Development of the gonads

During early embryonic development, cells from the dorsal endoderm of the yolk sac migrate along the hindgut to the gonadal ridge. These primordial germ cells (PGCs) multiply by mitosis and once they have reached the gonadal ridge they are called oogonia (diploid stem cells of the ovary).

Once oogonia enter this area they attempt to associate with the other somatic cells, derived from both the peritoneum and mesonephros. Development proceeds and the oogonia become fully surrounded by a layer of connective tissue cells (pre-granulosa cells) in an irregular manner. In this way, the rudiments of the ovarian follicles are formed.

The embryological origin of granulosa cells remains controversial. In the 1970s, evidence emerged that the first cells to make contact with the oogonia were of mesonephric origin. It was suggested that mesonephric cells already closely associated with the oogonia proliferated throughout development to form the granulosa cell layer.[1][2][3]

Recently this hypothesis has been challenged with some thorough histology. Sawyer et al. hypothesized that in sheep most of the granulosa cells develop from cells of the mesothelium (i.e., epithelial cells from the presumptive surface epithelium of the ovary).[4]

Just as in the male, there is a gubernaculum in the female, which effects a considerable change in the position of the ovary, though not so extensive a change as in that of the testis. The gubernaculum in the female lies in contact with the fundus of the uterus and adheres to this organ, and thus the ovary can only descend as far as to this level. The part of the gubernaculum between the ovary and the uterus ultimately becomes the proper ovarian ligament, while the part between the uterus and the labium majus forms the round ligament of the uterus. A pouch of peritoneum analogous to the vaginal process in the male accompanies it along the inguinal canal: it is called the canal of Nuck.

In rare cases, the gubernaculum may fail to contract adhesions to the uterus, and then the ovary descends through the inguinal canal into the labium majus; under these circumstances, its position resembles that of the testis.

The descent of the testes consists of the opening of a connection from the testis to its final location at the anterior abdominal wall, followed by the development of the gubernaculum, which subsequently pulls and translocates the testis down into the developing scrotum. Ultimately, the passageway closes behind the testis.

At an early period of fetal life the testes are placed at the back part of abdominal cavity, behind the peritoneum, and each is attached by a peritoneal fold, the mesorchium, to the mesonephros. From the front of the mesonephros a fold of peritoneum termed the inguinal fold grows forward to meet and fuse with a peritoneal fold, the inguinal crest, which grows backward from the antero-lateral abdominal wall. The testis thus acquires an indirect connection with the anterior abdominal wall. At the same time, a portion of the peritoneal cavity lateral to these fused folds is marked off as the future vaginal process.

Also, in the inguinal crest a structure, the gubernaculum testis, makes its appearance. This is at first a slender band, extending from that part of the skin of the groin which afterward forms the scrotum through the inguinal canal to the body and epididymis of the testis. As and contains the upper part of the gubernaculum, and ultimately also the internal spermatic vessels; the one below, the plica gubernatrix, contains the lower part of the gubernaculum.

The gubernaculum grows into a thick cord. It ends below at the abdominal inguinal ring in a tube of peritoneum, the vaginal process, which protrudes itself down the inguinal canal. By the fifth month the lower part of the gubernaculum still is a thick cord, while the upper part has disappeared. The lower part now consists of a central core of smooth muscle fibers, surrounded by a firm layer of striated muscle elements, connected, behind the peritoneum, with the abdominal wall.

As the testes develops, the main portion of the lower end of the gubernaculum is carried, following the skin to which it is attached, to the bottom of this pouch. Other bands are carried to the medial side of the thigh and to the perineum. The tube of peritoneum constituting the vaginal process projects itself downward into the inguinal canal, and emerges at the external inguinal ring, pushing before it a part of the obliquus internus and the aponeurosis of the obliquus externus, which form respectively the cremaster muscle and the external spermatic fascia. The vaginal process forms a gradually elongating pouch, which eventually reaches the bottom of the scrotum, and behind this pouch the testis is drawn by the growth of the body of the fetus, for the gubernaculum does not grow proportionately with the growth of other parts, and therefore the testis, being attached by the gubernaculum to the bottom of the scrotum, is prevented from rising as the body grows, and is instead drawn first into the inguinal canal and eventually into the scrotum. It seems certain also that the gubernacular cord becomes shortened as development proceeds, and this assists in causing the testis to reach the bottom of the scrotum.

Diagram of an indirect, scrotal inguinal hernia (median view from the left).

By the end of the eighth month the testis has reached the scrotum, preceded by the vaginal process, which communicates by its upper extremity with the peritoneal cavity. Just before birth the upper part of the vaginal process, at the internal inguinal ring, usually becomes closed, and this obliteration extends gradually downward to within a short distance of the testis. The process of peritoneum surrounding the testis is now entirely cut off from the general peritoneal cavity and constitutes the tunica vaginalis.

If the internal inguinal ring does not close properly, then there is a risk that other contents of the abdominal cavity protrudes through the passageway and cause indirect inguinal hernia.