Reproduction 721
Erection is controlled by two portions of the central ner-
vous system—the hypothalamus in the brain and the sacral por-
tion of the spinal cord. Conscious sexual thoughts originating
in the cerebral cortex act via the hypothalamus to control the
sacral region, which in turn increases parasympathetic nerve
activity to promote vasodilation and erection of the penis. Con-
scious thought is not required for erection, however, because
sensory stimulation of the penis can more directly activate the
sacral region of the spinal cord and cause an erection.
Male Fertility
The approximate volume of semen for each ejaculation is 1.5
to 5.0 milliliters. The bulk of this fluid (45% to 80%) is pro-
duced by the seminal vesicles and 15% to 30% is contributed
by the prostate. There are usually between 60 and 150 million
Figure 20.21 Role of nitric oxide in erection of the penis. Nitric oxide is released by parasympathetic axons in the penis,
and is also produced as a paracrine regulator by endothelial cells in blood vessels of the penis. (1) Nitric oxide activates guanylate
cyclase, causing the production of cyclic GMP (cGMP). (2) cGMP causes Ca^2 1 channels to close in the plasma membrane of the
vascular smooth muscle cells, decreasing the cytoplasmic Ca^2 1 concentration. (3) This produces relaxation of the smooth muscle cells,
causing vasodilation in the penis and engorgement of the erectile tissue. Viagra and related drugs inhibit the phosphodiesterase that
catalyzes the breakdown of cGMP, thereby increasing the availability of cGMP to promote an erection.
Parasympathetic axon
Nitric
oxide
Activates
guanylate
cyclase
Inhibits
Viagra
PDE
5' GMP
Ca2+
channels
close
cGMP
GTP
Decreased
cytoplasmic Ca2+
Ca2+
NO
Parasympathetic axon
ACh
Vascular
endothelial
cell
L-Arginine
O 2
Vascular
smooth muscle
cell
Smooth muscle
relaxes
Engorgement
of erectile
tissue
Erection
Stimulates
eNOS
1
3
2
CLINICAL APPLICATION
Erectile dysfunction is the inability to produce or sustain an
erection sufficient for sexual intercourse. This may be due to
physical problems that include heart disease, enlarged pros-
tate, atherosclerosis, and metabolic syndrome, or to such
psychological problems as depression and stress. Erection
requires nitric oxide (NO), which activates guanylate cyclase
in smooth muscle cells to raise cGMP as previously discussed
( fig. 20.21 ). Sildenafil ( Viagra ) and Tadalafil ( Cialis ) inhibit
the enzyme phosphodiesterase ( PDE ), which breaks down
cGMP. Specifically, they block the PDE-5 isoenzyme that pre-
dominates in the penis, raising the cGMP to promote erection
( fig. 20.21 ). These drugs have minimal effect on systemic blood
pressure, but vasodilation elsewhere in the body can produce
such side effects as headache, flushing, and nasal congestion.