The male gonads are the testes; they are the source of spermatozoa and also of male sex hormones called androgens. The other genital organs are the epididymides; the ductus, or vasa, deferentia; the seminal vesicles; the ejaculatory ducts; and the penis; as well as certain accessory structures, such as the prostate and the bulbourethral (Cowper) glands. The principal functions of these structures are to transport the spermatozoa from the testes to the exterior, to allow their maturation on the way, and to provide certain secretions that help form the semen.EXTERNAL GENITALIA
The penis
The penis, the male organ of copulation, is partly inside and partly outside the body. The inner part, attached to the bony margins of the pubic arch (that part of the pelvis directly in front and at the base of the trunk), is called the root of the penis. The second, or outer, portion is free, pendulous, and enveloped all over in skin; it is termed the body of the penis. The organ is composed chiefly of cavernous or erectile tissue that becomes engorged with blood to produce considerable enlargement and erection. The penis is traversed by a tube, the urethra, which serves as a passage both for urine and for semen.
The body of the penis, sometimes referred to as the shaft, is cylindrical in shape when flaccid but when erect is somewhat triangular in cross section, with the angles rounded. This condition arises because the right corpus cavernosum and the left corpus cavernosum, the masses of erectile tissue, lie close together in the dorsal part of the penis, while a single body, the corpus spongiosum, which contains the urethra, lies in a midline groove on the undersurface of the corpora cavernosa. The dorsal surface of the penis is that which faces upward and backward during erection.
The slender corpus spongiosum reaches beyond the extremities of the erectile corpora cavernosa and at its outer end is enlarged considerably to form a soft, conical, sensitive structure called the glans penis. The base of the glans has a projecting margin, the corona, and the groove where the corona overhangs the corpora cavernosa is referred to as the neck of the penis. The glans is traversed by the urethra, which ends in a vertical, slitlike, external opening. The skin over the penis is thin and loosely adherent and at the neck is folded forward over the glans for a variable distance to form the prepuce or foreskin. A median fold, the frenulum of the prepuce, passes to the undersurface of the glans to reach a point just behind the urethral opening. The prepuce can usually be readily drawn back to expose the glans.
The root of the penis comprises two crura, or projections, and the bulb of the penis. The crura and the bulb are attached respectively to the edges of the pubic arch and to the perineal membrane (the fibrous membrane that forms a floor of the trunk). Each crus is an elongated structure covered by the ischiocavernosus muscle, and each extends forward, converging toward the other, to become continuous with one of the corpora cavernosa. The oval bulb of the penis lies between the two crura and is covered by the bulbospongiosus muscle. It is continuous with the corpus spongiosum. The urethra enters it on the flattened deep aspect that lies against the perineal membrane, traverses its substances, and continues into the corpus spongiosum.
The two corpora cavernosa are close to one another, separated only by a partition in the fibrous sheath that encloses them. The erectile tissue of the corpora is divided by numerous small fibrous bands into many cavernous spaces, relatively empty when the penis is flaccid but engorged with blood during erection. The structure of the tissue of the corpus spongiosum is similar to that of the corpora cavernosa, but there is more smooth muscle and elastic tissue. A deep fascia, or sheet of connective tissue, surrounding the structures in the body of the penis is prolonged to form the suspensory ligament, which anchors the penis to the pelvic bones at the midpoint of the pubic arch.
The penis has a rich blood supply from the internal pudendal artery, a branch of the internal iliac artery, which supplies blood to the pelvic structures and organs, the buttocks, and the inside of the thighs. Erection is brought about by distension of the cavernous spaces with blood, which is prevented from draining away by compression of the veins in the area.
The penis is amply supplied with sensory and autonomic (involuntary) nerves. Of the autonomic nerve fibres the sympathetic fibres cause constriction of blood vessels, and the parasympathetic fibres cause their dilation. It is usually stated that ejaculation is brought about by the sympathetic system, which at the same time inhibits the desire to urinate and also prevents the semen from entering the bladder.
The scrotum
The scrotum is a pouch of skin lying below the pubic symphysis and just in front of the upper parts of the thighs. It contains the testes and lowest parts of the spermatic cord. A scrotal septum or partition divides the pouch into two compartments and arises from a ridge, or raphe, visible on the outside of the scrotum. The raphe turns forward onto the undersurface of the penis and is continued back onto the perineum (the area between the legs and as far back as the anus). This arrangement indicates the bilateral origin of the scrotum from two genital swellings that lie one on each side of the base of the phallus, the precursor of the penis or clitoris in the embryo. The swellings are also referred to as the labioscrotal swellings, because in females they remain separate to form the labia majora and in males they unite to form the scrotum.
The skin of the scrotum is thin, pigmented, devoid of fatty tissue, and more or less folded and wrinkled. There are some scattered hairs and sebaceous glands on its surface. Below the skin is a layer of involuntary muscle, the dartos, which can alter the appearance of the scrotum. On exposure of the scrotum to cold air or cold water, the dartos contracts and gives the scrotum a shortened, corrugated appearance; warmth causes the scrotum to become smoother, flaccid, and less closely tucked in around the testes. Beneath the dartos muscle are layers of fascia continuous with those forming the coverings of each of the two spermatic cords, which suspend the testes within the scrotum and contain each ductus deferens, the testicular blood and lymph vessels, the artery to the cremaster muscle (which draws the testes upward), the artery to each ductus deferens, the genital branch of the genitofemoral nerve, and the testicular network of nerves.
The scrotum is supplied with blood by the external pudendal branches of the femoral artery, which is the chief artery of the thigh, and by the scrotal branches of the internal pudendal artery. The veins follow the arteries. The lymphatic drainage is to the lymph nodes in the groin.
The testes
The two testes, or testicles, which usually complete their descent into the scrotum from their point of origin on the back wall of the abdomen in the seventh month after conception, are suspended in the scrotum by the spermatic cords. Each testis is 4 to 5 cm (about 1.5 to 2 inches) long and is enclosed in a fibrous sac, the tunica albuginea. This sac is lined internally by the tunica vasculosa, containing a network of blood vessels, and is covered by the tunica vaginalis, which is a continuation of the membrane that lines the abdomen and pelvis. The tunica albuginea has extensions into each testis that act as partial partitions to divide the testis into approximately 250 compartments, or lobules.
Each lobule contains one or more convoluted tubules, or narrow tubes, where sperm are formed. The tubules, if straightened, would extend about 70 cm (about 28 inches). The multistage process of sperm formation, which takes about 60 days, goes on in the lining of the tubules, starting with the spermatogonia, or primitive sperm cells, in the outermost layer of the lining. Spermatozoa (sperm) leaving the tubules are not capable of independent motion, but they undergo a further maturation process in the ducts of the male reproductive tract; the process may be continued when, after ejaculation, they pass through the female tract. Maturation of the sperm in the female tract is called capacitation.Each spermatozoon is a slender elongated structure with a head, a neck, a middle piece, and a tail. The head contains the cell nucleus. When the spermatozoon is fully mature, it is propelled by the lashing movements of the tail.
The male sex hormone testosterone is produced by Leydig cells. These cells are located in the connective (interstitial) tissue that holds the tubules together within each lobule. The tissue becomes markedly active at puberty under the influence of the interstitial-cell-stimulating hormone of the anterior lobe of the pituitary gland; this hormone in women is called luteinizing hormone. Testosterone stimulates the male accessory sex glands (prostate, seminal vesicles) and also brings about the development of male secondary sex characteristics at puberty. The hormone may also be necessary to cause maturation of sperm and to heighten the sex drive of the male. The testis is also the source of some of the female sex hormone estrogen, which may exert an influence on pituitary activity.
Each testis is supplied with blood by the testicular arteries, which arise from the front of the aorta just below the origin of the renal (kidney) arteries. Each artery crosses the rear abdominal wall, enters the spermatic cord, passes through the inguinal canal, and enters the upper end of each testis at the back. The veins leaving the testis and epididymis form a network, which ascends into the spermatic cord. The lymph vessels, which also pass through the spermatic cord, drain to the lateral and preaortic lymph nodes. Nerve fibres to the testis accompany the vessels; they pass through the renal and aortic nerve plexuses, or networks.
STRUCTURES OF THE SPERM CANAL
The epididymis, ductus deferens (or vas deferens), and ejaculatory ducts form the sperm canal. Together they extend from the testis to the urethra, where it lies within the prostate. Sperm are conveyed from the testis along some 20 ductules, or small ducts, which pierce the fibrous capsule to enter the head of the epididymis. The ductules are straight at first but become dilated and then much convoluted to form distinct compartments within the head of the epididymis. They each open into a single duct, the highly convoluted duct of the epididymis, which constitutes the “body” and “tail” of the structure. It is held together by connective tissue but if unraveled would be nearly 6 metres (20 feet) long. The duct enlarges and becomes thicker-walled at the lower end of the tail of the epididymis, where it becomes continuous with the ductus deferens.
The ductules from the testis have a thin muscular coat and a lining that consists of alternating groups of high columnar cells with cilia (hairlike projections) and low cells lacking cilia. The cilia assist in moving sperm toward the epididymis. In the duct of the epididymis the muscle coat is thicker and the lining is thick with tufts of large nonmotile cilia. There is some evidence that the ductules and the first portion of the duct of the epididymis remove excess fluid and extraneous debris from the testicular secretions entering these tubes. The blood supply to the epididymis is by a branch from the testicular artery given off before that vessel reaches the testis.
The ductus deferens, or vas deferens, is the continuation of the duct of the epididymis. It commences at the lower part of the tail of the epididymis and ascends along the back border of the testis to its upper pole. Then, as part of the spermatic cord, it extends to the deep inguinal ring. Separating from the other elements of the spermatic cord—the blood vessels, nerves, and lymph vessels—at the ring, the ductus deferens makes its way through the pelvis toward the base of the prostate, where it is joined by the seminal vesicle to form the ejaculatory duct. A part of the ductus that is dilated and rather tortuous, near the base of the urinary bladder, is called the ampulla.
The ductus deferens has a thick coat of smooth muscle that gives it a characteristic cordlike feel. The longitudinal muscle fibres are well developed, and peristaltic contractions (contractions in waves) move the sperm toward the ampulla. The mucous membrane lining the interior is in longitudinal folds and is mostly covered with nonciliated columnar cells, although some cells have nonmotile cilia. The ampulla is thinner-walled and probably acts as a sperm store.
ACCESSORY GLANDS
The prostate gland, seminal vesicles, and bulbourethral glands
These structures provide secretions to form the bulk of the seminal fluid of an ejaculate. The prostate gland is in the lesser or true pelvis, centred behind the lower part of the pubic arch. It lies in front of the rectum. The prostate is shaped roughly like an inverted pyramid; its base is directed upward and is immediately continuous with the neck of the urinary bladder. The urethra traverses its substance. The two ejaculatory ducts enter the prostate near the upper border of its posterior surface. The prostate is of a firm consistency, surrounded by a capsule of fibrous tissue and smooth muscle. It measures about 4 cm across, 3 cm in height, and 2 cm front to back (about 1.6 by 1.2 by 0.8 inch) and consists of glandular tissue contained in a muscular framework. It is imperfectly divided into three lobes. Two lobes at the side form the main mass and are continuous behind the urethra. In front of the urethra they are connected by an isthmus of fibromuscular tissue devoid of glands. The third, or median, lobe is smaller and variable in size and may lack glandular tissue. There are three clinically significant concentric zones of prostatic glandular tissue about the urethra. A group of short glands that are closest to the urethra and discharge mucus into its channel are subject to simple enlargement. Outside these is a ring of submucosal glands (glands from which the mucosal glands develop), and farther out is a large outer zone of long branched glands, composing the bulk of the glandular tissue. Prostate cancer is almost exclusively confined to the outer zone. The glands of the outer zone are lined by tall columnar cells that secrete prostatic fluid under the influence of androgens from the testis. The fluid is thin, milky, and slightly acidic.
The seminal vesicles are two structures, about 5 cm (2 inches) in length, lying between the rectum and the base of the bladder. Their secretions form the bulk of semen. Essentially, each vesicle consists of a much-coiled tube with numerous diverticula or outpouches that extend from the main tube, the whole being held together by connective tissue. At its lower end the tube is constricted to form a straight duct or tube that joins with the corresponding ductus deferens to form the ejaculatory duct. The vesicles are close together in their lower parts, but they are separated above where they lie close to the deferent ducts. The seminal vesicles have longitudinal and circular layers of smooth muscle, and their cavities are lined with mucous membrane, which is the source of the secretions of the organs. These secretions are ejected by muscular contractions during ejaculation. The activity of the vesicles is dependent on the production of the hormone androgen by the testes. The secretion is thick, sticky, and yellowish; it contains the sugar fructose and is slightly alkaline.
The bulbourethral glands, often called Cowper glands, are pea-shaped glands that are located beneath the prostate gland at the beginning of the internal portion of the penis. The glands, which measure only about 1 cm (0.4 inch) in diameter, have slender ducts that run forward and toward the centre to open on the floor of the spongy portion of the urethra. They are composed of a network of small tubes, or tubules, and saclike structures; between the tubules are fibres of muscle and elastic tissue that give the glands muscular support. Cells within the tubules and sacs contain droplets of mucus, a thick protein compound. The fluid excreted by these glands is clear and thick and acts as a lubricant; it is also thought to function as a flushing agent that washes out the urethra before the semen is ejaculated; it may also help to make the semen less watery and to provide a suitable living environment for the sperm.
Ejaculatory ducts
The two ejaculatory ducts lie on each side of the midline and are formed by the union of the duct of the seminal vesicle, which contributes secretions to the semen, with the end of the ductus deferens at the base of the prostate. Each duct is about 2 cm (about 0.8 inch) long and passes between a lateral and the median lobe of the prostate to reach the floor of the prostatic urethra. This part of the urethra has on its floor (or posterior wall) a longitudinal ridge called the urethral crest. On each side is a depression, the prostatic sinus, into which open the prostatic ducts. In the middle of the urethral crest is a small elevation, the colliculus seminalis, on which the opening of the prostatic utricle is found. The prostatic utricle is a short diverticulum or pouch lined by mucous membrane; it may correspond to the vagina or uterus in the female. The small openings of the ejaculatory ducts lie on each side of or just within the opening of the prostatic utricle. The ejaculatory ducts are thin-walled and lined by columnar cells.