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Ganong's Review of Medical Physiology, 23rd Edition

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406 SECTION IV Endocrine & Reproductive <strong>Physiology</strong><br />

Normally, erection is terminated by sympathetic vasoconstrictor<br />

impulses to the penile arterioles.<br />

Ejaculation<br />

Ejaculation is a two-part spinal reflex that involves emission,<br />

the movement <strong>of</strong> the semen into the urethra; and ejaculation<br />

proper, the propulsion <strong>of</strong> the semen out <strong>of</strong> the urethra at the<br />

time <strong>of</strong> orgasm. The afferent pathways are mostly fibers from<br />

touch receptors in the glans penis that reach the spinal cord<br />

through the internal pudendal nerves. Emission is a sympathetic<br />

response, integrated in the upper lumbar segments <strong>of</strong><br />

the spinal cord and effected by contraction <strong>of</strong> the smooth<br />

muscle <strong>of</strong> the vasa deferentia and seminal vesicles in response<br />

to stimuli in the hypogastric nerves. The semen is propelled<br />

out <strong>of</strong> the urethra by contraction <strong>of</strong> the bulbocavernosus muscle,<br />

a skeletal muscle. The spinal reflex centers for this part <strong>of</strong><br />

the reflex are in the upper sacral and lowest lumbar segments<br />

<strong>of</strong> the spinal cord, and the motor pathways traverse the first to<br />

third sacral roots and the internal pudendal nerves.<br />

PSA<br />

The prostate produces and secretes into the semen and the<br />

bloodstream a 30 kDa serine protease generally called prostate-specific<br />

antigen (PSA). The gene for PSA has two androgen<br />

response elements. PSA hydrolyzes the sperm motility<br />

inhibitor semenogelin in semen, and it has several substrates<br />

in plasma, but its precise function in the circulation is unknown.<br />

An elevated plasma PSA occurs in prostate cancer and<br />

is widely used as a screening test for this disease, though PSA<br />

is also elevated in benign prostatic hyperplasia and prostatitis.<br />

O<br />

H<br />

Cholesterol<br />

Pregnenolone<br />

17α-Hydroxypregnenolone<br />

Dehydroepiandrosterone<br />

OH<br />

Dihydrotestosterone<br />

In some<br />

target tissues<br />

5α-reductase,<br />

type 1 or type 2<br />

Vasectomy<br />

Bilateral ligation <strong>of</strong> the vas deferens (vasectomy) has proved to<br />

be a relatively safe and convenient contraceptive procedure.<br />

However, it has proven difficult to restore the patency <strong>of</strong> the<br />

vas in those wishing to restore fertility, and the current success<br />

rate for such operations, as measured by the subsequent production<br />

<strong>of</strong> pregnancy, is about 50%. Half <strong>of</strong> the men who have<br />

been vasectomized develop antibodies against spermatozoa,<br />

and in monkeys, the presence <strong>of</strong> such antibodies is associated<br />

with a higher incidence <strong>of</strong> infertility after restoration <strong>of</strong> the<br />

patency <strong>of</strong> the vas. However, the anti-sperm antibodies do not<br />

appear to have any other adverse effects.<br />

ENDOCRINE FUNCTION OF THE TESTES<br />

Chemistry & Biosynthesis <strong>of</strong> Testosterone<br />

Testosterone, the principal hormone <strong>of</strong> the testes, is a C 19 steroid<br />

with an –OH group in the 17 position (Figure 25–15). It is<br />

synthesized from cholesterol in the Leydig cells and is also<br />

formed from androstenedione secreted by the adrenal cortex.<br />

The biosynthetic pathways in all endocrine organs that form<br />

steroid hormones are similar, the organs differing only in the<br />

enzyme systems they contain. In the Leydig cells, the 11- and<br />

21-hydroxylases found in the adrenal cortex (see Figure 22–7)<br />

are absent, but 17α-hydroxylase is present. Pregnenolone is<br />

therefore hydroxylated in the 17 position and then subjected<br />

to side chain cleavage to form dehydroepiandrosterone. Androstenedione<br />

is also formed via progesterone and 17-hydroxyprogesterone,<br />

but this pathway is less prominent in humans.<br />

Dehydroepiandrosterone and androstenedione are then converted<br />

to testosterone.<br />

The secretion <strong>of</strong> testosterone is under the control <strong>of</strong> LH, and<br />

the mechanism by which LH stimulates the Leydig cells involves<br />

Progesterone<br />

17α-Hydroxyprogesterone<br />

Androstenedione<br />

FIGURE 25–15 Biosynthesis <strong>of</strong> testosterone. The formulas <strong>of</strong> the precursor steroids are shown in Figure 22–7. Although the main secretory<br />

product <strong>of</strong> the Leydig cells is testosterone, some <strong>of</strong> the precursors also enter the circulation.<br />

O<br />

Testosterone<br />

OH

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