Unique for its role in human reproduction, a gamete is a specialized sex cell carrying 23 chromosomes—one half the number in body cells.  Those with sperm producing and conducting (SPC) organs possess a gamete called a sperm cell (or spermatozoon), while those with egg producing and conducting (EPC) organs have a gamete known as an oocyte (or egg). At fertilization, the chromosomes in the sperm combine with the chromosomes in an oocyte. The function of SPC organs is to produce sperm and transfer them outside of the body, potentially into the egg conducting organs of another individual. The paired testes are a crucial component in this process, as they produce both sperm and androgens, the hormones that support the physiological function of SPC organs. In humans, the most important androgen for those with SPC organs is testosterone. Several accessory organs and ducts aid the process of sperm maturation and transport the sperm and other seminal components to the penis, which delivers sperm to the egg conducting organs. In this section, we examine each of these different structures, and discuss the process of sperm production and transport.

Scrotum

The testes are located in a skin-covered, highly pigmented, muscular sack called the scrotum that extends from the body behind the penis. This location is important in sperm production, which occurs within the testes, and proceeds more efficiently when the testes are kept 2 to 4°C below core body temperature.

The dartos muscle makes up the subcutaneous muscle layer of the scrotum. It continues internally to make up the scrotal septum, a wall that divides the scrotum into two compartments, each housing one testis. Descending from the internal oblique muscle of the abdominal wall are the two cremaster muscles, which cover each testis like a muscular net. By contracting simultaneously, the dartos and cremaster muscles can elevate the testes in cold weather (or water), moving the testes closer to the body and decreasing the surface area of the scrotum to retain heat. Alternatively, as the environmental temperature increases, the scrotum relaxes, moving the testes farther from the body core and increasing scrotal surface area, which promotes heat loss. Externally, the scrotum has a raised medial thickening on the surface called the raphae.

Testes

The testes (singular = testis) are the gonads —that is, the sperm producing organs. They produce both sperm and androgens, such as testosterone, and are active starting at puberty.

Paired ovals, the testes are each approximately 4 to 5 cm in length and are housed within the scrotum. They are surrounded by two distinct layers of protective connective tissue. The outer tunica vaginalis is a serous membrane that has both a parietal and a thin visceral layer. Beneath the tunica vaginalis is the tunica albuginea, a tough, white, dense connective tissue layer covering the testis itself. Not only does the tunica albuginea cover the outside of the testis, it also invaginates to form septa that divide the testis into 300 to 400 structures called lobules. Within the lobules, sperm develop in structures called seminiferous tubules. During the seventh month of the developmental period of a fetus, each testis moves through the abdominal musculature to descend into the scrotal cavity. This is called the “descent of the testis.” Cryptorchidism is the clinical term used when one or both of the testes fail to descend into the scrotum prior to birth.

The tightly coiled seminiferous tubules form the bulk of each testis. They are composed of developing sperm cells surrounding a lumen, the hollow center of the tubule, where formed sperm are released into the duct system of the testis. Specifically, from the lumens of the seminiferous tubules, sperm move into the straight tubules (or tubuli recti), and from there into a fine meshwork of tubules called the rete testes. Sperm leave the rete testes, and the testis itself, through the 15 to 20 efferent ductules that cross the tunica albuginea.

Inside the seminiferous tubules are six different cell types. These include supporting cells called sustentacular cells, as well as five types of developing sperm cells called germ cells. Germ cell development progresses from the basement membrane—at the perimeter of the tubule—toward the lumen. Let’s look more closely at these cell types.

Sustentacular Cells

Surrounding all stages of the developing sperm cells are elongate, branching Sustentacular cells (Sertoli cells).  These are a type of supporting cell that is typically found in epithelial tissue. Sustentacular cells secrete signaling molecules that promote sperm production and can control whether germ cells live or die. They extend physically around the germ cells from the peripheral basement membrane of the seminiferous tubules to the lumen. Tight junctions between these sustentacular cells create the blood–testis barrier, which keeps bloodborne substances from reaching the germ cells and, at the same time, keeps surface antigens on developing germ cells from escaping into the bloodstream and prompting an autoimmune response.

Germ Cells

The least mature cells, the spermatogonia (singular = spermatogonium), line the basement membrane inside the tubule. Spermatogonia are the stem cells of the testis, which means that they are still able to differentiate into a variety of different cell types throughout adulthood. Spermatogonia divide to produce primary and secondary spermatocytes, then spermatids, which finally produce formed sperm. The process that begins with spermatogonia and concludes with the production of sperm is called spermatogenesis.

Spermatogenesis

As just noted, spermatogenesis occurs in the seminiferous tubules that form the bulk of each testis. The process begins at puberty, after which time sperm are produced constantly throughout a person’s life. One production cycle, from spermatogonia through formed sperm, takes approximately 64 days. A new cycle starts approximately every 16 days, although this timing is not synchronous across the seminiferous tubules. Sperm counts—the total number of sperm a man produces—slowly decline after age 35, and some studies suggest that smoking can lower sperm counts irrespective of age.

The process of spermatogenesis begins with mitosis of the diploid spermatogonia. Because these cells are diploid (2n), they each have a complete copy of the father’s genetic material, or 46 chromosomes. However, mature gametes are haploid (1n), containing 23 chromosomes—meaning that daughter cells of spermatogonia must undergo a second cellular division through the process of meiosis.

Two identical diploid cells result from spermatogonia mitosis. One of these cells remains a spermatogonium, and the other becomes a primary spermatocyte, the next stage in the process of spermatogenesis. As in mitosis, DNA is replicated in a primary spermatocyte, and the cell undergoes cell division to produce two cells with identical chromosomes. Each of these is a secondary spermatocyte. Now a second round of cell division occurs in both of the secondary spermatocytes, separating the chromosome pairs. This second meiotic division results in a total of four cells with only half of the number of chromosomes. Each of these new cells is a spermatid. Although haploid, early spermatids look very similar to cells in the earlier stages of spermatogenesis, with a round shape, central nucleus, and large amount of cytoplasm. A process called spermiogenesis transforms these early spermatids, reducing the cytoplasm, and beginning the formation of the parts of a true sperm. The fifth stage of germ cell formation—spermatozoa, or formed sperm—is the end result of this process, which occurs in the portion of the tubule nearest the lumen. Eventually, the sperm are released into the lumen and are moved along a series of ducts in the testis toward a structure called the epididymis for the next step of sperm maturation.

Structure of Formed Sperm

Sperm are smaller than most cells in the body; in fact, the volume of a sperm cell is 85,000 times less than that of the oocyte. Approximately 100 to 300 million sperm are produced each day, whereas typically only one oocyte is released via ovulation per month. As is true for most cells in the body, the structure of sperm cells speaks to their function. Sperm have a distinctive head, mid-piece, and tail region. The head of the sperm contains the extremely compact haploid nucleus with very little cytoplasm. These qualities contribute to the overall small size of the sperm (the head is only 5 μm long). A structure called the acrosome covers most of the head of the sperm cell as a “cap” that is filled with lysosomal enzymes important for preparing sperm to participate in fertilization. Tightly packed mitochondria fill the mid-piece of the sperm. ATP produced by these mitochondria will power the flagellum, which extends from the neck and the mid-piece through the tail of the sperm, enabling it to move the entire sperm cell. The central strand of the flagellum, the axial filament, is formed from one centriole inside the maturing sperm cell during the final stages of spermatogenesis.

Sperm Transport

To fertilize an egg, sperm must be moved from the seminiferous tubules in the testes, through the epididymis, and—later during ejaculation—along the length of the penis and potentially out into the egg conducting organs.

Role of the Epididymis

From the lumen of the seminiferous tubules, the immotile sperm are surrounded by testicular fluid and moved to the epididymis (plural = epididymides), a coiled tube attached to the testis where newly formed sperm continue to mature. Though the epididymis does not take up much room in its tightly coiled state, it would be approximately 6 m (20 feet) long if straightened. It takes an average of 12 days for sperm to move through the coils of the epididymis, with the shortest recorded transit time in humans being one day. Sperm enter the head of the epididymis and are moved along predominantly by the contraction of smooth muscles lining the epididymal tubes. As they are moved along the length of the epididymis, the sperm further mature and acquire the ability to move under their own power. Once inside the egg conducting organs, they will use this ability to move independently toward the unfertilized egg. The more mature sperm are then stored in the tail of the epididymis (the final section) until ejaculation occurs.

Duct System

During ejaculation, sperm exit the tail of the epididymis and are pushed by smooth muscle contraction to the ductus deferens (also called the vas deferens). The ductus deferens is a thick, muscular tube that is bundled together inside the scrotum with connective tissue, blood vessels, and nerves into a structure called the spermatic cord. Because the ductus deferens is physically accessible within the scrotum, surgical sterilization to interrupt sperm delivery can be performed by cutting and sealing a small section of the ductus (vas) deferens. This procedure is called a vasectomy, and it is an effective form of birth control. Although it may be possible to reverse a vasectomy, clinicians consider the procedure permanent.

From each epididymis, each ductus deferens extends superiorly into the abdominal cavity through the inguinal canal in the abdominal wall. From here, the ductus deferens continues posteriorly to the pelvic cavity, ending posterior to the bladder where it dilates in a region called the ampulla (meaning “flask”).

Sperm make up only 5 percent of the final volume of semen, the thick, milky fluid that the male ejaculates. The bulk of semen is produced by three critical accessory glands of the male reproductive system: the seminal vesicles, the prostate, and the bulbourethral glands.

Seminal Vesicles

As sperm pass through the ampulla of the ductus deferens at ejaculation, they mix with fluid from the associated seminal vesicle. The paired seminal vesicles are glands that contribute approximately 60 percent of the semen volume. Seminal vesicle fluid contains large amounts of fructose, which is used by the sperm mitochondria to generate ATP to allow movement through the egg conducting organs.

The fluid, now containing both sperm and seminal vesicle secretions, next moves into the associated ejaculatory duct, a short structure formed from the ampulla of the ductus deferens and the duct of the seminal vesicle. The paired ejaculatory ducts transport the seminal fluid into the next structure, the prostate gland.

Prostate Gland

As shown in Figure 1, the centrally located prostate gland sits anterior to the rectum at the base of the bladder surrounding the prostatic urethra (the portion of the urethra that runs within the prostate). About the size of a walnut, the prostate is formed of both muscular and glandular tissues. It excretes an alkaline, milky fluid to the passing seminal fluid—now called semen—that is critical to first coagulate (thicken) and then de-coagulate the semen following ejaculation. The temporary thickening of semen helps retain it within the female reproductive tract, providing time for sperm to utilize the fructose provided by seminal vesicle secretions. When the semen regains its fluid state, sperm can then pass farther into the egg conducting organs.

The prostate normally doubles in size during puberty. At approximately age 25, it gradually begins to enlarge again. This enlargement does not usually cause problems; however, abnormal growth of the prostate, or benign prostatic hyperplasia (BPH), can cause constriction of the urethra as it passes through the middle of the prostate gland, leading to a number of lower urinary tract symptoms, such as a frequent and intense urge to urinate, a weak stream, and a sensation that the bladder has not emptied completely. By age 60, approximately 40 percent of men have some degree of BPH. By age 80, the number of affected individuals has jumped to as many as 80 percent. Treatments for BPH attempt to relieve the pressure on the urethra so that urine can flow more normally. Mild to moderate symptoms are treated with medication, whereas severe enlargement of the prostate is treated by surgery in which a portion of the prostate tissue is removed.

Another common disorder involving the prostate is prostate cancer. According to the Centers for Disease Control and Prevention (CDC), prostate cancer is the second most common cancer in those with SPC organs. However, some forms of prostate cancer grow very slowly and thus may not ever require treatment. Aggressive forms of prostate cancer, in contrast, involve metastasis to vulnerable organs like the lungs and brain. There is no link between BPH and prostate cancer, but the symptoms are similar. Prostate cancer is detected by a medical history, a blood test, and a rectal exam that allows physicians to palpate the prostate and check for unusual masses. If a mass is detected, the cancer diagnosis is confirmed by biopsy of the cells.

Bulbourethral Glands

The final addition to semen is made by two bulbourethral glands (or Cowper’s glands) that release a thick, salty fluid that lubricates the end of the urethra and the vagina, and helps to clean urine residues from the penile urethra. The fluid from these accessory glands is released after becoming sexually aroused, and shortly before the release of the semen. It is therefore sometimes called pre-ejaculate. It is important to note that, in addition to the lubricating proteins, it is possible for bulbourethral fluid to pick up sperm already present in the urethra, and therefore it may be able to cause pregnancy.

The Penis

The penis is the male organ of copulation (sexual intercourse). It is flaccid for non-sexual actions, such as urination, and turgid and rod-like with sexual arousal. When erect, the stiffness of the organ allows it to penetrate into the vagina and deposit semen.

The shaft of the penis surrounds the urethra. The shaft is composed of three column-like chambers of erectile tissue that span the length of the shaft. Each of the two larger lateral chambers is called a corpus cavernosum (plural = corpora cavernosa). Together, these make up the bulk of the penis. The corpus spongiosum, which can be felt as a raised ridge on the erect penis, is a smaller chamber that surrounds the spongy, or penile, urethra. The end of the penis, called the glans penis, has a high concentration of nerve endings, resulting in very sensitive skin that influences the likelihood of ejaculation. The skin from the shaft extends down over the glans and forms a collar called the prepuce (or foreskin). The foreskin also contains a dense concentration of nerve endings, and both lubricate and protect the sensitive skin of the glans penis. A surgical procedure called circumcision, often performed for religious or social reasons, removes the prepuce, typically within days of birth.

Both sexual arousal and REM sleep (during which dreaming occurs) can induce an erection. Penile erections are the result of vasocongestion, or engorgement of the tissues because of more arterial blood flowing into the penis than is leaving in the veins. During sexual arousal, nitric oxide (NO) is released from nerve endings near blood vessels within the corpora cavernosa and spongiosum. Release of NO activates a signaling pathway that results in relaxation of the smooth muscles that surround the penile arteries, causing them to dilate. This dilation increases the amount of blood that can enter the penis and induces the endothelial cells in the penile arterial walls to also secrete NO and perpetuate the vasodilation. The rapid increase in blood volume fills the erectile chambers, and the increased pressure of the filled chambers compresses the thin-walled penile venules, preventing venous drainage of the penis. The result of this increased blood flow to the penis and reduced blood return from the penis is erection. Depending on the flaccid dimensions of a penis, it can increase in size slightly or greatly during erection, with the average length of an erect penis measuring approximately 15 cm.

Testosterone

Testosterone, an androgen, is a steroid hormone produced by Interstitial cells (Leydig cells). The term “interstitial” reflects their location between the seminiferous tubules in the testes. In embryos, testosterone is secreted by Interstitial cells by the seventh week of development, with peak concentrations reached in the second trimester. This early release of testosterone results in the anatomical differentiation of the SPC organs. In childhood, testosterone concentrations are low. They increase during puberty, activating characteristic physical changes and initiating spermatogenesis.

Functions of Testosterone

The continued presence of testosterone is necessary to keep the SPC organs working properly, and Interstitial cells produce approximately 6 to 7 mg of testosterone per day. Testicular steroidogenesis (the manufacture of androgens, including testosterone) results in testosterone concentrations that are 100 times higher in the testes than in the circulation. Maintaining these normal concentrations of testosterone promotes spermatogenesis, whereas low levels of testosterone can lead to infertility. In addition to intra-testicular secretion, testosterone is also released into the systemic circulation and plays an important role in muscle development, bone growth, the development of secondary sex characteristics, and maintaining libido (sex drive). In those with EPC organs, the ovaries secrete small amounts of testosterone, although most is converted to estradiol. A small amount of testosterone is also secreted by the adrenal glands in both those with EPC and SPC organs.

Chapter Review

Gametes are the reproductive cells that combine to form offspring. Organs called gonads produce the gametes, along with the hormones that regulate human reproduction. The male gametes are called sperm. Spermatogenesis, the production of sperm, occurs within the seminiferous tubules that make up most of the testis. The scrotum is the muscular sac that holds the testes outside of the body cavity.

Spermatogenesis begins with mitotic division of spermatogonia (stem cells) to produce primary spermatocytes that undergo the two divisions of meiosis to become secondary spermatocytes, then the haploid spermatids. During spermiogenesis, spermatids are transformed into spermatozoa (formed sperm). Upon release from the seminiferous tubules, sperm are moved to the epididymis where they continue to mature. During ejaculation, sperm exit the epididymis through the ductus deferens, a duct in the spermatic cord that leaves the scrotum. The ampulla of the ductus deferens meets the seminal vesicle, a gland that contributes fructose and proteins, at the ejaculatory duct. The fluid continues through the prostatic urethra, where secretions from the prostate are added to form semen. These secretions help the sperm to travel through the urethra and into the female reproductive tract. Secretions from the bulbourethral glands protect sperm and cleanse and lubricate the penile (spongy) urethra.

The penis is the organ of copulation in those with SPC organs. Columns of erectile tissue called the corpora cavernosa and corpus spongiosum fill with blood when sexual arousal activates vasodilatation in the blood vessels of the penis. Testosterone regulates and maintains the sex organs and sex drive, and induces the physical changes of puberty. Interplay between the testes and the endocrine system precisely control the production of testosterone with a negative feedback loop.

Self Check

Answer the question(s) below to see how well you understand the topics covered in the previous section.

Glossary

blood–testis barrier: tight junctions between Sertoli cells that prevent bloodborne pathogens from gaining access to later stages of spermatogenesis and prevent the potential for an autoimmune reaction to haploid sperm

bulbourethral glands: (also, Cowper’s glands) glands that secrete a lubricating mucus that cleans and lubricates the urethra prior to and during ejaculation

corpus cavernosum: either of two columns of erectile tissue in the penis that fill with blood during an erection

corpus spongiosum: (plural = corpora cavernosa) column of erectile tissue in the penis that fills with blood during an erection and surrounds the penile urethra on the ventral portion of the penis

ductus deferens: (also, vas deferens) duct that transports sperm from the epididymis through the spermatic cord and into the ejaculatory duct; also referred as the vas deferens

ejaculatory duct: duct that connects the ampulla of the ductus deferens with the duct of the seminal vesicle at the prostatic urethra

epididymis: (plural = epididymides) coiled tubular structure in which sperm start to mature and are stored until ejaculation

gamete: haploid reproductive cell that contributes genetic material to form an offspring

glans penis: bulbous end of the penis that contains a large number of nerve endings

gonadotropin-releasing hormone (GnRH): hormone released by the hypothalamus that regulates the production of follicle-stimulating hormone and luteinizing hormone from the pituitary gland

gonads: reproductive organs (testes in those with SPC organs, ovaries in those with EPC organs) that produce gametes and reproductive hormones

inguinal canal: opening in abdominal wall that connects the testes to the abdominal cavity

Interstitial cells: cells between the seminiferous tubules of the testes that produce testosterone

penis: organ of copulation in those with SPC organs

prepuce: (also, foreskin) flap of skin that forms a collar around, and thus protects and lubricates, the glans penis; also referred as the foreskin

prostate gland: doughnut-shaped gland at the base of the bladder surrounding the urethra and contributing fluid to semen during ejaculation

scrotum: external pouch of skin and muscle that houses the testes

semen: ejaculatory fluid composed of sperm and secretions from the seminal vesicles, prostate, and bulbourethral glands

seminal vesicle: gland that produces seminal fluid, which contributes to semen

seminiferous tubules: tube structures within the testes where spermatogenesis occurs

sperm: (also, spermatozoon) male gamete

spermatic cord: bundle of nerves and blood vessels that supplies the testes; contains ductus deferens

spermatid: immature sperm cells produced by meiosis II of secondary spermatocytes

spermatocyte: cell that results from the division of spermatogonium and undergoes meiosis I and meiosis II to form spermatids

spermatogenesis: formation of new sperm, occurs in the seminiferous tubules of the testes

spermatogonia: (singular = spermatogonium) diploid precursor cells that become sperm

spermiogenesis: transformation of spermatids to spermatozoa during spermatogenesis

Sustentacular cells: cells that support germ cells through the process of spermatogenesis

testes: (singular = testis) sperm producing organs