DIFFERENTIATION OF THE GONADS

After the Migration of Germ Cells from the Yolk Sac, the Undifferentiated Primordial Gonad Develops into Either a Testis or an Ovary

The primordial germ cells do not originate in the gonad; instead, they migrate to the gonad from the yolk sac along the mesentery of the hindgut at about the fifth week of embryo development (Fig. 52-3A, B). The primordial germ cells of humans are first found in the endodermal epithelium of the yolk sac in the vicinity of the allantoic stalk, and from there the germ cells migrate into the adjoining mesenchyme. They eventually take up their position embedded in the gonadal ridges. Gonadal development fails to progress normally in the absence of germ cells. Thus, any event that interferes with germ cell migration may cause abnormal gonadal differentiation. The gonad forms from a portion of the coelomic epithelium, the underlying mesenchyme, and the primordial germ cells that migrate from the yolk sac. At 5 weeks' development, a thickened area of coelomic epithelium develops on the medial aspect of the urogenital ridge as a result of proliferation of both the coelomic epithelium and cells of the underlying mesenchyme. This prominence, which forms on the medial aspect of the mesonephros, is known as the gonadal ridge (see Fig. 52-3B, C). Migration of the primordial germ cells to the gonadal ridge establishes the "anlagen" for the primordial gonad. The primordial gonad at this early stage of development consists of both a peripheral cortex and a central medulla (see Fig. 52-3C) and has the capacity to develop into either an ovary or a testis. As discussed later, the cortex and medulla have different fates in the male and female. The germ cells themselves seem to direct the sexual development of the gonad. An embryo with an XY chromosome complement undergoes development of the medullary portion of the gonad to become a testis and the cortex regresses. On the other hand, XX germ cells appear to stimulate development of the cortex of the early gonad to become an ovary and the medulla regresses. DEVELOPMENT OF THE PRIMITIVE TESTIS. In male embryos, primordial germ cells migrate from the cortex of the gonad, in which they were originally embedded, into the primitive sex cords of the medulla (see Fig. 52-3D). The primitive sex cords become hollowed out and develop into the seminiferous tubules. The primordial germ cells give rise to spermatogonia, the first cells in the pathway to mature sperm (p. 1131). The sex cords give rise to Sertoli cells. The rete testis is a system of thin, interconnected tubules that develop in the dorsal part of the gonad; they drain the seminiferous tubules. The contents of the rete testes flow into the efferent ductules, which-as we will see later-develop from the adjoining tubules of the mesonephros. These tubular structures establish a pathway from the male gonad to the mesonephric duct, which-as we will also see later-evolves into the outlet for sperm. The cortex of the primordial gonad is a thin epithelial layer covering the coelomic surface of the testis. DEVELOPMENT OF THE PRIMITIVE OVARY.
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Figure 52-3 The early gonad and germ cell migration. A, The primordial germ cells originate in the endodermal endothelium of the yolk sac. B, The primordial germ cells migrate along the mesentery of the hindgut and reach the region of the urogenital ridge called the gonadal ridge. C, The indifferent gonad consists of an outer cortex and an inner medulla. D, The testis develops from the medulla of the indifferent gonad; the cortex regresses. E, The ovary develops from the cortex of the indifferent gonad; the medulla regresses.
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In female embryos, the medulla of the gonad regresses, the primary sex cords are resorbed, and the

interior of the gonad is filled with a loose mesenchyme that is highly permeated by blood vessels. However, the cortex greatly increases in thickness, with the primordial germ cells remaining embedded within (see Fig. 52-3E). Masses of cortical cells are split up on the inner surface of the cortex into groups and strands of cells, or secondary sex cords, surrounding one or several primordial germ cells, or oogonia, during growth of the gonad. These germ cells become primary oocytes that enter the initial stages of oogenesis.
The Embryonic Gonad Has Local Effects on Development of the Internal Genitalia and Also Determines the External Sexual Phenotype

As discussed in the following subchapter, several products of the developing male or female gonad have profound effects on differentiation of the internal sex ducts, as well as on development of the external genitalia. Thus, just as genetic sex determines the gonadal phenotype, products of the gonad primarily determine the sexual phenotype. Androgens produced by the developing testis cause development of the mesonephric or wolffian ducts. The para-mesonephric or mllerian ducts degenerate in the male under the influence of antimllerian hormone (AMH). In the female embryo, the mllerian ducts develop whereas the wolffian ducts degenerate. In the absence of a functioning testis, the left and right mllerian ducts develop according to the female phenotype, that is, as the fallo-pian tubes (oviducts), the uterus, and the upper third of the vagina (see the next section). Just as the absence of male hormones or androgens causes the internal genital ducts to follow a female pattern of differentiation, the absence of androgens causes the external genital development to be female. Conversely, testosterone and dihydrotestosterone (DHT) cause masculinization of the external genitalia (see the subchapter titled Differentiation of the External Genitalia).
Printed from STUDENT CONSULT: Medical Physiology (on 28 August 2006) 2006 Elsevier