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Gilbert SF. Developmental Biology. 6th edition. Sunderland (MA): Sinauer Associates; 2000.

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Developmental Biology. 6th edition.

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Spermatogenesis

While the reductive divisions of meiosis are conserved in every eukaryotic kingdom of life, the regulation of meiosis in mammals differs dramatically between males and females. The differences between oogenesis, the production of eggs, and spermatogenesis, the production of sperm, are outlined in Table 19.1.

Table 19.1. Sexual dimorphism in mammalian meioses.

Table 19.1

Sexual dimorphism in mammalian meioses.

Spermatogenesis is the production of sperm from the primordial germ cells. Once the vertebrate PGCs arrive at the genital ridge of a male embryo, they become incorporated into the sex cords. They remain there until maturity, at which time the sex cords hollow out to form the seminiferous tubules, and the epithelium of the tubules differentiates into the Sertoli cells. The initiation of spermatogenesis during puberty is probably regulated by the synthesis of BMP8B by the spermatogenic germ cells, the spermatogonia. When BMP8B reaches a critical concentration, the germ cells begin to differentiate. The differentiating cells produce high levels of BMP8B, which can then further stimulate their differentiation. Mice lacking BMP8B do not initiate spermatogenesis at puberty (Zhao et al. 1996).

The spermatogenic germ cells are bound to the Sertoli cells by N-cadherin molecules on both cell surfaces and by galactosyltransferase molecules on the spermatogenic cells that bind a carbohydrate receptor on the Sertoli cells (Newton et al. 1993; Pratt et al. 1993). The Sertoli cells nourish and protect the developing sperm cells, and spermatogenesis—the developmental pathway from germ cell to mature sperm—occurs in the recesses of the Sertoli cells (Figure 19.17). The processes by which the PGCs generate sperm have been studied in detail in several organisms, but we will focus here on spermatogenesis in mammals.

Figure 19.17. Drawing of a section of the seminiferous tubule, showing the relationship between Sertoli cells and the developing sperm.

Figure 19.17

Drawing of a section of the seminiferous tubule, showing the relationship between Sertoli cells and the developing sperm. As cells mature, they progress toward the lumen of the seminiferous tubule. (After Dym 1977.)

After reaching the gonad, the PGCs divide to form type A1 spermatogonia. These cells are smaller than the PGCs and are characterized by an ovoid nucleus that contains chromatin associated with the nuclear membrane. The A1 spermatogonia are found adjacent to the outer basement membrane of the sex cords. They are stem cells, and at maturity, they are thought to divide so as to make another type A1 spermatogonium as well as a second, paler type of cell, the type A2 spermatogonium. Thus, each type A1 spermatogonium is a stem cell capable of regenerating itself as well as producing a new cell type. The A2 spermatogonia divide to produce the A3 spermatogonia, which then beget the type A4 spermatogonia. It is possible that each of the type A spermatogonia are stem cells, capable of self-renewal. The A4 spermatogonium has three options: it can form another A4 spermatogonium (self-renewal); it can undergo cell death (apoptosis); or it can differentiate into the first committed stem cell type, the intermediate spermatogonium. Intermediate spermatogonia are committed to becoming spermatozoa, and they divide mitotically once to form the type B spermatogonia. These cells are the precursors of the spermatocytes and are the last cells of the line that undergo mitosis. They divide once to generate the primary spermatocytes—the cells that enter meiosis. It is not known what causes the spermatogonia to take the path toward differentiation rather than self-renewal; nor is it known what stimulates the cells to enter meiotic rather than mitotic division (Dym 1994).

Looking at Figures 19.17 and 19.18, we find that during the spermatogonial divisions, cytokinesis is not complete. Rather, the cells form a syncytium whereby each cell communicates with the others via cytoplasmic bridges about 1 μm in diameter (Dym and Fawcett 1971). The successive divisions produce clones of interconnected cells, and because ions and molecules readily pass through these intercellular bridges, each cohort matures synchronously. During this time, the spermatocyte nucleus often transcribes genes whose products will be used later to form the axoneme and acrosome.

Figure 19.18. The formation of syncytial clones of human male germ cells.

Figure 19.18

The formation of syncytial clones of human male germ cells. (After Bloom and Fawcett 1975.)

Each primary spermatocyte undergoes the first meiotic division to yield a pair of secondary spermatocytes, which complete the second division of meiosis. The haploid cells thus formed are called spermatids, and they are still connected to one another through their cytoplasmic bridges. The spermatids that are connected in this manner have haploid nuclei, but are functionally diploid, since a gene product made in one cell can readily diffuse into the cytoplasm of its neighbors (Braun et al. 1989). During the divisions from type A1 spermatogonium to spermatid, the cells move farther and farther away from the basement membrane of the seminiferous tubule and closer to its lumen (see Figure 19.17). Thus, each type of cell can be found in a particular layer of the tubule. The spermatids are located at the border of the lumen, and here they lose their cytoplasmic connections and differentiate into sperm cells. In humans, the progression from spermatogonial stem cell to mature sperm takes 65 days (Dym 1994).

WEBSITE

19.6 Gonial syncytia: Bridges to the future. The products of meiotic divisions are connected by cytoplasmic connections. The functions of these connections may differ between those cells producing sperm and those producing eggs. http://www.devbio.com/chap19/link1906.shtml

Spermiogenesis

The mammalian haploid spermatid is a round, unflagellated cell that looks nothing like the mature vertebrate sperm. The next step in sperm maturation, then, is spermiogenesis (or spermateliosis), the differentiation of the sperm cell. For fertilization to occur, the sperm has to meet and bind with the egg, and spermiogenesis prepares the sperm for these functions of motility and interaction. The processes of mammalian sperm differentiation is shown in Figure 7.2. The first steps involve the construction of the acrosomal vesicle from the Golgi apparatus. The acrosome forms a cap that covers the sperm nucleus. As the acrosomal cap is formed, the nucleus rotates so that the cap will be facing the basal membrane of the seminiferous tubule. This rotation is necessary because the flagellum is beginning to form from the centriole on the other side of the nucleus, and this flagellum will extend into the lumen. During the last stage of spermiogenesis, the nucleus flattens and condenses, the remaining cytoplasm (the “cytoplasmic droplet”) is jettisoned, and the mitochondria form a ring around the base of the flagellum.

One of the major changes in the nucleus is the replacement of the histones by protamines. Transcription of the gene for protamine is seen in the early haploid cells (spermatids), although translation is delayed for several days (Peschon et al. 1987). Protamines are relatively small proteins that are over 60% arginine. During spermiogenesis, the nucleosomes dissociate, and the histones of the haploid nucleus are eventually replaced by protamines. This causes the complete shutdown of transcription in the nucleus and facilitates its assuming an almost crystalline structure. The resulting sperm then enter the lumen of the tubule.

In the mouse, the entire development process from stem cell to spermatozoon takes 34.5 days. The spermatogonial stages last 8 days, meiosis lasts 13 days, and spermiogenesis takes up another 13.5 days. In humans, spermatic development takes nearly twice as long to complete. Because the type A1 spermatogonia are stem cells, spermatogenesis can occur continuously. Each day, some 100 million sperm are made in each human testicle, and each ejaculation releases 200 million sperm. Unused sperm are either resorbed or passed out of the body in urine. During his lifetime, a human male can produce 1012 to 1013 sperm (Reijo et al. 1995).

VADE MECUM

Spermatogenesis in mammals. The development of sperm is visualized with color-coded histological sections through a mammalian testis. Each stage is shown, from spermatogonium to flagellated spermatid.[Click on Gametogenesis]

WEBSITE

19.7 Gene expression during spermatogenesis. Transcription occurs both from the diploid spermatocyte nucleus and from the haploid spermatid nuclei. Posttranscriptional control is also important in regulating sperm gene expression. http://www.devbio.com/chap19/link1907.shtml

WEBSITE

19.8 The Nebenkern. Sperm mitochondria are often highly modified to fit the streamlined cell. The mitochondria of flies fuse together to form a structure called the Nebenkern, and this fusion is controlled by the fuzzy onions gene. http://www.devbio.com/chap19/link1908.shtml

By agreement with the publisher, this book is accessible by the search feature, but cannot be browsed.

Copyright © 2000, Sinauer Associates.
Bookshelf ID: NBK10095

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