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IN ANATOMY. REPRODUCTIVE SYSTEM - The reproductive system in some parts of its course shares structures in common with the urinary system. In this article the following structures will be dealt with. In the male the testes, epididymis, vasa deferentia, vesiculae seminales, prostate, penis and urethra. In the female the ovaries, Fallopian tubes, uterus, vagina and vulva.
Male Reproductive Organs. The testes or testicles are the glands in which the male reproductive cells are formed. They lie, one on each side, in the scrotum surrounded by the tunica vaginalis (see Coelom and Serous Membranes). Each is an oval gland about one and a half inches long with its long axis directed downward, backward and inward. There is a strong fibrous coat called the tunica albuginea, from which vertical and horizontal septa penetrate into the substance, thus dividing it into compartments or lobules in which the seminiferous tubes are coiled. It is estimated that the total length of these seminiferous tubes in the two glands is little short of a mile. (See fig. I.) At the posterior part of the testis the fibrous sheath is greatly thickened to form the mediastinum testis, and contains a plexus of tubules called the rete testis (see fig. I), into which the seminiferous tubes open. In this way the secretion of the gland is carried to its upper and back part, whence from fifteen to twenty small tubes (vasa efferentia) pass to the epididymis. Each of these is convoluted before opening, and forms what is known as a cones vasculosus. Under the microscope the seminiferous tubules are seen to consist of a basement membrane surrounding several layers of epithelial cells, some of which are constantly being transformed into spermatozoa or male sexual cells.
The epididymis (see fig. i) is a soft body lying behind the testis; it is enlarged above to form the globus major or head, while below is a lesser swelling, the globus minor or tail. The whole epididymis is made up of a convoluted tube about 20 ft. long, from which one long diverticulum (vas aberrans) comes off. Between the globus major and the testis two small vesicles called the hydatids of Morgagni are often found.
The vas deferens is the continuation of the tube of the epididymis and starts at the globus minor; at first it is convoluted, but soon becomes straight, and runs up on the inner (mesial) side of the epididymis to the external abdominal ring in the abdominal wall. On its way up it is joined by several other structures, to form the spermatic cord; these are the artery (spermatic) and veins (pampiniform plexus) of the testis, the artery of the vas, the ilio-inguinal, genito-crural and sympathetic nerves, and the testicular lymphatics. After entering the external abdominal ring, these structures pass obliquely through the abdominal wall, lying in the inguinal canal for an inch and a half, until the internal abdominal ring is reached. Here they separate and the vas passes down the side of the pelvis and turns XXIII. 5 ,??t,t??+,1 3 ?,i - a, ??tQ ' '44,V3"0.131, 104*,1 1 411,141,)S'Y04 vS T 4 ta, k 11, g.
From A. F. Dixon, Cunningham's Text-book of Anatomy. FIG. I. - Diagram to illustrate the structure of the testis and epididymis.
c.v. Coni vasculosi. s.t. Seminiferous c. Globus major. tubule.
g.m'. Globus minor. v.d. Vas deferens.
r. v. Rete testis v.e. Vas efferens.
s. Septula testis. v.r. Tubuli recti.
while the paroOphoron, like the organ of Giraldes in the male, is probably formed from some separate tubes (see fig. 8, Ep. 0. and Par. 0.).
The Wolffian duct, which, in the early embryo, carries the excretion of the mesonephros to the cloaca, forms eventually the body and tail of the epididymis, the vas deferens, and ejaculatory duct in the male, the vesicula seminalis being developed as a pouch in its course. In the female this duct is largely done away with, but remains as the collecting tube of the epoOphoron, and in some mammals as the duct of Gartner, which runs down the side of the vagina to open into the vestibule.
The Miillerian duct, as it approaches the cloaca, joins its fellow of the opposite side, so that there is only one opening into the ventral cloacal wall. In the male the lower part only of it remains as the uterus masculinus (fig. 8, U.M.), but in the female the Fallopian tubes, uterus, and probably the vagina, are all formed from it (fig. 8, F.T. and U.). In both sexes a small hydatid or vesicle is liable to be formed at the beginning of both the Wolffian and Miillerian duct (fig. 8, P.H. and S.H.); in the male these are close together in front of the globus major of the epididymis, and are known as the sessile and pedunculated hydatids of Morgagni. In the female there is a hydatid among the fimbriae of the Fallopian tube which of course is Miillerian and corresponds to the sessile hydatid in the male, while another is often found at the beginning of the collecting tube of the epoOphoron and is probably formed by a blocked mesonephric tubule. This is the pedunculated hydatid of the male. The development of the vagina, as Berry Hart (Journ. Anat. and Phys. xxxv. 330). has pointed out, is peculiar. Instead of the two Miillerian ducts joining to form the lumen of its lower third, as they do in the case of the uterus and its upper two-thirds, they become obliterated, and their place is taken by two solid cords of cells, which Hart thinks are derived from the Wolffian ducts and are therefore probably of ectodermal origin, though this is open to doubt. These cords later become canalized and the septum between them is obliterated.
The common chamber, or cloaca, into which the alimentary, urinary and reproductive tubes open in the foetus, has the urinary bladder (the remains of the allantois) opening from its ventral wall (see Placenta and Urinary System).
During development the alimentary or anal part of the cloaca is separated from the urogenital, and in the article Alimentary System the hitherto accepted method of this separation is described. The question has, however, lately been reinvestigated by F. Wood Jones, who says that the anal part is completely shut off from the urogenital and ends in a blind pouch which grows toward the surface and meets a new ectodermal depression, the main point being that the permanent anus is not, according to him, any part of the original cloacal aperture, but a new perforation. This description is certainly more in harmony with the malformations occurring in this region than the old one, and only awaits confirmatory evidence to be generally accepted.
The external generative organs have at first the same appearance in the two sexes, and consist of a swelling, the genital eminence, in the ventral wall of the cloaca. This in the male becomes the penis and in the female the clitoris. Throughout the generative system the male organs depart most from the undifferentiated type, and in the case of the genital eminence two folds grow together and enclose the urogenital passage, thus making the urethra perforate the penis, while in the female these two folds remain separate as the labia minora or nymphae. Sometimes in the male the folds fail to unite completely, and then there is an opening into the urethra on the under surface of the penis - a condition known as hypospadias.
In the undifferentiated condition the integument surrounding the genital opening is raised into a horseshoelike swelling with its convexity over the pubic symphysis and its concavity toward the anus; the lateral parts of this remain separate in the female and form the labia majora, but in the male they unite to form the scrotum. The median part forms the mons Veneris or mons Jovis.
The Descent of the Testis
It has been shown that the testis is formed in the loin region of the embryo close to the kidney, and it is only in the later months of foetal life that it changes this position for that of the scrotum. In the lower part of the genital ridge a fibro-muscular cord is formed which stretches from the lower part of the testis to the bottom of the scrotum; it is known as the gubernaculum testis, and by its means the testis is directed into the scrotum. Before the testis descends, a pouch of peritoneum called the processus vaginalis passes down in front of the gubernaculum through the opening in the abdominal wall, which afterwards becomes the inguinal canal, into the scrotum, and behind this the testis descends, carrying with it the mesonephros and mesonephric duct. These, as has already been pointed out, form the epididymis and vas deferens. At the sixth month the testis lies opposite the abdominal ring, and at the eighth reaches the bottom of the scrotum and invaginates the processus vaginalis from behind. Soon after birth the communication between that part of the processus vaginalis which now surrounds the testis and the general cavity of the peritoneum disappears, and the part which remains forms the tunica vaginalis. Sometimes the testis fails to pass beyond the inguinal canal, and the term " cryptorchism " is used for such cases.
In the female the ovary undergoes a descent like that of the testis, but it is less marked owing to the fact that the gubernaculum becomes attached to the Miillerian duct where that duct joins its fellow to form the uterus; hence the ovary does not descend lower than the level of the top of the uterus, and the part of the gubernaculum running between it and the uterus remains as the ligament of the ovary, while the part running from the uterus to the labium is the round ligament. In rare cases the ovary may be drawn into the labium just as the testis is drawn into the scrotum.
In the Urochorda, the class to which Salpa, Pyrosoma and the sea squirts (Ascidians) belong, male and female generative glands (gonads) are present in the same individual; they are therefore hermaphrodite.
In the Acrania (Amphioxus) there are some twenty-six pairs of gonads arranged segmentally along the side of the pharynx and intestine and bulging into the atrium. Between them and the atrial wall, however, is a rudimentary remnant of the coelom, through which the spermatozoa or ova (for the sexes are distinct) burst into the atrial cavity. There are no genital ducts.
In the Cyclostomata (lampreys and hags) only one median gonad is found, and its contents (spermatozoa or ova) burst into the coelom and then pass through the genital pores into the urogenital sinus and so to the exterior. It is probable that the single gonad is accounted for by the fact that its fellow has been suppressed.
In the Elasmobranchs or cartilaginous fishes there are usually two testes or two ovaries, though in the dogfish one of the latter is suppressed. From each testis, which in fish is popularly known as the soft roe, vasa efferentia lead into the mesonephros, and the semen is conducted down the vas deferens or mesonephric duct into the urogenital sinus, into which also the ureters open. Sometimes one or more thin-walled diverticula - the sperm sacs - open close to the aperture of the vas deferens. In the female the ova are large, on account of the quantity of yolk, and they burst into the coelum, from which they pass into the large Miillerian ducts or oviducts. In the oviparous forms, such as the common dogfish (Scyllium), there is an oviducal gland which secretes a horny case for the egg after it is fertilized, and these cases have various shapes in different species. Some of the Elasmobranchs, e.g., the spiny dogfish (Acanthias), are viviparous, and in these the lower part of the oviduct is enlarged and acts as a uterus. In male elasmobranchs the anterior part of the Miillerian duct persists. Paired intromittent organs (claspers) are developed on the pelvic fins of the males; these conduct the semen into the cloaca of the female.
In the teleostean and ganoid fishes (Teleostomi) the nephridial ducts are not always used as genital ducts, but special coelomic ducts are formed (see Coelom and Serous Mem Branes) .
In the Dipnoi or mudfish long coiled Miillerian ducts are present, but the testes either pour their secretion directly into the coelom or, as in Protopterus, have ducts which are probably coelomic in origin.
In both the Teleostomi and Dipnoi the testes and ovaries are paired.
True hermaphroditism is known among fishes, the hag (Myxine) and the sea perch (Serranus) being examples. In many others it occurs as an abnormality.
In the Amphibia both ovaries and testes are symmetrical. In the snakelike forms which are found in the order Gymnophiona the testes are a series of separate lobules extending for a long distance, one behind the other, and joined by a connecting duct from which vasa efferentia pass into the Malpighian capsules of the kidneys, and so the sperm is conducted to the mesonephric duct, which acts both as vas deferens and ureter. The Miillerian ducts or oviducts are long and often coiled in Amphibia, and usually open separately into the cloaca. There is no penis, but in certain forms, especially the Gymnophiona, the cloaca is protrusible in the male and acts as an intromittent organ. Corpora adiposa or fat bodies are present in all Amphibians, and probably nourish the sexual cells during the hibernating period.
In Reptilia two testes and ovaries are developed, though they are often asymmetrical in position. In Lizards the vas deferens and ureter open into the cloaca by a common orifice; as they do in the human embryo. In these animals there are two penes, which can be protruded and retracted through the vent; but in the higher reptiles (Chelonia and Crocodilia) there is a single median penis rising from the ventral wall of the cloaca, composed of erectile tissue and deeply grooved on its dorsal surface for the passage of the sperm.
In birds the right ovary and oviduct degenerates, and the left alone iš functional. In the male the ureter and vas deferens open separately into the cloaca, and in the Ratitae (ostriches) and Anseres (ducks and geese) a well-developed penis is present in the male. In the ostrich this is fibrous, and bifurcated at its base, suggesting the crura penis of higher forms.
Among the Mammalia the Monotremata (Ornithorhynchus and Echidna) have bird-like affinities. The left ovary is larger than the right, and the oviducts open separately into the cloaca and do not fuse to form a uterus. The testes retain their abdominal position; and the vasa deferentia open into the base of the penis, which lies in a separate sheath in the ventral wall of the cloaca, and shows an advance on that of the reptiles and birds in that the groove is now converted into a complete tunnel. In the female there is a well-developed clitoris, having the same relations as the penis.
In the marsupials the cloaca is very short, and the vagina and rectum open separately into it. The two uteri open separately and three vaginae are formed, two lateral and one median. The two lateral join together below to form a single median lower vagina, and it is by means of these that the spermatozoa pass up into the oviducts. The upper median vagina at first does not open into the lower one, but during parturition a communication is established which in some animals remains permanent (see J. P. Hill, Proc. Linn. Soc. N.S. Wales, 1899 and 1900). This tripartite arrangement of the upper part of the marsupial vagina is of especial interest in connexion with the views of the embryology of the canal detailed by Berry Hart and already referred to.
When, as in marsuf ials, the two uteri open separately into the vagina by two ora, the arrangement is spoken of as uterus duplex. When the two uteri join below and open by one os externum, it is known as uterus bipartitus. When the uterus bifurcates above and has two horns for the reception of the Fallopian tubes (oviducts), but is otherwise single, the term uterus bicornis is given to it, while the 'single uterus of man and other Primates is called uterus simplex. From the marsupials upward the ovarian end of the Fallopian tube has the characteristic fimbriated appearance noticed in human anatomy.
In some mammals, such as the sow and the cow, the Wolffian duct is persistent in the female and runs along the side of the vagina as the duct of Gartner. It is possible that the lateral vaginae of the marsupials are of Wolffian origin.
In marsupials the testes descend into the scrotum, which lies in these animals in front of instead of behind the penis. In some mammals, such as the elephant, they never reach the scrotum at all; while in others, e.g. many rodents, they can be drawn up into the abdomen or lowered into the scrotum. The subject of the descent of the testicles has been very fully treated by H. Klaatsche, " Ueber den Descensus testiculorum," Morph. Jahrb., Bd. xvi.
The prostate is met with in its most simple forms in marsupials, in which it is a mere thickening of the mucous membrane of the urethra; in the sheep it forms a bilateral elongated mass of gland tissue lying behind the urethra and surrounded by a welldeveloped layer of striped muscle. In the sloth it is said to be altogether absent, while in many of the insectivores and rodents it consists of many lobes which usually show a bilateral arrangement. The vesiculae seminales are usually present in the Eutheria or higher mammals, and sometimes, as in the hedgehog, are very large, though they are absent in the Carnivora. Cowper's glands are usually present and functional throughout From C. S. Wallace's Prostatic Enlargement. FIG. 9. - Transverse Section of Sheep's Prostate.
life. The uterus masculinus is also usually present, but there is grave doubt whether the large organ called by this name in the rabbit should not rather be regarded as homologous with part of the vesiculae seminales. The penis shows many diversities of arrangement; above the marsupials its two crura obtain an attachment to the ischium. In many mammals it is quite hidden by the skin in the flaccid condition, and its external orifice may range from the perineum in the marsupials to the middle of the ventral wall of the abdomen in the ruminants. In the Marsupialia, Rodentia, Chiroptera, Carnivora and some Primates an os penis is developed in connexion with the corpora cavernosa.
The clitoris is present in all mammals; sometimes, as in the female hyena, it is very large, and at others, as in the lemur, it is perforated by the urethra.
For further details and literature, see Oppel's Lehrbuch der vergleich. mikroskop. Anatomie der Wirbelthiere, Bd. iv. (Jena, 1904); also Gegenbaur's Vergleich. Anat. der Wirbelthiere, and Wiedersheim's Comparative Anatomy of Vertebrates, translated by W. N. Parker (London, 1907). (F. G. P.)
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