopleural (mesoderm and endoderm)
11 Neural groove
12 Neural ridge

Due to the lateral folding, the intermediate mesoderm is shifted ventrally and loses its connection with the Quiz
somites and the lateral mesoderm.
Quiz 01
The nephrogenic cord develops out of Fig. 2 - Development of the Legenda
the intermediate mesoderm 9 and extends intermediate mesoderm (stage 10)
from the cervical to the caudal region. It becomes Fig. 2
segmented like the paraxial mesoderm (somites) (Fig. Transverse section of an
embryo at the beginning of
1). This segmentation is easily seen in the cranial
the folding (ca. 28 days). In
region, and is rudimentary in the middle region. In this diagram the nephrogenic
the caudal region it isno longer present, as we will cord is clearly distinguished
describe in detail below. Due to the growth of the inner in the intermediate
structures of the embryo, the tissue lying most mesoderm. The black arrow
laterally is displaced ventrally. This leads not only to indicates the border between
a separation of the nephrogenic cord from the the paraxial and intermediate
paraxial and lateral mesoderm, but also to a mesoderm.
coalescence of certain median structures such as the
two dorsal aortas 12 which fuse to form the
definitive (median) aorta.

1 Neural tube
2 Amniotic cavity
3 Paraxial mesoderm
4 Intermediate mesoderm
5 Yolk sac (umbilical vesicle)

Fig. 3 - Isolation of the nephrogenic cord (stage 11) Fig. 4 - Isolation of the nephrogenic cord (stage 12) Legenda
Fig. 3, Fig. 4
The intermediate mesoderm
moves ventrally and loses its
connection (black arrow) to
the somites and the lateral
mesoderm. Also observe its
approach towards the
intraembryonic coelom

1 Neural tube 5 Yolk sac (umbilical vesicle)


2 Amniotic cavity 6 Notochord
3 Dermatomyome 7 Aorta
4 Nephrogenic cord that has moved away from the 8 Intraembryonic coelom
paraxial mesoderm (somite) (black arrow)
20.2 Development of the renal anlage
 Inleiding
 Pronephros (forekidney): transitory structure
 Mesonephros: transitory kidney
o Mesonephros: first excretory organ

 Metanephros: definitive kidney

Inleiding
The kidneys develop along a cranio-caudal gradient. Typically, the development passes through three stages: Color code
 Pronephros for the
urinary system
 Mesonephros module

 Metanephros Urine excreting system:


yellow-beige-
orange
In principle, for all three developmental stages with similar designations, the same embryonic anlage - the Urine forming system:
intermediate mesoderm - is involved. pink-lilac-violet
The term "holonephros" can thus be employed to designate all the parts of the urinary system that arise from it.
The first two developmental stages have a transitory character and the definitive kidneys actually develop from
the metanephros stage.

Pronephros (forekidney): transitory structure


The pronephros develops during the 4th week, beginning in the cranial part of the nephrogenic cord and it
atrophies during the 5th week. Three characteristic pronephros structures can be distinguished:
 The pronephros duct 10 in the neck region
 The pronephros tubules 11

 The external or coelom glomeruli 13 that have been shown to exist in humans (1)
Beginning with the 4th week, conforming to the cranio- Fig. 5 - Early pronephros development Legenda
caudal gradient, the pronephros in the neck region (stages 10-12)
divides into independent masses of cells, the Fig. 5
nephrotomes 11 . Schematic sagittal section
Each nephrotome develops into anepithelialized through a
pronephros glomerulus. Laterally, they form 4-week-old embryo. One can
the pronephros tubules that can partly bind with the recognize the segmented
coelom. Via the fusion of these tubules between two part of the nephrogenic cord,
nephrotomes the hollow pronephros duct arises that the pronephros and the
is the anlage of the pronephric collecting duct. In unsegmented part, the
humans, this pronephros system corresponds more to mesonephros. In this
a primitive and transient structure that is stage 12 the
functionally of no importance. According to the mesonephric duct already
classical view, the pronephros duct stops in the discharges into the cloaca.
caudal region at the level of the 13th -14th somite and
then goes over into themesonephric duct (Wolffian
duct).

1 Nephrogenic cord
2 Mesonephric duct (Wolff)
1+2 Mesonephros
3 Intestinal tube
4 Cloaca
5 Atrophying nephrotomes
6 Yolk sac (umbilical vesicle)
7 Allantois
8 Outflow of the mesonephric duct into the cloaca

Meer info
The origin of the mesonephric duct (Wolffian duct):
At present it is not sure whether the mesonephric duct (Wolffian duct) arises from the joining up of
these pronephros tubules or from the first mesonephric vesicles. (2)
Mesonephros: transitory kidney Quiz 02
Quiz

The mesonephros differentiates itself during the 4th week and after the 8th week it degenerates. It replaces the Quiz
pronephros and develops from three structures:
Quiz 17
 Nephrogenic cord 10 in the dorso-lumbal region
 Mesonephric duct 11

 Glomerular capillary network 13


The mesonephric duct forms on the dorsal side of the Fig. 6 - Development of the Legenda
nephrogenic cord at the level of the 9th somite. mesonephros
Initially it consists of a solid mesenchymal cord of Fig. 6
cells 11 . It releases itself from the nephrogenic Sagittal section of a 5-week-
old embryo. The pronephros
cord and is finally localized under the ectoderm,
atrophies while the
which probably plays an inductive role in its formation mesonephric duct grows
(3). caudally and fuses with the
Released from the nephrogenic cord, it develops in cloaca wall. In this stage, it
the caudal direction andcanalizes itself at the same goes through a
time 12 , in order to finally end in the cloaca. As mesenchymal- epithelial
soon as it is canalized one calls it themesonephric transformation and forms a
duct (Wolffian duct). At the site where the central lumen. Only the
caudal part remains
mesonephric duct (Wolffian duct) discharges into the
mesenchymal. Observe the
cloaca, the rear wall of the bladderforms. ureter bud at its caudal end.

1 Nephrogenic cord
2 Mesonephric duct
1+2 Mesonephros
3 Intestine
4 Cloaca
5 Atrophied nephrotome
6 Yolk sac (umbilical vesicle)
7 Allantois
8 Outflow of the mesonephric duct
into the cloaca
9 Ureter bud (anlage)

Fig. 7 - Development of the mesonephros as the first excretory organ Legenda


Fig. 7
Transverse section along A
(see previous figure).
The urogenital ridge projects
into the lumen of the coelom.
With the S-shaped
mesonephric tubules the
mesonephric duct (Wolffian
duct) forms a transitory
precursor of the adult
excretory system. The medial
end of the mesonephric
tubule is closed and forms a
funnel (Bowman’s capsule)
that surrounds a tuft of
capillaries (the glomerulus).
The capillaries come from
lateral branches of the dorsal
aorta and drain into the
inferior cardinal vein.
This functional unit is also
termed the excretory unit of
1 Neural tube the mesonephros.
2 Notochord
3 Aorta dorsalis
4 Dorsal mesentery
5 Intestinal tube
6 Ectoderm
7 Somite
8 Inferior cardinal vein
9 Mesonephric duct (Wolffian duct)
10 Mesonephric tubule
11 Urogenital ridge

Meer info Quiz


The mesonephric duct (Wolffian duct) Quiz 03
Animal experiments have shown that the growth and the caudal extension of the mesonephric duct depend on
the presence of the extracellular molecule fibronectin (an integrin). The differentiation of the solid cord into a
duct depends on the secretion of the BMP4 (bone morphogenetic protein 4) by the ectoderm (4).

Mesonephros: first excretory organ

Shortly after the differentiation of the mesonephric duct, through mesenchymal-epithelial transformation, the Quiz
mesonephric vesicles 11 arise out of the nephrogenic cord, which represents a mesoderm condensate. Quiz 15
Via the mesonephric tubules they connect up with the mesonephric duct 14 . This differentiation takes
place bilaterally in the area between the upper thoracic region (Th1) and the lumbar region (L3). A cranio-caudal
gradient is also visible here. To the extent in which new mesonephric vesicles and tubules develop caudally, the
cranial elements begin to atrophy so that never more than 30 pairs exist in the mesonephros. They form the
excretory system that closely resembles the adult nephrons.

Medially, the mesonephric vesicle dead-ends in that it forms a funnel (Bowman's capsule). Each of these
funnels surrounds a tuft of capillaries (glomerulus), which have also arisen in the nephrogenic tissue and come
from lateral (visceral) branches of the dorsal aorta. They drain into the inferior cardinal vein.
The capsule with glomerulus together form a renal corpuscle. A renal corpuscle and its associated tubule are
called a nephron and the functional unit an excretory mesonephric unit.

The production of urine begins in the mesonephros during the 6th week 17 . After the 10th week these nephrons become inactive and
atrophy. While in the female all mesonephric tubules atrophy completely, in the male a few that lie caudally remainin order to develop into
the testicular efferent ducts.

Metanephros: definitive kidney

The metanephros develops from three intermediate mesoderm structures of the


sacral region:
 Ureter anlage 14
 Metanephric vesicle 13

 Glomerular capillary network 13

The ureter anlage 14 is an epithelial Fig. 9 - Development of the Legenda


diverticulum from the caudal part of metanephros: 5th week
the mesonephric duct (Wolffian duct) Fig. 9
in the area of the first sacral vertebra Sagittal section of a 5-
(S1). week-old embryo -
The anlage intrudes into the metanephric development of the
vesicle and forms the extra- and metanephros.
intrarenal excretory passages. In the caudal region of
The metanephric blastema 13 the nephrogenic cord one
corresponds to the sacral part of the observes the
nephrogenic cord below L3. It is development of the
mesenchymal tissue out of which the metanephro-
metanephric vesicles arise. From these genic blastema that is in
originate the nephrons (= functional contact with the ureter
units of the kidneys). anlage. In this stage the
At present it is still not clear whether the pronephros has
glomerular capillary network develops disappeared almost
through vasculogenesis (direct 1a Pronephros (atrophying) completely. The
development of vessels from the 1b Mesonephros (atrophying) mesonephros is also in
metanephric vesicles) or through 2 Mesonephric duct the process of
angiogenesis (development from existing (Wolffian duct) atrophying.
vessels of the metanephros) (5). 3 Nephrogenic cord
4 Ureter anlage
5 Metanephric blastema
6 Liver anlage
7 Cloaca

20.3 The upper urinary tract: development


of the metanephros
 Ureter anlage and metanephric blastema: reciprocal induction
 Morphologic differentiation of the urinary tract
o The ureter anlage and the formation of the evacuating urinary tract
o The metanephric blastema and the formation of the nephron
 Functional development

 Ascent of the kidneys

Ureter anlage and metanephric


Quiz
blastema: reciprocal induction Quiz 21

The interaction between the ureter anlage (epithelial tissue) and the metanephric
blastema (mesenchyma) is of decisive importance for renal development. The
development of the kidneys represents a classical model of a sequential and
reciprocal induction between epithelium and mesenchyma. For this reason it is
frequently used for investigating the molecular cell mechanisms that play a role in the
entire organogenesis.
Renal development comprises a whole series of developmental processes such as
forming an epithelial tree structure, interactive tissue induction, differentiation,
polarization, migration, cell adhesion and finally the epithelio-
mesenchymaltransformation.

During the genesis of the metanephros the metanephric blastema first induces the
branching of the ureter anlage, which, for its part, then lets the metanephric
vesicleform into a predetermined blastema. Through the transformation into epithelial
tissue, the renal tubules form and finally the nephrons emerge. The most recent
molecular-biologic research, mainly on transgenic mice, has shown that several factors
are involved in this process. These various factors can be gone into only briefly in this
chapter; those wishing to dig deeper can have a look at the cited references.

Meer info
Genes and their products that are important in the formation of the upper urinary tract.

Morphologic differentiation of the


urinary tract
The ureter anlage and the formation of the evacuating
urinary tract
The ureter anlage 14 is an epithelial diverticulum which arises from the caudal Quiz
portion of the mesonephric duct (Wolffian duct) at the level of the first sacral vertebra. Quiz 04
Its enlarged end grows in the dorso-cranial direction and projects into themetanephric
blastema.
Quiz
It is the origin of the intra- and extrarenal excretory passages:
Quiz 08
 Ureter
 Renal pelvis
Quiz
 Major and minor calices
Quiz 09

 Collecting ducts

Fig. 10 - Schematic cut through the Fig. 11 - Enlargement of the inset Legenda
kidney (sagittal section) in Fig. 10
Fig. 10, 11
The branching of the
collecting ducts and
calices from the renal
pelvis is shown. The
enlargement in B
shows the collecting
duct and the various
elements that form the
nephron.
Meer info
Histological picture of
the vessel
architecture of the
renal cortex

1 Ureter 8 Distal tubule Meer info


2 Major calix 9 Proximal tubule Meer information on
3 Minor calix 10 Glomerulus fig. 11.
4 Renal pelvis 11 Connecting tubule
5 Collecting duct 13 Intermediate tubule
6 Metanephric vesicle
7 Kidney lobe
The cranial end of the ureter anlage Fig. 12 - Development of the Legenda
subdivides dichotomically many times due metanephric outflow
to the inductive effect of the metanephric Fig. 12
blastema. The renal excretory
passages develop from
the ureter anlage. This
development is
described in greater
detail in an interactive
diagram of region C
(see below).

1 Cloaca
2 Ureter anlage
3 Metanephric blastema
2+3 Metanephros
4 Mesonephric duct (Wolffian duct)
5 Nephrogenic cord
4+5 Mesonephros

The renal pelvis arises from the swollen end of the ureteric bud 16 that subdivides
4 times over the course of the 4th to 6th weeks. 16 branches arise from this that
partially then coalesce again in order to finally form 2 to 4 major calices.
At around the 7th week the minor calices start to develop. They discharge into
themajor calices. Further dichotomic branching follow - up to the 15th generation
(roughly 32 weeks). Thereby the caliber of the tubules is reduced more and more so
that finally 1-3 millions collecting ducts are formed in the periphery of the
metanephric blastema 18 .

The metanephric blastema and the formation of the


nephron
The process of nephron formation is complicated and thus subdivided into various Quiz
steps: Quiz 10

 The metanephric blastema surrounds each newly formed collecting duct. It


condenses in order to form peritubular cell aggregates.

 Through induction signals (see: beginning of the chapter), derived from the
ureter anlage, the mesenchymal cells transform themselves and form vesicles.

 These vesicles grow longer and form an "S"-shaped tubule with three
sections.
o Development of the distal section into the distal part (distal Meer info
convoluted limb, thick ascending limb and thin ascending limb) Histology of the renal
o Development of the middle section into the proximal part (thin corpuscle
descending limb, proximal straight and convoluted tubule)
Scanning electron
microscopy (SEM) of a
o Development of the proximal section in the glomerular capsule vascular cast
21 . The epithelial vesicle secretes angiogenic factors. Thereby, (following corrosion of
over the course of the further development, endothelial cells are adherent tissue).
brought into the glomerular capsule. As soon as the afferent vessels
come into close contact with the vesicular epithelium, it flattens and
forms a cup with a bilaminar structure, Bowman's
capsule (formation of the renal corpuscle: interactive
diagram, overview over all the pictures).
 At the same time as the formation of the renal corpuscle, the distal end of the
epithelial vesicle fuses with the neighboring collecting duct. The metanephros
thus becomes able to function and can filter the plasma from the glomeruli.
Through the proximal tubule the glomerular filtrate (primary urine) gets into Meer info
the intermediate tubule, the distal tubule, connecting tubules and collecting Renal diuresis in
duct. In these tubules the secondary urine arises through resorption and fetuses and newborns.
secretion processes. It then reaches the renal pelvis and, via the ureter, the
bladder. During the pregnancy, the fetal urine is excreted into the amniotic
cavity.
Physiology of the kidneys Quiz
The fetal kidneys do not have to maintain the water and electrolyte households in the Quiz 11
organism nor do they have to clean the blood from substances (mainly end products
of cellular metabolism in the form of urea, uric acid and creatinine ) that are usually
eliminated with the urine. These functions are performed by the placenta. The kidneys
are, however, involved in the production of amniotic fluid. This is why an agenesis of
the kidneys on both sides leads to an oligohydramnios (deficiency of amniotic fluid),
which is described in greater detail in the chapter on pathology.
Naturally, besides their excretory function, the kidneys also have an endocrine one in
that they secrete hormones (renin, erythropoetin, prostaglandin and kallikrein). Even
though some of the endocrine functions only begin at birth, the embryonic kidneys
nevertheless have an important task to fulfill, namely the production of erythropoetin.
In addition, renin-positive cells can already be found in the mesonephros of 5-6 week
old embryos (19).

Summary:
 The urine-forming system (filtration, resorption and secretion): distal,
intermediate, and proximal tubules and the renal corpuscles (Bowman's
capsule with capillaries) arise from the metanephric blastema of
themetanephros.

 The urine excreting system - the ureter, the renal pelvis, the major and
minor calices as well as the collecting ducts - arise from the ureteric bud
anlage from the Wollfian duct.

Functional development Quiz 05


Quiz

Quiz
Quiz 24

At birth the kidneys have a multilobular Fig. 13 - Development of the Legenda


appearance, due to the development of smooth kidney surface
the ureter anlage in the metanephric Fig. 13
blastema. Normally, towards the end of In humans renal
the fetal period, the lobes are lobulation is easily
seen only in the
considerably smoothed, but they still exist embryonic and fetal
until after birth. periods, but can extend
Completion of the smoothing follows into childhood.
during childhood by the increase in At birth, however, it is
volume of the connective tissue and strongly attenuated.
theincrease in size of the In adults, the cortical
nephrons without any change in their zones of the individual
lobes, "A to E", fuse
number. With only few exceptions, adult
and the smoothing of
kidneys no longer exhibit any their depressions leads
lobulation. to a smooth and
Since the renal architecture is finalized uniform renal surface
between the 5th and 15th weeks of structure.
intrauterine development; organogenesis
of the kidneys lasts well beyond the
embryonic phase until far into the fetal
period. 1 Renal medulla
2 Calix minor
3 Renal cortex

Fig. 14 - Anatomy of the kidney at the end of its development Legenda


Fig. 14
The renal
parenchyma is divided
into an outer cortical
zone and an inner
medullar zone. In its
interior the kidney has
a hollow space, the
renal sinus, into which
the papillas intrude
and where the trunks
of the renal vessels
are found.
Meer info
Histology

Quiz
Quiz 12

1 Ureter 7 Cortex
2 Renal pelvis 8 Capsula renis
3 Vena renalis 9 Medullary rays
4 Arteria renalis 10 Papilla renalis
5 Major calix 11 Sinus renalis
6 Minor calix 12 Columna renalis
13 Medullary pyramid

Fig. 15 - Vascular supply of the kidney at Legenda


the end of the development
Fig. 15
Detail of the vascular
supply of a renal lobe.
One sees the renal
pyramid surrounded
by the interlobar
vessels as well as
their continuations,
the arcuate vessels.
From them originate
the interlobular
vessels that form the
afferent arterioles,
followed by a capillary
network (glomerulus;
not shown here).

Quiz
Quiz 06
10 Papilla renalis
14 Interlobar artery
15 Interlobar vein
16 Arcuate artery
17 Arcuate vein
18 Interlobular arteries and veins

Ascent of the kidneys Quiz 13


Quiz

The metanephros is formed in the sacral region at the level of the first sacral vertebra Quiz
(S1) and the bifurcation of the aorta. In the adult, the kidney lies at the upper lumbar Quiz 19
level. The "migration" (ascent) of the kidney occurs between the 6th and 9th week and
the kidneys finally come to lie at the level of the 12th thoracal vertebra (Th12) under the
suprarenal glands.
The mechanism that leads to this ascent of the kidneys is not an active migration but
rather much more the result of the differing growths of the sacral and lumbar regions,
which lead to an unfolding of the lower pole of the embryonic body. During its ascent,
the kidney is supplied by a number of transitory vessels that all originate from the
aorta. The definitive renal arteries stem from the lumbar region of the aorta, while the
transitory vessels normally disappear.

It must also be mentioned here that Fig. 16 - Migration of the kidneys Legenda
during their development the kidneysturn Stage 15 (ca. 36 days)
90° towards the vertebral cords, so that Fig. 16
the hili are medially oriented at the end, Ascent of the kidney at
while they originally face ventrally. the end of the 5th
week. The kidneys still
(Animation). find themselves in the
sacro-lumbar region.
Various anomalies are connected with this The mesonephros,
ascent and will be described in detail in which in this stage is
the chapter on pathology. still to be found under
the undifferentiated
gonads, is in the
process of atrophying.

1 Gonad
2 Mesonephros
3 Allantois
4 Tuberculum genitale
5 Cloacal membrane
6 Cloaca
7 Posterior intestine
8 Metanephros
9 Mesonephric duct
(Wolffian duct)
10 Ureter anlage

Fig. 17 - Migration of the kidneys Fig. 18 - Migration of the kidneys Legenda


Stage 18 (ca. 44 days) Stage 23 (ca. 56 days)
Fig. 17
Ascent of the kidneys
at the end of the 7th
week. At this point the
ureter - independent of
the mesonephric duct
(Wolffian duct) -
inserts into the
bladder. The
mesonephros has
disappeared.

Fig. 18
Ascent of the kidneys
at the end of the 8th
week. The kidneys find
themselves at the
upper lumbar level in
1 Gonad 1 Gonad their definitive position
3 Bladder being formed 3 Bladder being formed and a
4 Genital tubercle 4 Genital tubercle
5 Cloacal membrane 6a Rectum
6 Rectum 7 Posterior intestine
7 Posterior intestine Metanephros 8 Metanephros
8 Metanephros 9 Mesonephric duct
9 Mesonephric duct (Wolffian duct)
(Wolffian duct) 10a Ureter
11 Urogenital orifice
12 Anal orifice

20.4 Lower urinary system


 Subdividing the cloaca
o Separating the cloaca
o The perineum and the urorectal septum
 Development of the bladder

 Development of the male and female urethra

Subdividing the cloaca Quiz 22


Quiz

We have seen that the upper urinary system - consisting of the collecting ducts, the
calices, the renal pelvis, and the ureters - arises from the ureter anlage.
The lower urinary system - composed of the bladder and the urethra - is formed from
the endoderm of the posterior intestine.

Separating the cloaca


In stage 13 13 the cloaca is the common end of the rectal tube and the urogenital
tract. Towards the outside it is closed by the cloacal membrane.
Between the 4th and 6th weeks the urorectal septum separates the cloaca into
aprimary urogenital sinus (ventrally) and the rectum (dorsally).
 The bladder and the pelvic limb of the urethra arise from the primary
urogenital sinus and the caudal portion of the urethra comes from
the definitive urogenital sinus (see Fig. 26)

 The urorectal septum divides the cloacal membrane into two membranes:
theurogenital membrane (ventrally) and the anal membrane (dorsally).
These two membranes atrophy 19 , like the bucco-pharyngeal
membrane 11 , in order to form the intestinal and urogenital openings.

Fig. 19 - Development of the cloaca, Fig. 20 - Migration of the kidneys, Legenda


Stage 13, roughly 32 days Stage 23, roughly 56 days
Fig. 19
The white arrow
indicates the direction
of growth of the
urorectal septum. This
separates the cloaca
into a urogenital sinus
(ventrally) and the
rectum (dorsally).
Fig. 20
In this diagram the
kidneys are found in
their definitive position
1 Urorectal septum 3 Urogenital orifice
at the level of the
2 Cloacal membrane 4 Anal orifice
upper lumbar region.
5 Cloaca 6 Urogenital sinus
The urorectal septum
8 Allantoïs 7 Rectum
(white arrow) has
8a Future bladder
divided the cloaca. The
urogenital sinus,
bounded on the
outside by the
urogenital orifice, lies
ventrally. The rectum,
which opens to the
outside through the
anal orifice, is found
behind it. The cloacal
membrane disappears
in stage 19.
The perineum and the urorectal septum
Today, the urorectal septum is no longer regarded as an isolated cellular layer of
mesoderm cells that slowly grow towards the cloacal membrane. It consists of two
mesodermal structures that are fused together.
An upper fold (Tourneux), located frontally, grows caudally. Near the cloacal
membrane two lateral folds (Rathke) form that fuse at the median level.

They subdivide the cloaca and also grow Fig. 21 - Development of the urorectal Legenda
in the direction of the upper fold septum
(Tourneux) that is located frontally. A Fig. 21
disorder in the formation of these two The urorectal septum
structures leads to recto-urethral or consists of two
different mesoderm
rectovesical fistulas. Connective tissue structures.
and the perineal musculature, which Over the course of the
keep the pelvic organs in place, arise 4th week an upper fold
from the mesoderm, which surrounds the (Tourneux) divides the
rectal tube. The central fibrous part of the cloaca in the cranio-
perineum corresponds anatomically to the caudal direction. The
region between the anal and urogenital two lower, lateral folds
(Rathke) are
orifices.
responsible for the
1 Peritoneal cavity
division in the lower
2 Upper fold (Tourneux) (pink arrow)
section.
Lower fold (Rathke) (blue arrow)

Fig. 22 - Separation of the cloaca Fig. 23 - Separation of the cloaca Legenda


and the formation of the perineum and the formation of the perineum

1 Peritoneal cavity 5 Urogenital orifice


2 Upper fold (Tourneux; pink arrow) ; 6 Anal orifice
Lower folds (Rathke; blue arrows) 7 Perineum
3 Primary urogenital sinus
4 Anal canal

Development of the bladder


The bladder develops from the upper part of the urogenital sinus (UGS) and is
connected with the allantois.
The allantois is obliterated during the development and forms a fibrous cord, the
urachus, which following birth becomes the median umbilical ligament.

While the cloaca is being divided, the caudal, originally common part of the
mesonephric duct (Wolffian duct) and the ureter anlage is taken up into the upper,
postero-lateral wall of the urogenital sinus (future bladder).
The rapid growth of the back wall of the Fig. 24 - Development of the bladder Legenda
urogenital sinus has the result that the Fig. 24
common lowest part of the ureter and the Dorsal view of the
mesonephric duct (Wolffian duct) are both entry level of the
taken up into the bladder wall. Further mesonephric canal
complicated growth processes have the (common extremities
result that the ureteral orifices and the of the mesonephric
orifice locations of the mesonephric duct duct (Wolffian duct)
(Wolffian duct) go through a cranio- and the ureter) while it
caudal positionexchange during the is incorporated into
course of the further development. The the dorsal wall of the
ureteral openings appear to migrate bladder.
thereby in a cranio-lateral direction and
the mesonephros orifices appear to be
shifted caudo-medially. The triangular Animation
zone that is thus created is termed Reversal of the orifice
the vesical trigonum. In males, the 1 Primary urogenital sinus locations of the ureter
wolffian duct forms the future deferent 2 Mesonephric duct (Wolffian duct) and mesonephric duct
duct on both sides. 3 Ureter anlage (Wolffian duct) in the
4 Common drainage urinary bladder wall
passages
5 Pelvic limb of the urogenital sinus

A Part of the genital tract


B Part of the urinary tract

The trigonum thus originates from the mesoderm while the ventral bladder wall has an endodermal origin.
Later, though, the trigonum will be completely covered by endodermal epithelial cells. The smooth musculature of
the bladder develops during the 12th week from the splanchnopleural mesoderm, which coats the endoderm on
the outside.

Development of the male and female


urethra
The urethra forms itself from the lower part of the urogenital sinus (UGS).

 In a man the prostate and membranous part of the urethra arise from the Meer info
pelvic part of the UGS while the spongy urethra comes from the phallic part In the genital tract
(urethral plate). module you will learn
more about the
development and
 In a woman the whole urethra and part of the vagina arise from the pelvic part differentiation of the
of the UGS while the phallic part (urethral plate) forms the vestibule and the urogenital sinus
labia minora. inmales and females.

Fig. 26 - Development of the urethra out of the cloaca


SUR Septum urorectal