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Functional Anatomy of the Hypothalamic–Pituitary–Gonadal Axis and 1 the Male Reproductive Tract

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Functional Anatomy of the Hypothalamic–Pituitary–Gonadal Axis and 1 the Male Reproductive Tract Cambridge University Press 978-1-107-01212-7 - Fertility Preservation in Male Cancer Patients Editor-in-Chief John P. Mulhall Excerpt More information Section 1 Anatomy and physiology Chapter Functional anatomy of the hypothalamic–pituitary–gonadal axis and 1 the male reproductive tract Nelson E. Bennett Jr. Anatomy of reproductive function The reproductive functional axis of the male can be divided into three major subdivisions: (1) the hypo- thalamus, (2) the pituitary gland, and (3) the testis. Each level elaborates a signal, or transmitter molecule, that stimulates or inhibits the subsequent level of the axis. The end result is the production and expulsion of semen that contains spermatozoa. This chapter exam- ines the hypothalamic–pituitary–gonadal (HPG) axis, and reviews the functional anatomy of the testis, epi- didymis, vas deferens, seminal vesicles, prostate, and penis. Hypothalamus and anterior pituitary gland The control of male sexual and reproductive func- tion begins with secretion of gonadotropin-releasing hormone (GnRH) by the hypothalamus (Fig. 1.1). This hormone in turn stimulates the anterior pituitary gland to secrete two downstream hormones (termed gonadotropins). These hormones are luteinizing hor- mone (LH) and follicle-stimulating hormone (FSH). LH is the primary stimulus for the testicular secre- tion of testosterone, while FSH mainly stimulates spermatogenesis. Gonadotropin-releasing hormone (GnRH) Figure 1.1. Feedback regulation of the hypothalamic– The neuronal cells of the arcuate nuclei of the hypo- pituitary–gonadal (HPG) axis in males. Positive (stimulatory) effects are shown by + and inhibitory (negative feedback) effects by –. thalamus secrete GnRH, a 10-amino-acid peptide. The GnRH, gonadotropin-releasing hormone; LH, luteinizing hormone; endingsoftheseneuronsterminateinthemedian FSH, follicle-stimulating hormone. eminence of the hypothalamus, where they release GnRH into the hypothalamic–hypophysial portal vas- and stimulates the release of the two gonadotropins, cular system. The GnRH is transported to the anter- LH and FSH [1]. The output of GnRH is influenced by ior pituitary gland via the hypophysial portal blood three types of rhythmicity: seasonal, on a timescale of Fertility Preservation in Male Cancer Patients, ed. John P.Mulhall, Linda D. Applegarth, Robert D. Oates and Peter N. Schlegel. Published by Cambridge University Press. C Cambridge University Press 2013. 1 © in this web service Cambridge University Press www.cambridge.org Cambridge University Press 978-1-107-01212-7 - Fertility Preservation in Male Cancer Patients Editor-in-Chief John P. Mulhall Excerpt More information Section 1: Anatomy and physiology months and peaking in the spring; circadian, resulting effect, operating through the hypothalamus and anter- in highest testosterone levels during the early morning ior pituitary gland, reduces the testosterone secre- hours; and pulsatile, with peaks occurring every 90– tion back toward the desired operating level (see 120 minutes on average [2]. The intensity of this hor- Chapter 30). On the contrary, a testosterone-poor mone’s stimulus is determined (1) by the frequency of environment allows the hypothalamus to secrete large the cycles of secretion and (2) by the quantity of GnRH amounts of GnRH, with a corresponding increase released with each cycle. LH secretion by the anterior in LH and FSH from the anterior pituitary and an pituitary gland is also cyclical. LH follows the pulsatile increase in testicular testosterone secretion. release of GnRH. On the other hand, FSH secretion changes slowly with the fluctuation of GnRH secretion Testosterone and FSH over a period of many hours. In the seminiferous tubules, both FSH and testoster- one are necessary for the maintenance of spermatoge- Gonadotropic hormones: LH and FSH nesis. Specific FSH-dedicated receptors on the Sertoli Luteinizing hormone and follicle-stimulating cells induce Sertoli cell growth and elaboration of var- hormone are glycoproteins that are secreted by ious spermatogenic substances. Simultaneously, the gonadotropic cells in the anterior pituitary gland. In paracrine action of testosterone and dihydrotestos- the absence of GnRH secretion from the hypothal- terone from the interstitial Leydig cells stimulates and amus, the gonadotropes in the pituitary gland secrete supports spermatogenesis in the seminiferous tubule. essentially no LH or FSH. They exert their effects on their target tissues in the testes via the cyclic adenosine Inhibin monophosphate (cAMP) second messenger system. Inhibin is a glycoprotein, like both LH and FSH. It This, in turn, activates specific enzyme systems in the has a molecular weight of 36 000 daltons. Inhibin is respective target cells. dimeric in structure, and the two monomers are linked together by a single disulfide bond. The monomers are Testosterone and LH termed ␣ and ␤ subunits. The ␣ subunit is conserved in Testosterone is secreted by the Leydig cells in the inter- the different types of inhibin, but the ␤ subunit varies. stitium of the testes in response to stimulation by In humans, the Sertoli cells secrete inhibin B (␣␤B). LH from the anterior pituitary gland. The quantity of Inhibin B selectively suppresses FSH secretion in the testosterone secreted is nearly directly proportional to anterior pituitary gland by inhibiting transcription of the level of LH stimulation. Mature Leydig cells are the gene encoding the ␤ subunit of FSH [3]. Add- normally found in a child’s testes for a few weeks after itionally, inhibin has a slight effect on the hypotha- birth, but then involute until puberty. The secretion lamus to inhibit secretion of GnRH. Inhibin is released of LH at puberty causes testicular interstitial cells that from the Sertoli cells in response to robust, rapid look like fibroblasts to evolve into functional Leydig spermatogenesis. The end result is to diminish the cells. pituitary secretion of FSH. Conversely, when the sem- iniferous tubules fail to produce sperm, inhibin pro- Negative feedback of testosterone duction diminishes, resulting in a marked increase in FSH secretion. This potent inhibitory feedback effect The testosterone secreted by the testes in response on the anterior pituitary gland provides an important to LH inhibits the secretion of LH from the anterior negative feedback mechanism for control of spermato- pituitary. The bulk of this inhibition is most likely genesis, operating simultaneously with and in paral- from the direct effect of testosterone on the hypothal- lel to the negative feedback mechanism for control of amus to decrease the secretion of GnRH. A decrease testosterone secretion. in GnRH secretion results in a parallel decrease in secretion of both LH and FSH by the anterior pitu- itary. This decrease in LH, in turn, decreases the secre- Testis tion of testosterone by the testes. Hence, whenever Embryologically, the testes develop at the urogenital serum level of testosterone exceeds the body’s pre- ridge and descend into the scrotum via the inguinal set homeostatic level, the automatic negative feedback canal at birth. These two paired organs are suspended 2 © in this web service Cambridge University Press www.cambridge.org Cambridge University Press 978-1-107-01212-7 - Fertility Preservation in Male Cancer Patients Editor-in-Chief John P. Mulhall Excerpt More information Chapter 1: Functional anatomy of the HPG axis and the male reproductive tract pampiniform plexus is that it functions to efficiently maintain the optimal temperature for spermatogen- esis, which is below body temperature. Skandhan and Rajahariprasad hypothesized that the process of IC spermatogenesis results in a large amount of heat, ASTR IAC which has to be regulated [5]. The pampiniform plexus . IL EPIG and the human scrotal skin act as a radiator for the EXT robust heat generation. The scrotal skin is devoid of subcutaneous fat, and the presence of high sweat-gland density enables heat transmission. Upon exposure to cold temperatures, the scrotal surface is minimized by contraction, preventing temperature loss, and cremas- IN T . S PENISPENIS ter muscles retract the testes closer to the abdomen, for A P H temperature maintenance. The rich anastomoses between the testicular (internal spermatic) and vasal arteries allow mainte- nance of testicular viability if the internal spermatic artery is transected. In the testis, the artery gives rise to centrifugal arteries that pierce the testicular TESTIS parenchyma. Further branches divide into arterioles that bring in blood to peritubular and intertubular capillaries [6]. In some men, up to 90% of testicular blood supply derives from the testicular artery. Testicular venous drainage is through the pampini- form plexus, which in the region of the internal Figure 1.2. Vascular anatomy of the spermatic cord and testis. inguinal ring gives origin to the testicular vein [7]. (Reproduced from Gray H. Anatomy of the Human Body. Philadelphia, PA: Lea & Febiger, 1918; Bartleby.com, 2000.) See color The left testicular vein discharges into the left renal plate section. vein at a right angle, whereas the right testicular vein discharges directly into the inferior vena cava at an on the spermatic cords and are covered by numer- oblique angle. All testicular veins have valves. In the ous layers of tissue. Upon emerging from the inguinal region of the fourth lumbar vertebra the testicular ring in utero, they are covered by the tunica vaginalis, veins divide into two trunks, one lateral and one internal spermatic fascia,
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