FIGURE The tunica albuginea is a connective tissue capsule forming the outer part of each testis.
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1 Testicular Histology (see p in text) FIGURE The tunica albuginea is a connective tissue capsule forming the outer part of each testis. 2. Septa are extensions of the tunica albuginea that divide the testes into lobules. 3. Located within the lobules are the seminiferous tubules, which are the site of sperm cell production. 4. Interstitial cells are endocrine cells found in the connective tissue around the seminiferous tubules. They produce testosterone. 5. The tubulus rectus is a short, straight tube into which a seminiferous tubule empties. 6. The rete testis is a tubular network into which the tubuli recti empty. 7. The efferent ductules are conducting tubes formed by the union of the rete testis. They leave the testis and connect to the epididymis. 8. The epididymis is a comma-shaped structure on the posterior testis. The duct of the epididymis receives sperm cells from the efferent ductules and is the site of sperm cell maturation, which takes from one to ten days. 9. The duct of the epididymis joins the ductus deferens or vas deferens, which transports sperm cells toward the outside of the body. Sperm Cell Development (see pp in text) 1. Spermatogenesis is the formation of sperm cells, or spermatozoa (sing., spermatozoon), by meiosis. 2. Spermatogenesis accomplishes three goals. A. The number of chromosomes (diploid number; 2n) is reduced by half (haploid number; n) because the sperm cell is formed by meiosis. In humans the diploid number is 46 and the haploid number is 23. Why is it necessary for the number of chromosomes to be reduced by half (hint: the number of chromosomes in the oocyte is also reduced in half by meiosis)? 1
2 B. In meiosis there is crossing over (exchange of genetic material between chromatids), and there is random assortment of homologous chromosomes. Therefore, all of the sperm cells are genetically different from each other. Because the same process occurs in the development of the oocyte, this means every individual is genetically unique from other individuals (unless they are identical twins). C. The sperm cells become specialized to swim and find the mate of their lives. A sperm cell consists of a head, middle piece, and tail. FIGURE ) The head contains the nucleus (with the genetic material), and is covered by the acrosome (a cap that contains enzymes that allow the sperm cell to digest its way into the oocyte). 2) A middle piece that contains the organelles responsible for moving the tail. 3) The tail or flagellum moves back and forth, enabling the sperm cell to swim. Why are mitochondria found in the middle piece of the sperm cell? 3. The steps of spermatogenesis. FIGURE 28.5 A. Spermatogonia are the cells from which sperm cells arise. The spermatogonia divide by mitosis. One daughter cell remains a spermatogonium that can divided again by mitosis. One daughter cell becomes a primary spermatocyte. B. The primary spermatocyte divides by meiosis to form secondary spermatocytes. C. The secondary spermatocytes divide by meiosis to form spermatids. D. The spermatids differentiate to form sperm cells. 4. Support cells. A. Spermatogenesis takes place within the seminiferous tubules. The process begins at the periphery of the seminiferous tubules and ends with the release of sperm cells into the lumen of the tubule. 2
3 B. Sustentacular cells (Sertoli, cells) extend from the periphery to the lumen and assist the process of spermatogenesis in several ways. 1) The sustentacular cells surround and provide nourishment to the developing cells. 2) The sustentacular cells are joined by tight junctions. This separates the primary spermatocyte and subsequent cells from the immune system. This is called the blood-testes barrier. Without this barrier the immune system would destroy the sperm cells. 3) The sustentacular cells produce chemicals that promote sperm cell formation, such as dihydrotestosterone, estradiol, and androgen-binding protein. 4) The sustentacular cells produce the hormone inhibin (more later). PHYSIOLOGY OF MALE REPRODUCTION Regulation of Sex Hormone Secretion FIGURE Gonadotropin releasing hormone (GnRH) produced by the hypothalamus stimulates the release of the gonadotropins FSH and LH from the anterior pituitary. Gonadotropins influence the activity of the gonads (testes or ovaries). A. Follicle-stimulating hormone (FSH) 1) FSH stimulates the development of the seminiferous tubules and the process of spermatogenesis (sperm cell production). 2) FSH stimulates the activity of sustentacular cells in the seminiferous tubules. Sustentacular cells provide nourishment for the developing sperm cells. Sustentacular cells also produce the hormone inhibin which inhibits FSH production (negative-feedback mechanism that regulates sperm cell production). GnRH, FSH, Inhibin, which causes FSH B. Luteinizing hormone (LH). 1) LH stimulates interstitial cells (Leydig cells) to produce androgens. Androgens are a group of hormones that stimulate the development of male sexual characteristics. The major androgen is testosterone. 2) Testosterone is necessary for normal sustentacular cell function, spermatogenesis, and the development of the sex organs and secondary sexual characteristics. 3) Testosterone has a negative-feedback effect on the hypothalamus, inhibiting GnRH secretion, and on the anterior pituitary, inhibiting LH and FSH secretion. GnRH, LH, Testosterone, which causes GnRH, LH, FSH 3
4 Puberty 1. Before birth the placenta secretes human chorionic gonadotropin (HCG), which stimulates the testes to secrete testosterone. The testosterone is responsible for the development of male sexual characteristics in the fetus and descent of the testes. 2. After birth, there is no longer HCG stimulation. From birth to puberty there is little testosterone secretion. The testosterone that is produced inhibits GnRH secretion from the hypothalamus. 3. Puberty is the change from a child into a sexually functional adult. A. For unknown reasons, testosterone no longer inhibits the hypothalamus as much, and production of GnRH increases. The result is increased FSH, LH, and testosterone production. B. There is speculation that decreased secretion of melatonin by the pineal body plays a role in this process. Effects of Testosterone 1. Primary sexual characteristics. From puberty to age 20, testosterone causes the penis, the scrotum, and the testes to enlarge. Testosterone also is necessary for spermatogenesis. 2. Secondary sexual characteristics. A. Development of secondary sex organs, such as the prostate gland and seminal vesicles. B. Growth of hair over the pubis, under the arms (axilla), on the chest, and on the face (beard) 1) Hair changes from vellus hairs (short, fine, unpigmented hairs) to terminal hairs (long, coarse, pigmented hairs). 2) If genetically predisposed, testosterone can lead to baldness. C. Lowering of the voice because of enlargement of the larynx. D. The skin increases in thickness. Increased melanin production makes the skin darker. Excessive secretion of sebum by the sebaceous glands plays a role in acne production. E. Increased muscular development as a result of increased protein synthesis. F. Thicker and longer bones because of increased bone growth. G. Increase metabolic rate (5 to 10%). Male Sexual Behavior and the Male Sex Act 1. Sex drive. A. Testosterone is responsible, at least in part, for sex drive or libido in males. B. In castrated adult males, there is, in general, a decrease in sex drive. 1) Sexual function can range from complete loss of sex drive to continued, normal sexual activity. 2) Testosterone from the adrenal glands may be responsible for retention of sex drive 3) Treatment with testosterone can restore sex drive. 4
5 2. Male climacteric. A. After puberty, testosterone is produced throughout life. However, in the late 40's to 50's the level of testosterone declines. B. Symptoms can include decreased sexual function and occasionally hot flashes, feelings of suffocation, and psychic disorders similar to menopausal symptoms observed in women. Erection 1. Erection occurs when the spongy tissue of the corpora cavernosa and corpus spongiosum fills with blood. A. Blood vessels dilate and there is increased blood flow into the spongy tissues of the penis. B. Expansion of the spongy tissues compresses veins draining the penis, which reduces blood flow out of the penis. 2. Mechanism of blood vessel dilation in the penis. FIGURE 28.9 A. Action potentials reaching presynaptic terminals cause voltage-gated Ca 2+ channels to open. B. Ca 2+ move into the presynaptic terminal and cause the release of acetylcholine (ACh) into the synaptic cleft. C. ACh activates a G protein mediated response and smooth muscle cells in the wall of blood vessels relax, resulting in blood vessel dilation. D. Ca 2+ also activate nitric oxide synthase, an enzyme that promotes the formation of nitric oxide (NO) from arginine (an amino acid). E. NO diffuses into smooth muscle and activates guanylate cyclase. This enzyme promotes the conversion of guanine triphosphate (GTP) to cyclic GMP (cgmp). The increase in cgmp results in smooth muscle relaxation. F. Remember that chemical signals that stimulate a response from a cell are usually quickly turned off inactivating the chemical signal. Thus, an enzyme inactivates cgmp. Viagra is used to treat erectile dysfunction (ED) because it blocks the activity of the enzyme that inactivates cgmp. Thus, cgmp builds up in smooth muscle cells and they relax. 5
6 Level in spinal cord T10 - T12 Aware of stimulation To brain Sexual stimuli From brain S2 - S4 Parasympathetic Sympathetic Penis Bladder Internal urinary sphincter Glands Duct system Testes External urinary sphincter Bulbospongiosus muscle 3. Neural control of erection. A. The brain. 1) Psychic erection can occur due to thoughts, visual images, etc. Impulses from the brain pass to parasympathetic and sympathetic centers in the spinal cord. The parasympathetic control is the most important. 2) Erection can result from stimulation of the genitals or other parts of the body. The stimulation is detected by the brain and output is sent to the spinal cord centers. 3) Impotence (inability to have an erection) can be produced by anxiety, fear, or depression. B. The spinal cord. Stimulation of the penis (especially the glans penis) activates a parasympathetic reflex in the spinal cord. This reflex can occur independently of input from the brain, for example, following spinal cord injury. 6
7 Emission Level in spinal cord Aware of stimulation To brain T10 - T12 S2 - S4 Sympathetic Penis Semen Bladder Internal urinary sphincter Glands Testes External urinary sphincter Bulbospongiosus muscle 1. Emission is the accumulation of semen (sperm cells plus glandular secretions) in the prostatic urethra. 2. A sympathetic reflex causes contraction of smooth muscle in the epididymis, ductus deferens, and ejaculatory duct. This results in sperm cell transport. 3. A sympathetic reflex causes contraction of smooth muscle in the seminal vesicles and prostate gland, and glandular secretions are mixed with the sperm cells to form semen. 7
8 Ejaculation Level in spinal cord Aware of emission To brain T10 - T12 S2 - S4 Sympathetic (different pathway than emission) Penis Semen Bladder Internal urinary sphincter Glands Duct system Testes External urinary sphincter Bulbospongiosus muscle 3. Ejaculation is the ejection of the semen from the body. A. Semen within the prostatic urethra activates a somatic motor reflex that causes repeated contraction of the external urinary sphincter and the bulbospongiosus muscle, propelling the semen through the membranous and spongy urethra. B. A sympathetic reflex causes contraction of the internal urinary sphincter, preventing the ejection of the semen into the urinary bladder (retrograde ejaculation) and preventing the mixing of urine with the semen. The prostate gland continues to contract, adding secretions to the semen. 2. Orgasm is the pleasurable feeling associated with ejaculation. It is strictly a cerebral phenomenon and it is possible to experience orgasm without ejaculation, and vice versa. 3. Resolution is a phase after ejaculation in which the penis becomes flaccid, there is an overall feeling of satisfaction, and the male is unable to have another erection and ejaculation. The length of the resolution phase can vary from a few minutes to hours in the young or as long as a month in the elderly. 8
9 Anabolic Steroids 1. Anabolic steroids are artificial variants of testosterone. Some people believe that using anabolic steroids can increase athletic performance. Why would they believe this? 2. The development of muscle mass when using anabolic steroids is dose dependent. A. At low, relatively safe dosages, there is little increase in muscle mass. B. At high dosages (5 to 10 times those used in most controlled studies), muscle mass increases. In athletic events that require increased strength, this can be an advantage. Current evidence suggests that anabolic steroids do not increase endurance. 3. The use of anabolic steroids has a down side. Which of the following side effects would be expected from taking anabolic steroids? Testosterone production Spermatogenesis Size of testes Acne Hair growth in women Voice in women Sex drive in women Size of clitoris Lower or higher 4. In addition to the above changes, anabolic steroids produce other undesirable and dangerous side effects. A. Impotence in males. Apparently the anabolic steroids do not affect the brain in the same way as natural testosterone. However, they do inhibit natural testosterone production. B. Breast reduction in women and breast enlargement in men. Women stop menstruating. C. Increase in aggressive behavior. D. Liver and cardiovascular disease. 9
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