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surface of the hymen is directly continuous with the vaginal wall, and on it are to be seen slight ridges continuous with the vaginal rugæ.

Developmentally the hymen appears to be a portion of the vagina.

On each side of the vaginal opening, and close against the medial side of the attached margin of the labium minus, is the minute opening of the duct of the glandula vestibularis major (O.T. Bartholin's gland). This is usually just large enough to be visible to the unaided eye.

Numerous minute mucous glands, the glandulæ vestibulares minores, open on the surface of the mucous membrane of the vestibule, between the urethral and vaginal orifices. The opening of the ductus para-urethralis at the side of the urethral orifice has been already noted,

p. 1285.

Clitoris.—The clitoris is the morphological equivalent of the penis, and is composed of a body and two crura. Upon the summit of the body is a minute glans. Unlike the penis, the clitoris is not traversed by the urethra.

The corpus clitoridis is composed for the most part of erectile tissue resembling that of the penis in the male. It is about an inch or an inch and a half in length, and is bent upon itself, forming an angle open downwards. The body of the clitoris tapers towards its distal end, which is covered by the glans clitoridis. The organ is enclosed in a dense fibrous coat, and is divided by an incomplete septum, the septum corporum cavernosorum, into two symmetrical and somewhat cylindrical portions, the corpora cavernosa clitoridis. These represent the corpora cavernosa penis of the male, and diverge from one another at the root of the clitoris to form the crura clitoridis. A ligamentum suspensorium clitoridis passes from the fibrous coat of the body of the clitoris to the symphysis pubis (Fig. 1036)

The glans clitoridis is a small mass of erectile tissue which is fitted over the pointed end of the body. It possesses, like the glans penis, which it represents, a very sensitive epithelium. The præputium, or fold of skin which covers it, and the frenulum clitoridis, which is attached to it inferiorly, are continuous with the labia minora (Fig. 1036).

The crura clitoridis diverge from the body posteriorly, and are attached to the sides of the pubic arch. Each is continuous with one of the corpora cavernosa, and has a firm fibrous sheath, which is covered by the corresponding ischio-cavernosus or erector clitoridis muscle. In structure the crura and body of the clitoris resemble the corpora cavernosa penis, while the glans more closely resembles the bulbus vestibuli, with which it is continuous through a structure known as the pars intermedia.

In the seal and some other animals, a bone, which represents the os penis of the male, is developed in the septum of the clitoris. This bone receives the name os clitoridis.

Arteries and Nerves of the Clitoris.—Each crus receives a branch, the arteria profunda clitoridis, from the internal pudendal, while the glans is supplied by branches of the arteria dorsalis clitoridis.

The nerve-supply of the clitoris is derived partly from the hypogastric sympathetic plexus and partly from the dorsal norves of the clitoris, which are branches of the pudendal nerves.

Bulbus Vestibuli.—The bulbus vestibuli is a mass of erectile tissue, in the female, which corresponds developmentally to the corpus cavernosum urethræ of the male. In the female the fusion of the two halves of this structure is not nearly so complete as in the male, for the vagina and urethra separate the bulbus vestibuli into a right and a left portion which are only slightly connected in front by a narrow median part called the pars intermedia. Each half of the bulb is thick and massive posteriorly, and more pointed in front, where it joins the pars intermedia. It rests against the lateral wall of the vagina, and upon the superficial aspect of the fascia inferior of the urogenital diaphragm. It represents onehalf of the corpus cavernosum urethræ of the male. Superficially it is covered by the bulbo-cavernosus muscle. The pars intermedia lies above the opening of the urethra, and becomes continuous with the tissue of the glans clitoridis.

The bulbus vestibuli is for the most part composed of minute convoluted bloodvessels, held together by a very small amount of connective tissue. These vessels frequently anastomose with one another, and those of each half communicate with the vessels of the pars intermedia and the glans clitoridis.

ply of the bulb is derived, on each side, from the arteria bulbi vestibuli, nternal pudendal.

GLANDULA VESTIBULARES MAJORES. er vestibular glands (O.T. glands of Bartholin) are placed one on e lower part of the vagina, and represent the bulbo-urethral glands in




Larger vestibular glands




e. They are often overlapped by the posterior ends of the bulbus vestibuli, covered by the bulbo-cavernosus muscle. Each is about the size and of a small bean, and possesses a long slender duct which opens into the udendi in the angle between the attached border of the labium minus and the 1 opening.



General Account. - In tracing the developmental history of the uro-genital ages we may for convenience begin with an embryo of fifteen days old. About this - a duct, which runs in a longitudinal direction, and occupies a position on the lateral

of the protovertebral somites, begins to develop on each side of the body. With the eption of the anterior portion of the cloaca and the proximal part of the allantois, this ct, which has received the name of primary excretory or Wo an duct, is the earliest med structure from which, or in connexion with which, the parts of the adult ino-genital system arise.

The Wolffian duct serves as the canal, or duct, for the primitive secretory organsle pronephros and the mesonephros of the embryo. With the atrophy of these the uct suffers modification, yet both sexes in the adult possess structures which have heir embryonic origin from the Wolffian duct. In the male the duct of the epididymis, Che ductus deferens, and the ejaculatory duct, are to be looked upon as directly developed from the Wolffian duct of the embryo; while in the female the longitudinal duct of the ep-oöphoron and the appendices vesiculosi are rudimentary structures having a like origin. Further, the ureter and its pelvis arise in both sexes as an outgrowth from the Wolfian duct (Fig. 1037). In the male the vesicula seminalis is developed as a diverticulum of the Wolffian duct.

The primitive secretory organs, the pronephros and the mesonephros, develop in connexion with the anterior part of the Wolffian duct (p. 48), and, during the early life of the embryo, the latter of these is a most important structure. Even in the embryo






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the pronephros is a vestigial organ, and its development in all higher vertebrates is very incomplete. It disappears almost as soon as it is formed, and it is replaced by the far more important mesonephros. With the development of the permanent kidney the mesonephros atrophies, yet some of its tubules persist in the adult. The ductuli

efferentes, the ductuli aberrantes, and the rudimentary paradidymis (organ of Giraldés) in the male, and the rudimentary tubules of the ep-oöphoron and of the par-oöphoron in the female, are structures which one their origin to the tubules of the mesonephros.

Soon after the formation of the Wolffian ducts two other longitudinally disposed canals, called the Müllerian ducts, are developed. These open at their cephalic ends into the body cavity, and at their caudal ends, unlike the Wolffian ducts, they unite with one another in the median plane. From them are formed, in the female—the uterine tubes, the uterus, and the vagina ; and in the male—the appendices of the testis and the utriculus prostaticus.

The Wolffian and Müllerian ducts open at their caudal ends into the ventral or urogenital part of the cloaca, which in the course of development becomes transformed into the bladder and the uro-genital canal of the embryo. The developing ureter at first

arises as a diverticulum from the Wolffian duct, at a U

short distance from the point where the latter joins

the cloaca. Soon, however, the ureters acquire inde CM

pendent openings into the cloaca, which become gradu. ally shifted further from one another and from those of the Wolffian ducts. The ureters are now found to open into the anterior portion of the cloaca which lies nearer to the head of the embryo than the part with which the Wolffian ducts are connected. This cephalic portion of the anterior subdivision of the cloaca which receives the ureters becomes the bladder and the upper part of the urethra.

The caudal part, lying

below the level of the entrance of the Wolffian ducts III

is called the uro-genital canal, and is represented in the adult male by the lower part of the prostatic and by the membranous portions of the urethra ; in the

female by the lower part of the urethra and the part of Fig. 1037.-DIAGRAM TO ILLUSTRATE THE

the uro-genital fissure which immediately surrounds MANNER IN WHICH THE URETER, THE the openings of the urethra and vagina (Figs. 1044Ductus DEFERENS, AND THE URINARY 1045). The united Müllerian ducts open into the lower BLADDER ARISE IN THE EMBRYO.

part of the cloaca or uro-genital canal between the The structures developed from the cloaca Wolffian ducts of opposite sides. In the male the

are indicated in blue, those from the position of this opening, which is represented in the Wolffian duct in red, and the ectoderm adult by the orifice of the utriculus prostaticus, remains in black.

almost unchanged ; in the female, on the other hand, The manner in which the rectum and

bladder become separated and acquire a downgrowth from the fused Müllerian ducts gives openings into the ectodermal cloacal origin to a new passage, the vagina, which establishes fossa is shown in II. and III. (A. H. an opening behind that of the urethra in the uro Young and A. Robinson).

genital fissure of the adult.

After the complete separation of the cloaca into Ur. Uro-genital canal.

anterior or uro-genital and posterior or rectal sub

divisions, the rectum establishes a communication with Pelvis of kidney. WD. Wolffian duct. the exterior in the floor of the shallow depression

known as the ectodermal cloacal fossa. At a little later time the uro-genital canal also joins this fossa at a point in front of the opening of the rectum. The ectodermal cloacal fossa lies in front of the vestigial tail, and extend forwards as far as a tubercle known as the cloacal tubercle, which later gives rise to the genital eminence and a pair of elevations called the labio-scrotal folds. The genital eminence becomes converted into the clitoris or penis according to the sex. The labioscrotal folds extending backwards on each side form the labia majora of the female, and,







A. Allantois.
B. Bladder.
C. Cloaca.
CM. Ectoderm of

cloacal fossa. K.

U. Ureter.
VD. Ductus deferens.
VS. Seminal vesicle.

y, give rise to the scrotum in the male. In the female the slit-like ro-genital canal retains its position, and its margins becoming elongated, Linora. The vaginal opening arises, as we have seen, by a downgrowth lerian ducts which lies in front of the rectum and behind the primitive

The latter becomes the urethra in the female. In the male the slithe uro-genital canal is prolonged anteriorly by an active growth at the al eminence, and its margins uniting, give rise to the penile portion of


Neural tube


Wolftian duct


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Umbilical vein

Iffian Duct.—The Wolffian duct arises in the mesoderm, about the as a solid cord of cells occupying a position immediately to the lateral side ertebral somites and to the medial side of the body cavity (Fig. 1038). When d the duct lies immediately beneath the ectoderm, and as it grows backwards cloaca it is often

intimately conthe ectoderm.

Notochord connexion of the

Muscle plate the ectoderm, in tages, is by some supposed to in

Mesentery imitive ectodermal

Mesonephric the canal, but by ad apparently with ason, to be a trace opening of ducts on ace of the body, such s in connexion with -xcretory organs of

animals. During hird week the cellular

which represents the Gan duct acquires a

Alimentary canal Body cavity en, and about the end he same week the duct Fig. 1038.— TRANSVERSE SECTION THROUGH THE TRUNK OF A HUMAN ets growth reaches the


as the aca has become divided into dorsal and ventral subdivisions, the Wolffian duct is en to end in the caudal part of the ventral subdivision, which becomes the bladder nd uro-genital canal (Fig. 1040).

The mesonephros or Wolffian body is developed in the mesoderm of the “intermediate ell mass," immediately adjoining the Wolffian duct, and consists of a number of transversely arranged canals or tubules, each of which opens by one end into the Wolffian duct, while its other extremity ends blindly. These transverse tubules, like the canal into which they open, are at first solid cellular structures, and only later acquire a distinct lumen. Increasing rapidly in size and number, the tubules become twisted and tortuous, and the blind end of each dilates to form a capsule invaginated upon itself and containing a bunch of capillary blood-vessels similar to the glomeruli of the adult kidney. It would appear that primitively one tubule is developed in the portion of the intermediate cell mass (nephrotome) corresponding to each mesodermic somite, but, in higher vertebrates at all events, such a correspondence between the number of somites and the number of tubules cannot be demonstrated. In the posterior part of the mesonephros the number of tubules is very numerous, and greater than the number of segments in this region. The tubules of the mesonephros arise in all segments from the sixth cervica] to the third lumbar. The tubules in the anterior part atrophy and disappear at a very early time, even while others are being formed towards the caudal end of the embryo. When at its greatest development (fifth to eighth week) the mesonephros forms a relatively large glandular mass, composed of tubules resembling in a general way those of the adult kidney, which projects into the dorsal part of the body cavity, and extends from the region of the liver to the caudal end of the body cavity. Along its lateral aspect lies the Wolffian duct.

As soon

In anamniate vertebrates, fishes, and amphibia, the mesonephros persists as the secretory organ of the adult.


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Neural tube

Post anal gut



THE TIME AT WHICH THE WOLFFIAN DUCTS REACH THE CLOACA. (Drawn from a model constructed by Prof. F. Keibel.)


Wolffian ducts open into the anterior part of the cloaca. (Drawn from a model constructed by Prof. F. Keibel.)

Pronephros.-From what is known regarding the development of lower animals, it seems certain that the Wolffian duct originally served as the duct of the still earlier secretory organ

Body of vertebra

Spinal medulla
Spinal ganglion

Arch of vertebra

Ilium (cartilage).

Wolffian duct

Body cavity

Müllerian duct

Genital gland


ABOUT 7 WEEKS. (Specimen in the possession of Prof. J. Symington.)

the Pronephros. In man the pronephros arises nearer to the head end of the embryo than the later formed mesonephros, and its tubules can only with difficulty be distinguished

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