Development Of The Urinary And Reproductive Organs
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The development of the urinary system begins during
prenatal development Prenatal development () includes the development of the embryo and of the fetus during a viviparous animal's gestation. Prenatal development starts with fertilization, in the germinal stage of embryonic development, and continues in fetal devel ...
, and relates to the development of the
urogenital system The genitourinary system, or urogenital system, are the organs of the reproductive system and the urinary system. These are grouped together because of their proximity to each other, their common embryological origin and the use of common pathw ...
– both the organs of the
urinary system The urinary system, also known as the urinary tract or renal system, consists of the kidneys, ureters, urinary bladder, bladder, and the urethra. The purpose of the urinary system is to eliminate waste from the body, regulate blood volume and bl ...
and the
sex organ A sex organ (or reproductive organ) is any part of an animal or plant that is involved in sexual reproduction. The reproductive organs together constitute the reproductive system. In animals, the testis in the male, and the ovary in the female, a ...
s of the
reproductive system The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are als ...
. The development continues as a part of
sexual differentiation Sexual differentiation is the process of development of the sex differences between males and females from an undifferentiated zygote. Sex determination is often distinct from sex differentiation; sex determination is the designation for the deve ...
. The urinary and reproductive organs are developed from the
intermediate mesoderm Intermediate mesoderm or intermediate mesenchyme is a narrow section of the mesoderm (one of the three primary germ layers) located between the paraxial mesoderm and the lateral plate of the developing embryo. The intermediate mesoderm develop ...
. The permanent organs of the adult are preceded by a set of structures which are purely embryonic, and which with the exception of the ducts disappear almost entirely before birth. These embryonic structures are on either side; the
pronephros Pronephros is the most basic of the three excretory organs that develop in vertebrates, corresponding to the first stage of kidney development. It is succeeded by the mesonephros, which in fish and amphibians remains as the adult kidney. In amnio ...
, the
mesonephros The mesonephros ( el, middle kidney) is one of three excretory organs that develop in vertebrates. It serves as the main excretory organ of aquatic vertebrates and as a temporary kidney in reptiles, birds, and mammals. The mesonephros is included ...
and the metanephros of the
kidney The kidneys are two reddish-brown bean-shaped organs found in vertebrates. They are located on the left and right in the retroperitoneal space, and in adult humans are about in length. They receive blood from the paired renal arteries; blood ...
, and the
Wolffian Christian Wolff (less correctly Wolf, ; also known as Wolfius; ennobled as Christian Freiherr von Wolff in 1745; 24 January 1679 – 9 April 1754) was a German philosopher. Wolff is characterized as the most eminent German philosopher between Le ...
and
Müllerian duct Paramesonephric ducts (or Müllerian ducts) are paired ducts of the embryo that run down the lateral sides of the genital ridge and terminate at the sinus tubercle in the primitive urogenital sinus. In the female, they will develop to form the fal ...
s of the
sex organ A sex organ (or reproductive organ) is any part of an animal or plant that is involved in sexual reproduction. The reproductive organs together constitute the reproductive system. In animals, the testis in the male, and the ovary in the female, a ...
. The pronephros disappears very early; the structural elements of the mesonephros mostly degenerate, but the
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sper ...
is developed in their place, with which the Wolffian duct remains as the duct in males, and the Müllerian as that of the female. Some of the tubules of the mesonephros form part of the permanent kidney.


Pronephros and Wolffian duct


Wolffian duct

In the outer part of the
intermediate mesoderm Intermediate mesoderm or intermediate mesenchyme is a narrow section of the mesoderm (one of the three primary germ layers) located between the paraxial mesoderm and the lateral plate of the developing embryo. The intermediate mesoderm develop ...
, immediately under the ectoderm, in the region from the fifth
cervical segment The spinal cord is a long, thin, tubular structure made up of nervous tissue, which extends from the medulla oblongata in the brainstem to the lumbar region of the vertebral column (backbone). The backbone encloses the central canal of the spi ...
to the third
thoracic segment The spinal cord is a long, thin, tubular structure made up of nervous tissue, which extends from the medulla oblongata in the brainstem to the lumbar region of the vertebral column (backbone). The backbone encloses the central canal of the spi ...
, a series of short
evagination Endodermic evagination relates to the inner germ layers of cells of the very early embryo, from which is formed the lining of the digestive tract, of other internal organs, and of certain glands, implies the extension of a layer of body tissue to ...
s from each segment grows
dorsally Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
and extends
caudally Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
, fusing successively from before backward to form the pronephric duct. This continues to grow caudally until it opens into the ventral part of the cloaca; beyond the
pronephros Pronephros is the most basic of the three excretory organs that develop in vertebrates, corresponding to the first stage of kidney development. It is succeeded by the mesonephros, which in fish and amphibians remains as the adult kidney. In amnio ...
it is termed the
Wolffian duct The mesonephric duct (also known as the Wolffian duct, archinephric duct, Leydig's duct or nephric duct) is a paired organ that forms during the embryonic development of humans and other mammals and gives rise to male reproductive organs. Stru ...
. Thus, the Wolffian duct is what remains of the pronephric duct after the atrophy of the pronephros.


Pronephros

The original evaginations form a series of transverse tubules each of which communicates by means of a funnel-shaped ciliated opening with the
abdominal cavity The abdominal cavity is a large body cavity in humans and many other animals that contains many organs. It is a part of the abdominopelvic cavity. It is located below the thoracic cavity, and above the pelvic cavity. Its dome-shaped roof is the th ...
, and in the course of each duct a glomerulus also is developed. A secondary glomerulus is formed ventral to each of these, and the complete group constitutes the pronephros. In humans, the pronephros is just rudimentary, and undergoes rapid atrophy and disappears.


Wolffian duct


Mesonephros

On the medial side of the Wolffian duct, from the sixth cervical to the third lumbar segments, a series of tubules, the Wolffian tubules, develops. They increase in number by outgrowths from the original tubules. They change from solid masses of cells to instead become hollowed in the center. One end grows toward and finally opens into the Wolffian duct, the other dilates and is invaginated by a tuft of capillary bloodvessels to form a glomerulus. The tubules collectively constitute the
mesonephros The mesonephros ( el, middle kidney) is one of three excretory organs that develop in vertebrates. It serves as the main excretory organ of aquatic vertebrates and as a temporary kidney in reptiles, birds, and mammals. The mesonephros is included ...
. The mesonephros persists and form the permanent kidneys in fish and amphibians, but in reptiles, birds, and mammals, it atrophies and for the most part disappears rapidly as the permanent kidney ( metanephros) develops beginning during the sixth or seventh week, so that by the beginning of the fifth month only the ducts and a few of the tubules of the mesonephros remain.


Development in male

In the male the
Wolffian duct The mesonephric duct (also known as the Wolffian duct, archinephric duct, Leydig's duct or nephric duct) is a paired organ that forms during the embryonic development of humans and other mammals and gives rise to male reproductive organs. Stru ...
persists, and forms for example the epididymis, the
ductus deferens The vas deferens or ductus deferens is part of the male reproductive system of many vertebrates. The ducts transport sperm from the epididymis to the ejaculatory ducts in anticipation of ejaculation. The vas deferens is a partially coiled tube ...
, the
ejaculatory duct The ejaculatory ducts (''ductus ejaculatorii'') are paired structures in male anatomy. Each ejaculatory duct is formed by the union of the vas deferens with the duct of the seminal vesicle. They pass through the prostate, and open into the ureth ...
,
seminal vesicle The seminal vesicles (also called vesicular glands, or seminal glands) are a pair of two convoluted tubular glands that lie behind the urinary bladder of some male mammals. They secrete fluid that partly composes the semen. The vesicles are 5†...
and efferent ducts. In the female, on the other hand, the Wolffian bodies and ducts atrophy, leaving behind only remnants in the adult, involving e.g. the
development of the suspensory ligament of the ovary The suspensory ligament of the ovary, also infundibulopelvic ligament (commonly abbreviated IP ligament or simply IP), is a fold of peritoneum that extends out from the ovary to the wall of the pelvis. Some sources consider it a part of the broad ...
.


Müllerian duct

Shortly after the formation of the Wolffian ducts a second pair of ducts is developed; these are the
Müllerian ducts Paramesonephric ducts (or Müllerian ducts) are paired ducts of the embryo that run down the lateral sides of the genital ridge and terminate at the sinus tubercle in the primitive urogenital sinus. In the female, they will develop to form the fal ...
. Each arises on the lateral aspect of the corresponding Wolffian duct as a tubular invagination of the cells lining the abdominal cavity. The orifice of the invagination remains open, and undergoes enlargement and modification to form the abdominal
ostium of the fallopian tube The fallopian tubes, also known as uterine tubes, oviducts or salpinges (singular salpinx), are paired tubes in the human female that stretch from the uterus to the ovaries. The fallopian tubes are part of the female reproductive system. In ot ...
. The ducts pass backward lateral to the Wolffian ducts, but toward the posterior end of the embryo they cross to the medial side of these ducts, and thus come to lie side by side between and behind the latter—the four ducts forming what is termed the ''common genital cord'', to distinguish it from the ''genital cords of the germinal epithelium'' seen later in this article. The Müllerian ducts end in an epithelial elevation, the
Müllerian eminence Sinus tubercle (also known as sinual tubercle or Müllerian eminence) is the proliferation of endoderm induced by the paramesonephric ducts. It is located in the developing fetus between the orifices of the mesonephric ducts on the urogenital s ...
, on the ventral part of the cloaca between the orifices of the Wolffian ducts. At a later stage the eminence opens in the middle, connecting the Müllerian ducts with the cloaca.


Atrophy in males

In the male the Müllerian ducts atrophy, but traces of their anterior ends are represented by the
appendices testis The appendix testis (or hydatid of Morgagni) is a vestigial remnant of the Müllerian duct, present on the upper pole of the testis and attached to the tunica vaginalis. It is present about 90% of the time. Clinical significance Torsion The appe ...
( hydatids of Morgagni of the male), while their terminal fused portions form the utriculus in the floor of the
prostatic urethra The prostatic urethra, the widest and most dilatable part of the urethra canal, is about 3 cm long. It runs almost vertically through the prostate from its base to its apex, lying nearer its anterior than its posterior surface; the form of ...
.


Development in females

In the female the Müllerian ducts persist and undergo further development. The portions which lie in the ''genital cord'' fuse to form the
uterus The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The uter ...
and
vagina In mammals, the vagina is the elastic, muscular part of the female genital tract. In humans, it extends from the vestibule to the cervix. The outer vaginal opening is normally partly covered by a thin layer of mucosal tissue called the hymen ...
. This fusion of the Müllerian ducts begins in the third month, and the septum formed by their fused medial walls disappears from below upward. The parts outside this cord remain separate, and each forms the corresponding
Fallopian tube The fallopian tubes, also known as uterine tubes, oviducts or salpinges (singular salpinx), are paired tubes in the human female that stretch from the uterus to the ovaries. The fallopian tubes are part of the female reproductive system. In ot ...
. The
ostium of the fallopian tube The fallopian tubes, also known as uterine tubes, oviducts or salpinges (singular salpinx), are paired tubes in the human female that stretch from the uterus to the ovaries. The fallopian tubes are part of the female reproductive system. In ot ...
remains from the anterior extremity of the original tubular invagination from the abdominal cavity. About the fifth month a ring-like constriction marks the position of the cervix of the uterus, and after the sixth month the walls of the uterus begin to thicken. For a time the vagina is represented by a solid rod of epithelial cells. A ring-like outgrowth of this epithelium occurs at the lower end of the uterus and marks the future
vaginal fornix The fornices of the vagina (sing. fornix of the vagina or fornix vaginae) are the superior portions of the vagina, extending into the recesses created by the vaginal portion of cervix. The word "fornix" is Latin for "arch". Structure There are t ...
. At about the fifth or sixth month the lumen of the vagina is produced by the breaking down of the central cells of the epithelium. The
hymen The hymen is a thin piece of mucosal tissue that surrounds or partially covers the external vaginal opening. It forms part of the vulva, or external genitalia, and is similar in structure to the vagina. In children, a common appearance of the h ...
represents the remains of the Müllerian eminence.


Metanephros and definitive kidney

The metanephros is the definite, permanent, but yet immature kidney. It arises from two directions. On one hand, the precursor of the ureter buds from the Wolffian duct, while on the other hand, the precursor of the renal tubules develop from the metanephrogenic blastema. The ureteric bud subsequently grows into the latter mass, forming the parts of the nephron. Other changes include e.g. the translocation of the ureteric opening directly into the cloaca.


Ureteric bud

The rudiments of the permanent
kidney The kidneys are two reddish-brown bean-shaped organs found in vertebrates. They are located on the left and right in the retroperitoneal space, and in adult humans are about in length. They receive blood from the paired renal arteries; blood ...
s make their appearance about the end of the first or the beginning of the second month. Each kidney originates as a
ureteric bud The ureteric bud, also known as the metanephric diverticulum, is a protrusion from the mesonephric duct during the development of the urinary and reproductive organs. It later develops into a conduit for urine drainage from the kidneys, which, in ...
from the caudal end of the Wolffian duct, which, in turn, originates from
intermediate mesoderm Intermediate mesoderm or intermediate mesenchyme is a narrow section of the mesoderm (one of the three primary germ layers) located between the paraxial mesoderm and the lateral plate of the developing embryo. The intermediate mesoderm develop ...
. The ureteric bud starts close to where the Wolffian duct opens into the cloaca, and grows dorsalward and rostralward along the posterior abdominal wall, where its blind extremity expands and subsequently divides into several buds, which form the rudiments of the
renal pelvis The renal pelvis or pelvis of the kidney is the funnel-like dilated part of the ureter in the kidney. It is formed by the covnvergence of the major calyces, acting as a funnel for urine flowing from the major calyces to the ureter. It has a mucous ...
and
renal calyces The renal calyces are chambers of the kidney through which urine passes. The minor calyces surround the apex of the renal pyramids. Urine formed in the kidney passes through a renal papilla at the apex into the minor calyx; two or three minor c ...
; by continued growth and subdivision it gives rise to the collecting duct system of the kidney. The other, more superficial, portion of the diverticulum, on the other hand, becomes the ureter.


Metanephrogenic blastema

The
renal corpuscle A renal corpuscle (also called malpighian body) is the blood-filtering component of the nephron of the kidney. It consists of a glomerulus - a tuft of capillaries composed of endothelial cells, and a glomerular capsule known as Bowman's capsul ...
s and
renal tubule The nephron is the minute or microscopic structural and functional unit of the kidney. It is composed of a renal corpuscle and a renal tubule. The renal corpuscle consists of a tuft of capillaries called a glomerulus and a cup-shaped structure ca ...
s, in contrast, are developed from the metanephrogenic blastema instead of from the ureteric bud. The metanephrogenic blastema is moulded over the growing end of the latter, and becomes a part of the metanephros in this way. The renal tubules of the metanephros, unlike those of the pronephros and mesonephros, do not open into the Wolffian duct. Instead, the tubules rapidly elongate to form the parts of the
nephron The nephron is the minute or microscopic structural and functional unit of the kidney. It is composed of a renal corpuscle and a renal tubule. The renal corpuscle consists of a tuft of capillaries called a glomerulus and a cup-shaped structure ...
: the
proximal tubule The proximal tubule is the segment of the nephron in kidneys which begins from the renal pole of the Bowman's capsule to the beginning of loop of Henle. It can be further classified into the proximal convoluted tubule (PCT) and the proximal strai ...
s, the loops of Henle and the
distal convoluted tubule The distal convoluted tubule (DCT) is a portion of kidney nephron between the loop of Henle and the collecting tubule. Physiology It is partly responsible for the regulation of potassium, sodium, calcium, and pH. On its apical surface (lumen ...
s. These last join and establish communications with the collecting duct system derived from the ultimate ramifications of the ureteric bud. In the other end, the renal tubules give rise to
Bowman's capsule Bowman's capsule (or the Bowman capsule, capsula glomeruli, or glomerular capsule) is a cup-like sac at the beginning of the tubular component of a nephron in the mammalian kidney that performs the first step in the filtration of blood to form ur ...
s and
glomeruli ''Glomerulus'' () is a common term used in anatomy to describe globular structures of entwined vessels, fibers, or neurons. ''Glomerulus'' is the diminutive of the Latin ''glomus'', meaning "ball of yarn". ''Glomerulus'' may refer to: * the filter ...
.


Other changes

The mesoderm around the tubules becomes condensed to form the connective tissue of the kidney. The ureter opens at first into the hind-end of the Wolffian duct; after the sixth week it separates from the Wolffian duct, and opens independently into the part of the cloaca which ultimately becomes the
urinary bladder The urinary bladder, or simply bladder, is a hollow organ in humans and other vertebrates that stores urine from the kidneys before disposal by urination. In humans the bladder is a distensible organ that sits on the pelvic floor. Urine enters ...
. The renal tubules become arranged into
renal pyramids The renal medulla is the innermost part of the kidney. The renal medulla is split up into a number of sections, known as the renal pyramids. Blood enters into the kidney via the renal artery, which then splits up to form the segmental arteries whi ...
, and the lobulated condition of the kidneys exists for some time after birth, while traces of it may be found even in the adult. The kidney of the ox and many other animals, on the other hand, remains lobulated throughout life.


Urinary bladder

The
urinary bladder The urinary bladder, or simply bladder, is a hollow organ in humans and other vertebrates that stores urine from the kidneys before disposal by urination. In humans the bladder is a distensible organ that sits on the pelvic floor. Urine enters ...
is formed partly from the endodermal cloaca and partly from the ends of the Wolffian ducts. In other words, the
allantois The allantois (plural ''allantoides'' or ''allantoises'') is a hollow sac-like structure filled with clear fluid that forms part of a developing amniote's conceptus (which consists of all embryonic and extraembryonic tissues). It helps the embryo ...
takes no share in its formation. After the separation of the rectum from the dorsal part of the cloaca, the ventral part becomes the primary urogenital sinus. The urogenital sinus, in turn, divides into the superficial definitive urogenital sinus and the deeper anterior vesico-urethral portion.


Definitive urogenital sinus

The definitive urogenital sinus consists of a caudal phallic portion and an intermediate narrow channel, the pelvic portion.


Vesico-urethral portion

The vesico-urethral portion is the deepest, continuous with the allantois. It absorbs the ends of the Wolffian ducts and the associated ends of the renal diverticula, and these give rise to the
trigone of urinary bladder The trigone (a.k.a. vesical trigone) is a smooth triangular region of the internal urinary bladder formed by the two ureteric orifices and the internal urethral orifice. The area is very sensitive to expansion and once stretched to a certai ...
and part of the
prostatic urethra The prostatic urethra, the widest and most dilatable part of the urethra canal, is about 3 cm long. It runs almost vertically through the prostate from its base to its apex, lying nearer its anterior than its posterior surface; the form of ...
. The remainder of the vesico-urethral portion forms the body of the bladder and part of the prostatic urethra; its apex is prolonged to the umbilicus as a narrow canal, the
urachus The urachus is a fibrous remnant of the allantois, a canal that drains the urinary bladder of the fetus that joins and runs within the umbilical cord. The fibrous remnant lies in the space of Retzius, between the transverse fascia anteriorly and ...
, which later is obliterated and becomes the
median umbilical ligament In human anatomy, the median umbilical ligament is an unpaired midline ligamentous structure upon the lower inner surface of the anterior abdominal wall. It is covered by the median umbilical fold. The median umbilical ligament represents the re ...
of the adult.


Sex organs

Until about the ninth week of gestational age,The University of North Carolina at Chapel Hill - Embryo images nr 024
/ref> the
sex organs A sex organ (or reproductive organ) is any part of an animal or plant that is involved in sexual reproduction. The reproductive organs together constitute the reproductive system. In animals, the testis in the male, and the ovary in the female, a ...
of males and females look the same, and follow a common development. This includes the development of a genital tubercle and a membrane dorsally to it, covering the developing
urogenital opening The urogenital opening is where bodily waste and reproductive fluids are expelled to the environment outside of the body cavity. In some organisms, including birds and many fish, discharge from the urological, digestive, and reproductive system ...
, and the development of labioscrotal folds. Even after differentiation can be seen between the sexes, some stages are common, e.g. the disappearing of the membrane. On the other hand, sex-dependent development include further protrusion of the genital tubercle in the male to form the penis. Furthermore, the labioscrotal folds evolve into the scrotum in males, while they evolve into labia in females.


Diagram of internal differentiation

A.—Diagram of the primitive urogenital organs in the embryo previous to sexual distinction. * 3. Ureter. * 4. Urinary bladder. * 5. Urachus. * cl. Cloaca. * cp. Elevation which becomes clitoris or penis. * i. Lower part of the intestine. * ls. Fold of integument from which the labia majora or scrotum are formed. * m, m. Right and left Müllerian ducts uniting together and running with the Wolffian ducts in gc, the genital cord. * ot. The genital ridge from which either the ovary or testis is formed. * ug. Sinus urogenitalis. * W. Left Wolffian body. * w, w. Right and left Wolffian ducts. B.—Diagram of the female type of sexual organs. * C. Greater vestibular gland, and immediately above it the urethra. * cc. Corpus cavernosum clitoridis. * dG. Remains of the left Wolffian duct, such as give rise to the duct of Gärtner, represented by dotted lines; that of the right side is marked w. * f. The abdominal opening of the left uterine tube. * g. Round ligament, corresponding to gubernaculum. * h. Situation of the hymen. * i. Lower part of the intestine. * l. Labium majus. * n. Labium minus. * o. The left ovary. * po. Epoophoron. * sc. Corpus cavernosum urethrae. * u. Uterus. The uterine tube of the right side is marked m. * v. Vulva. * va. Vagina. * W. Scattered remains of Wolffian tubes near it (paroöphoron of Waldeyer). C.—Diagram of the male type of sexual organs. * C. Bulbo-urethral gland of one side. * cp. Corpora cavernosa penis cut short. * e. Caput epididymis. * g. The gubernaculum. * i. Lower part of the intestine. * m. Müllerian duct, the upper part of which remains as the hydatid of Morgagni; the lower part, represented by a dotted line descending to the prostatic utricle, constitutes the occasionally existing cornu and tube of the uterus masculinus. * pr. The prostate. * s. Scrotum. * sp. Corpus cavernosum urethrae. * t. Testis in the place of its original formation. * t’, together with the dotted lines above, indicates the direction in which the testis and epididymis descend from the abdomen into the scrotum. * vd. Ductus deferens. * vh. Ductus aberrans. * vs. The vesicula seminalis. * W. Scattered remains of the Wolffian body, constituting the organ of Giraldès, or the paradidymis of Waldeyer.


See also

*
Development of the reproductive system The development of the reproductive system is the part of embryonic growth that results in the sex organs and contributes to sexual differentiation. Due to its large overlap with development of the urinary system, the two systems are typically ...


References


In-line


General


Henry Gray (1821–1865). Anatomy of the Human Body. 1918. 3. The Urogenital ApparatusUNSW Embryology - Development of the Kidney and Reproduction SystemsHow the Body Works / Sex Development / Sexual Differentiation / Duct Differentiation
- The Hospital for Sick Children (GTA - Toronto, Ontario, Canada) {{DEFAULTSORT:Development Of The Urinary And Reproductive Organs Embryology of urogenital system