REPRODUCTIVE SYSTEM

FUNCTIONS OF THE REPRODUCTIVE SYSTEM

The human species could not survive without functional male and female reproductive systems. The
reproductive systems play essential roles in the development of the structural and functional differences
between males and females, influence human behavior, and produce offspring. However, a reproductive
system, unlike other organ systems, is not necessary for the survival of an individual human.

The reproductive system performs the following functions:

1. Production of gametes. The reproductive system produces gametes: sperm cells in the testes of
males and oocytes (eggs) in the ovaries of females.

2. Fertilization. The reproductive system enhances fertilization of the oocyte by the sperm. The
duct system in male snourishes sperm until they are mature and are deposited in the female
reproductive tract by the penis. The female reproductive system receives the male’s sperm and
transports them to the fertilization site.

3. Development and nourishment of a new individual. The female reproductive system nurtures
the development of a new individual in the uterus until birth and provides nourishment (milk) after
birth.

4. Production of reproductive hormones. Hormones produced by the reproductive system control

its development and the development of the gender-specific body form. These hormones are also
essential for the normal function of the reproductive system and for reproductive behavior.

FORMATION OF GAMETES

The testes in males and the ovaries in females produce gametes or sex cells. The formation of gametes
in males and females occurs by a type of cell division called meiosis. Meiosis occurs only in the testis
and ovary. During meiosis, one cell undergoes two consecutive cell divisions to produce four daughter
cells, each having half as many chromosomes as the parent cell.

The two divisions of meiosis are called meiosis I and meiosis II. Like mitosis, each division of meiosis
has prophase, metaphase, anaphase, and telophase. However, there are distinct differences between
meiosis and mitosis.
Before meiosis begins, all the chromosomes are duplicated. At the beginning of meiosis, each of the 46
chromosomes consists of 2 chromatids connected by a centromere. The chromosomes align as pairs in
a process called synapsis. Because each chromosome consists of 2 chromatids, the pairing of the
chromosomes brings 2 chromatids of each chromosome close together. Occasionally, part of a
chromatid of 1 chromosome breaks off and is exchanged with part of another chromatid from the other
chromosome. This event, called crossing over, allows the exchange of genetic material between
chromosomes.

The chromosomes align along the center of the cell and then the pairs of chromosomes are separated
to each side of the cell. As a consequence, when meiosis I is complete, each daughter cell has 1
chromosome from each of the pairs or 23 chromosomes. Each of the 23 chromosomes in each daughter
cell consists of 2 chromatids joined by a centromere.

It is during the first meiotic division that the chromosome number is reduced from 46 (23 pairs) to 23
chromosomes. The first meiotic division is therefore called a reduction division.

The second meiotic division is similar to mitosis. The chromosomes, each consisting of 2 chromatids
align along the center of the cell. Then the chromatids separate at the centromere, and each daughter
cell receives 1 of the chromatids from each chromosome. When the centromere separates, each of the
chromatids is called a chromosome. Consequently, each of the 4 daughter cells produced by meiosis
contains 23 chromosomes.

During fertilization, the zygote receives 1 set of chromosomes (23) from each parent. Although half the
genetic material of a zygote comes from each parent, the genetic makeup of each zygote is unique.

MALE REPRODUCTIVE SYSTEM

The male reproductive system consists of the testes (sing. testis), a series of ducts, accessory glands,
and supporting structures. The ducts include the epididymides (sing. epididymis), the ducta deferentia
and the urethra. Accessory glands include the seminal vesicles, the prostate gland, and the
bulbourethral glands. Supporting structures include the scrotum and the penis (figure 19.3). The sperm
cells are very heat-sensitive and must develop at a temperature slightly less than normal body
temperature. The testes, in which the sperm cells develop, are located outside the body cavity in the
scrotum, where the temperature is lower. Sperm cells travel from each testis to the prostate gland and
then empty into the urethra within the prostate gland. The urethra exits the pelvis, passes through the
penis, and opens to the outside of the body.

Scrotum

The scrotum is a saclike structure containing the testes. It is divided into right and left internal
compartments by an incomplete connective tissue septum. Externally, the scrotum consists of skin.
Beneath the skin are a layer of loose connective tissue and a layer of smooth muscle called
the dartos muscle.

In cold temperatures, the dartos muscle contracts, causing the skin of the scrotum to become firm and
wrinkled and reducing the overall size of the scrotum. At the same time, extensions of abdominal
muscles into the scrotum, called cremaster muscles, contract. Consequently, the testes are pulled
nearer the body, and their temperature is elevated. During warm weather or exercise, the dartos and
cremaster muscles relax, the skin of the scrotum becomes loose and thin, and the testes descend away
from the body, which lowers their temperature. The response of the dartos and cremaster muscles is
important in regulating the temperature in the testes. If the testes become too warm or too cold, normal
sperm cell development does not occur.

Testes

The testes or male gonads, are oval organs, each about 4–5 cm long, within the scrotum. The outer part
of each testis consists of a thick, white connective tissue capsule. Extensions of the capsule project into
the interior of the testis and divide each testis into about 250 cone-shaped lobules. The lobules
contain seminiferous tubules, in which sperm cells develop. Delicate connective tissue surrounding
the seminiferous tubules contains clusters of endocrine cells called interstitial cells, or Leydig
cells, which secrete testosterone. The seminiferous tubules contain germ cells and sustentacular
cells, or Sertoli cells. Sustentacular cells are large and extend from the periphery to the lumen of the
seminiferous tubule. They nourish the germ cells and produce a number of hormones.

Spermatogenesis

Spermatogenesis is the formation of sperm cells. Before puberty, the testes remain relatively simple
and unchanged from the time of their initial development.

The interstitial cells are not prominent, and the seminiferous tubules are small and not yet functional.
At the time of puberty, the interstitial cells increase in number and size, the seminiferous tubules
enlarge, and spermatogenesis begins.

Germ cells are partially embedded in the sustentacular cells. The most peripheral germ cells
are spermatogonia, which divide through mitosis. Some daughter cells produced from these mitotic
divisions remain as spermatogonia and continue to divide by mitosis. Other daughter cells
form primary spermatocytes which divide by meiosis and become sperm cells.

Spermatogenesis

A primary spermatocyte contains 46 chromosomes, each consisting of 2 chromatids. Each primary

spermatocyte passes through the first meiotic division to produce 2 secondary spermatocytes. Each
secondary spermatocyte undergoes the second meiotic division to produce 2 smaller cells
called spermatids each having 23 chromosomes. After the second meiotic division, the spermatids
undergo major structural changes to form sperm cells. Much of the cytoplasm of the spermatids is
eliminated, and each spermatid develops a head, midpiece, and flagellum (tail) to become a sperm
cell, or spermatozoon nucleus of the sperm cell is located in the head of the sperm cell. Just anterior
to the nucleus is a vesicle called the acrosome which contains enzymes that are released during the
process of fertilization and are necessary for the sperm cell to penetrate the oocyte, or egg cell.

At the end of spermatogenesis, the developing sperm cells are located around the lumen of the
seminiferous tubules, with their heads directed toward the surrounding sustentacular cells and their
tails directed toward the center of the lumen. Finally, sperm cells are released into the lumen of the
seminiferous tubules.
Ducts

After their production, sperm cells are transported through the seminiferous tubules and a series of
ducts to the exterior of the body.

Epididymis
The seminiferous tubules of each testis empty into a tubular network called the rete testis. The rete
testis empties into 15–20 tubules called the efferent ductules. The efferent ductules carry sperm cells
from the testis to a tightly coiled series of threadlike tubules that form a comma-shaped structure on
the posterior side of the testis called the epididymis. The sperm cells continue to mature within the
epididymis, developing the capacity to swim and the ability to bind to the oocyte. Sperm cells taken
directly from the testes are not capable of fertilizing oocytes, but after maturing for several days in the
epididymis, the sperm cells develop the capacity to function as gametes. Final changes in sperm cells,
called capacitation occur after ejaculation of semen into the vagina and prior to fertilization.

Ductus Deferens
The ductus deferens or vas deferens, emerges from the epididymis and ascends along the posterior
side of the testis to become associated with the blood vessels and nerves that supply the testis. These
structures form the spermatic cord. Each spermatic cord consists of the ductus deferens, testicular
artery and veins, lymphatic vessels, and testicular nerve. It is surrounded by the cremaster muscle and
two connective tissue sheaths.

Each ductus deferens extends, in the spermatic cord, through the abdominal wall by way of the inguinal
canal. Each ductus deferens then crosses the lateral wall of the pelvic cavity and loops behind the
posterior surface of the urinary bladder to approach the prostate gland. The total length of the ductus
deferens is about 45 cm.

Just before reaching the prostate gland, the ductus deferens increases in diameter to become
the ampulla of the ductus deferens. The wall of the ductus deferens contains smooth muscle, which
contracts in peristaltic waves to propel the sperm cells from the epididymis through the ductus deferens.

Seminal Vesicle and Ejaculatory Duct

Near the ampulla of each ductus deferens is a sac-shaped gland called the seminal vesicle. A short
duct extends from the seminal vesicle to the ampulla of the ductus deferens. The ducts from the seminal
vesicle and the ampulla of the ductus deferens join at the prostate gland to form
the ejaculatory duct. Each ejaculatory duct extends into the prostate gland and ends by joining the
urethra within the prostate gland.

Urethra
The male urethra extends from the urinary bladder to the distal end of the penis. The urethra
can be divided into three parts: the prostatic urethra, which passes through the prostate gland;
the membranous urethra, which passes through the floor of the pelvis and is surrounded by the
external urinary sphincter; and the spongy urethra, which extends the length of the penis and opens
at its end. The urethra is a passageway for both urine and male reproductive fluids. However, urine and
the reproductive fluids do not exit the urethra at the same time. While male reproductive fluids are
passing through the urethra, a sympathetic reflex causes the internal urinary sphincter to contract, which
keeps semen from passing into the urinary bladder and prevents urine from entering the urethra.
Penis

The penis is the male organ of copulation and functions in the transfer of sperm cells from the male to
the female. The penis contains three columns of erectile tissue. Engorgement of this erectile tissue with
blood causes the penis to enlarge and become firm, a process called erection. Two columns of erectile
tissue form the dorsal portion and the sides of the penis and are called the corpora cavernosa. The
third, smaller erectile column occupies the ventral portion of the penis and is called the corpus
spongiosum. It expands over the distal end of the penis to form a cap, the glans penis. The spongy
urethra passes through the corpus spongiosum, including the glans penis, and opens to the exterior as
the external urethral orifice.

The shaft of the penis is covered by skin that is loosely attached to the connective tissue surrounding
the penis. The skin is firmly attached at the base of the glans penis, and a thinner layer of skin tightly
covers the glans penis. The skin of the penis, especially the glans penis, is well supplied with sensory
receptors. A loose fold of skin, called the prepuce, or foreskin, covers the glans penis.

Glands

The seminal vesicles are glands consisting of many saclike structures located next to the ampulla of the
ductus deferens. There are two seminal vesicles.

Each is about 5 cm long and tapers into a short duct that joins the ampulla of the ductus deferens to
form the ejaculatory duct, as previously mentioned.

The prostate gland consists of both glandular and muscular tissue and is about the size and shape of
a walnut. The prostate gland surrounds the urethra and the two ejaculatory ducts. It consists of a capsule
and numerous partitions. The cells lining the partitions secrete prostatic fluid. There are 10–20 short
ducts that carry secretions of the prostate gland to the prostatic urethra.

The bulbourethral glands, or Cowper glands, are a pair of small, mucus-secreting glands located near
the base of the penis. In young adults, each is about the size of a pea, but they decrease in size with
age. A single duct from each gland enters the urethra.

Secretions

Semen is a mixture of sperm cells and secretions from the male reproductive glands. The seminal
vesicles produce about 60% of the fluid, the prostate gland contributes approximately 30%, the testes
contribute 5%, and the bulbourethral glands contribute 5%.

The bulbourethral glands and the mucous glands in the urethra produce a mucous secretion, which
lubricates the urethra, helps neutralize the contents of the normally acidic urethra, provides a small
amount of lubrication during intercourse, and helps reduce acidity in the vagina.

Testicular secretions include sperm cells and a small amount of fluid. The thick, mucus like secretion of
the seminal vesicles contains the sugar fructose and other nutrients that nourish sperm cells. The
seminal vesicle secretions also contain proteins that weakly coagulate after ejaculation and enzymes
that are thought to help destroy abnormal sperm cells. Prostaglandins, which stimulate smooth muscle
contractions, are present in high concentrations in the secretions of the seminal vesicles and can cause
contractions of the female reproductive tract, which help transport sperm cells through the tract.

The thin, milky secretions of the prostate have an alkaline pH and help neutralize the acidic urethra, as
well as the acidic secretions of the testes, the seminal vesicles, and the vagina. The increased pH is
important for normal sperm cell function. The movement of sperm cells is not optimal until the pH is
increased to between 6.0 and 6.5. In contrast, the pH of vaginal secretions is between 3.5 and 4.0.
Prostatic secretions also contain proteolytic enzymes that break down the coagulated proteins of the
seminal vesicles and make the semen more liquid. The normal volume of semen is 2–5 milliliters (mL),
with each milliliter of semen typi-cally containing about 100 million sperm cells.

PHYSIOLOGY OF MALE REPRODUCTION

The male reproductive system depends on both hormonal and neural mechanisms to function normally.
Hormones control the development of reproductive structures, the development of secondary sexual
characteristics, spermatogenesis, and some aspects of sexual behavior. The mature neural mechanisms
are primarily involved in controlling the sexual act and in the expression of sexual behavior.

Puberty In Males

Puberty is the sequence of events by which a child is transformed into a young adult. The reproductive
system matures and assumes its adult functions, and the structural differences between adult males
and females become more apparent.

In boys, puberty commonly begins between the ages of 12 and 14 and is largely completed by age 18.
Before puberty, small amounts of testosterone, secreted by the testes and the adrenal cortex, inhibit
GnRH, LH, and FSH secretion. Beginning just before puberty and continuing throughout puberty,
developmental changes in the hypothalamus cause the hypothalamus and the anterior pituitary gland
to become much less sensitive to the inhibitory effect of testosterone, and the rate of GnRH, LH, and
FSH secretion increases. Consequently, elevated FSH levels promote spermatogenesis, and elevated LH
levels cause the interstitial cells to secrete larger amounts of testosterone. Testosterone still has a
negative-feedback effect on the hypothalamus and anterior pituitary gland, but GnRH, LH, and FSH
secretion occurs at substantially higher levels.

Effects of Testosterone
Testosterone is the major male hormone secreted by the testes. Testosterone influences reproductive
organs and nonreproductive structures. During puberty, testosterone causes the enlargement and
differentiation of the male genitals and the reproductive duct system. It is necessary for
spermatogenesis and for the development of male secondary sexual characteristics. The secondary
sexual characteristics are those structural and behavioral changes, other than in the reproductive
organs, that develop at puberty and distinguish males from females.

.
Secondary sexual characteristics in males include hair distribution and growth, skin texture, fat
distribution, skeletal muscle growth, and changes in the larynx. After puberty, testosterone maintains
the adult structure of the male genitals, reproductive ducts, and secondary sexual characteristics.

Male Sexual Behavior And The Male Sex Act

Testosterone is required for normal sexual behavior. Testosterone enters certain cells within the
brain, especially within the hypo-thalamus, and influences their functions. The blood levels of
testosterone remain relatively constant throughout the lifetime of a male, from puberty until about 40
years of age. Thereafter, the levels slowly decline to approximately 20% of this value by 80 years of age,
causing a slow decrease in sex drive and fertility.

The male sex act is a complex series of reflexes that result in erection of the penis, secretion of
mucus into the urethra, emission, and ejaculation. Emission is the movement of sperm cells, mucus,
prostatic secretions, and seminal vesicle secretions into the prostatic, membranous, and spongy
urethra. Ejaculation is the forceful expulsion of the secretions that have accumulated in the urethra to
the exterior.

Sensations, normally interpreted as pleasurable, occur during the male sex act and result in an
intense sensation called an orgasm or climax. In males, orgasm is closely associated with ejaculation,
although they are separate functions and do not always occur simultaneously. A phase
called resolution occurs after ejaculation. During resolution, the penis becomes flaccid, an overall
feeling of satisfaction exists, and the male is unable to achieve erection and a second ejaculation.

Sensory Impulses and Integration

Sensory action potentials from the genitals are carried to the sacral region of the spinal cord,
where reflexes that result in the male sex act are integrated. Action potentials also travel from the spinal
cord to the cerebrum to produce conscious sexual sensations. Rhythmic massage of the penis, especially
the glans, and sur-rounding tissues, such as the scrotal, anal, and pubic regions, pro-vide important
sources of sensory action potentials. Engorgement of the prostate gland and seminal vesicles with
secretions or irritation of the urethra, urinary bladder, ducta deferentia, and testes can also cause sexual
sensations.

Psychological stimuli, such as sight, sound, odor, or thoughts, have a major effect on male sexual
reflexes. Ejaculation while sleeping (nocturnal emission) is a relatively common event in young males
and is thought to be triggered by psychological stimuli associated with dreaming.

Erection, Emission, and Ejaculation

Erection is the first major component of the male sex act. Parasympathetic action potentials
from the sacral region of the spinal cord cause the arteries that supply blood to the erectile tissues to
dilate. Blood then fills small venous sinuses called sinusoids in the erectile tissue and compresses the
veins, which reduces blood flow from the penis. The increased blood pressure in the sinusoids causes
the erectile tissue to become inflated and rigid. Parasympathetic action potentials also cause the
mucous glands within the urethra and the bulbourethral glands to secrete mucus.
Failure to achieve erections, or erectile dysfunction (ED), sometimes called impotence, can be a major
source of frustration. The inability to achieve erections can be due to reduced testosterone secretion
resulting from hypothalamic, pituitary, or testicular complications. In other cases, ED can be due to
defective stimulation of the erectile tissue by nerve fibers or reduced response of the blood vessels to
neural stimulation. Some men can achieve erections by taking oral medications, such as sildenafil
(Viagra), tadalafil (Cialis), or verdenafil (Livitra), or by having specific drugs injected into the base of the
penis. These drugs increase blood flow into the erectile tissue of the penis, resulting in erection for
many minutes.

Before ejaculation, the ductus deferens begins to contract rhythmically, propelling sperm cells and
testicular fluid from the epididymis through the ductus deferens. Contractions of the ductus deferens,
seminal vesicles, and ejaculatory ducts cause the sperm cells, testicular secretions, and seminal fluid to
move into the urethra, where they mix with prostatic secretions released by contraction of the prostate.

Emission is stimulated by sympathetic action potentials that originate in the lumbar region of the spinal
cord. Action potentials cause the reproductive ducts to contract and stimulate the seminal vesicles and
the prostate gland to release secretions. Consequently, semen accumulates in the urethra.

Ejaculation results from the contraction of smooth muscle in the wall of the urethra and skeletal muscles
surrounding the base of the penis. Just before ejaculation, action potentials are sent to the skeletal
muscles that surround the base of the penis. Rhythmic contractions are produced that force the semen
out of the urethra, resulting in ejaculation. In addition, muscle tension increases throughout the body.

Infertility In Males

Infertility is reduced or diminished fertility. Th emost common cause of infertility in males is a low
sperm cell count. If the sperm cell count drops to below 20 million sperm cells per milliliter, the male is
usually sterile.

The sperm cell count can decrease because of damage to the testes as a result of trauma, radiation,
cryptorchidism or infections, such as mumps, which block the ducts in the epididymis. Reduced sperm
cell counts can also result from inadequate secretion of luteinizing hormone and follicle-stimulating
hormone, which can be caused by hypothyroidism, trauma to the hypothalamus, infarctions of the
hypothalamus or anterior pituitary gland, or tumors. Decreased testosterone secretion reduces the
sperm cell count as well.

Even when the sperm cell count is normal, fertility can be reduced if sperm cell structure is abnormal,
as occurs due to chromosomal abnormalities caused by genetic factors. Reduced sperm cell motility
also results in infertility. A major cause of reduced sperm cell motility is the presence of antisperm
antibodies, which are produced by the immune system and bind to sperm cells.

In cases of infertility due to low sperm cell count or reduced motility, fertility can sometimes be achieved
by collecting several ejaculations, concentrating the sperm cells, and inserting them into the female’s
reproductive tract, a process called artificial insemination.
FEMALE REPRODUCTIVE SYSTEM

The female reproductive organs consist of the ovaries, the uterine tubes (or fallopian tubes), the uterus,
the vagina, the external genitalia, and the mammary glands. The internal reproductive organs of the
female are located within the pelvis, between the urinary bladder and the rectum. The uterus and the
vagina are in the midline, with an ovary to each side of the uterus. The internal reproductive organs are
held in place within the pelvis by a group of ligaments. The most conspicuous is the broad
ligament, which spreads out on both sides of the uterus and attaches to the ovaries and uterine tubes.

Ovaries

The two ovaries (are small organs suspended in the pelvic cavity by ligaments. The suspensory
ligament extends from each ovary to the lateral body wall, and the ovarian ligament attaches the
ovary to the superior margin of the uterus. In addition, the ovaries are attached to the posterior surface
of the broad ligament by folds of peritoneum called the mesovarium. The ovarian arteries, veins, and
nerves traverse the suspensory ligament and enter the ovary through the mesovarium.

A layer of visceral peritoneum covers the surface of the ovary. The outer part of the ovary is composed
of dense connective tissue and contains ovarian follicles. Each of the ovarian follicles contains
an oocyte the female sex cell. Loose connective tissue makes up the inner part of the ovary, where
blood vessels, lymphatic vessels, and nerves are located.

Oogenesis and Fertilization

The formation of female gametes begins during fetal development, even before the female is born. By
the fourth month of development, the ovaries contain 5 million oogonia the cells from which oocytes
develop. By the time of birth, many of the oogonia have degenerated, and the remaining ones have
begun meiosis. Also, some data indicate that oogonia can form after birth from stem cells, but the
extent to which this occurs, and how long it occurs, is not clear. As in meiosis in males, the genetic
material is duplicated, and two cell divisions occur. Meiosis stops, however, during the first meiotic
division at prophase I. The cell at this stage is called a primary oocyte, and at birth there are about 2
million of them. From birth to puberty, many primary oocytes degenerate. The number of primary
oocytes decreases to around 300,000 to 400,000; of these, only about 400 will complete development
and be released from the ovaries. Nearly all others degenerate after partial development.

Ovulation is the release of an oocyte from an ovary. Just before ovulation, the primary oocyte
completes the first meiotic division to produce a secondary oocyte and a polar body. Unlike meiosis
in males, cytoplasm is not split evenly between the two cells. Most of the cytoplasm of the primary
oocyte remains with the secondary oocyte. The polar body either degenerates or divides to form two
polar bodies. The secondary oocyte begins the second meiotic division but stops in metaphase II.

After ovulation, the secondary oocyte may be fertilized by a sperm cell. Fertilization begins when a
sperm cell penetrates the cytoplasm of a secondary oocyte. Subsequently, the secondary oocyte
completes the second meiotic division to form 2 cells, each containing 23 chromosomes. One of these
cells has very little cytoplasm and is another polar body that degenerates. In the other, larger cell, the
23 chromosomes from the sperm join with the 23 from the female gamete to form a zygote and
complete fertilization.

The zygote has 23 pairs of chromosomes (a total of 46 chromosomes). All cells of the human body
contain 23 pairs of chromosomes, except for the male and female gametes. The zygote divides by
mitosis to form 2 cells, which divide to form 4 cells, and so on. The mass of cells formed may eventually
implant in, or attach to, the uterine wall and develop into a new individual .
Follicle Development

As we discussed, when a female is in her mother’s uterus, her ovaries have already begun oocyte
formation. The primary oocytes present at birth are surrounded by a primordial follicle. A primordial
follicleis a primary oocyte surrounded by a single layer of flat cells, called granulosa cells puberty
begins, some of the primordial follicles are converted to primary follicles when the oocyte enlarges
and the single layer of granulosa cells becomes enlarged and cuboidal. Subsequently, several layers of
granulosa cells form, and a layer of clear material called the zona pellucida is deposited around the
primary oocyte.

Approximately every 28 days, hormonal changes stimulate some of the primary follicles to continue to
develop. The primary follicle becomes a secondary follicle as fluid-filled spaces called vesicles form
among the granulosa cells, and a capsule called the theca forms around the follicle.

The secondary follicle continues to enlarge, and when the fluid-filled vesicles fuse to form a single, fluid-
filled chamber called the antrum, the follicle is called the mature follicle, or graafian follicle. The
primary oocyte is pushed off to one side and lies in a mass of granulosa cells called the cumulus cells.

The mature follicle forms a lump on the surface of the ovary. During ovulation, the mature follicle
ruptures, forcing a small amount of blood, follicular fluid, and the secondary oocyte, surround-ed by
the cumulus cells, into the peritoneal cavity. In most cases, only one of the follicles that begin to develop
forms a mature follicle and undergoes ovulation. The other follicles degenerate. After ovulation, the
remaining cells of the ruptured follicle are transformed into a glandular structure called the corpus
luteum. If pregnancy occurs, the corpus luteum enlarges in response to a hormone secreted by the
placenta called human chorionic gonadotropin hormone (hCG). If pregnancy does not occur, the
corpus luteum lasts for 10–12 days and then begins to degenerate.

Uterine tubes

A uterine tube, also called a fallopian tube or oviduct is associated with each ovary. The uterine tubes
extend from the area of the ovaries to the uterus. They open directly into the peritoneal cavity near
each ovary and receive the secondary oocyte. The opening of each uterine tube is surrounded by long,
thin processes called fimbriae.

The fimbriae nearly surround the surface of the ovary. As a result, as soon as the secondary oocyte is
ovulated, it comes into contact with the surface of the fimbriae. Cilia on the fimbriae surface sweep the
oocyte into the uterine tube. Fertilization usually occurs in the part of the uterine tube near the ovary,
called the ampulla. The fertilized oocyte then travels to the uterus, where it embeds in the uterine wall
in a process called implantation.

Uterus

The uterus is as big as a medium-sized pear. It is oriented in the pelvic cavity with the larger, rounded
part directed superiorly. The part of the uterus superior to the entrance of the uterine tubes is called
the fundus. The main part of the uterus is called the body, and the narrower part, the cervix is directed
inferiorly. Internally, the uterine cavity in the fundus and uterine body continues through the cervix as
the cervical canal, which opens into the vagina. The cervical canal is lined by mucous glands.

The uterine wall is composed of three layers: a serous layer, a muscular layer, and a layer of
endometrium. The outer layer, called the perimetrium or serous layer, of the uterus is formed from
visceral peritoneum. The middle layer, called the myometrium or muscular layer, consists of smooth
muscle, is quite thick, and accounts for the bulk of the uterine wall. The innermost layer of the uterus is
the endometrium which consists of simple columnar epithelial cells with an underlying connective
tissue layer. Simple tubular glands, called spiral glands, are formed by folds of the endometrium. The
superficial part of the endometrium is sloughed off during menstruation.

The uterus is supported by the broad ligament and the round ligament. In addition to these ligaments,
much support is provided inferiorly to the uterus by skeletal muscles of the pelvic floor. If ligaments
that support the uterus or muscles of the pelvic floor are weakened, as may occur due to childbirth, the
uterus can extend inferiorly into the vagina, a condition called a prolapsed uterus. Severe cases require
surgical correction.

Vagina

The vagina is the female organ of copulation; it receives the penis during intercourse. It also allows
menstrual flow and childbirth. The vagina extends from the uterus to the outside of the body. The
superior portion of the vagina is attached to the sides of the cervix, so that a part of the cervix extends
into the vagina.

The wall of the vagina consists of an outer muscular layer and an inner mucous membrane. The muscular
layer is smooth muscle and contains many elastic fibers. Thus, the vagina can increase in size to
accommodate the penis during intercourse, and it can stretch greatly during childbirth. The mucous
membrane is moist stratified squamous epithelium that forms a protective surface layer. Lubricating
fluid passes through the vaginal epithelium into the vagina.

In young females, the vaginal opening is covered by a thin mucous membrane called the hymen. In
rare cases, the hymen may completely close the vaginal orifice and it must be removed to allow
menstrual flow. More commonly, the hymen is perforated by one or several holes. The openings in the
hymen are usually greatly enlarged during the first sexual intercourse. The hymen can also be perforated
or torn earlier in a young female’s life during a variety of activities, including strenuous exercise. The
condition of the hymen is therefore an unreliable indicator of virginity.

The region between the vagina and the anus is the clinical perineum. The skin and muscle of this
region can tear during childbirth. To prevent such tearing, an incision called an episiotomy is
sometimes made in the clinical perineum. Traditionally, this clean, straight incision has been thought to
result in less injury, less trouble in healing, and less pain. However, many studies report less injury and
pain when no episiotomy is performed.

Mammary Glands

The mammary glands are the organs of milk production and are located in the breasts. The mammary
glands are modified sweat glands. Externally, each of the breasts of both males and females has a
raised nipple surrounded by a circular, pigmented area called the areola

In prepubescent children, the general structure of the male and female breasts is similar, and both males
and females possess a rudimentary duct system. The female breasts begin to enlarge during puberty,
under the influence of estrogen and progesterone. Some males also experience a minor and temporary
enlargement of the breasts at puberty. Occasionally, the breasts of a male can become permanently
enlarged, a condition called gynecomastia. Causes of gynecomastia include hormonal imbalances and
the abuse of anabolic steroids.

Each adult female breast contains mammary glands consisting of usually 15–20
glandular lobes covered by adipose tissue. It is primarily this superficial adipose tissue that gives the
breast its form. Each lobe possesses a single lactiferous duct that opens independently to the surface
of the nipple. The duct of each lobe is formed as several smaller ducts, which originate
from lobules, converge. Within a lobule, the ducts branch and become even smaller. In the milk-
producing, or lactating, mammary gland, the ends of these small ducts expand to form secretory sacs
called alveoli. Myoepithelial cells sur-round the alveoli and contract to expel milk from the alveoli.

The breasts are supported by suspensory ligaments that extend from the fascia over the pectoralis
major muscles to the skin over the breasts

The nipples are very sensitive to tactile stimulation and contain smooth muscle. When the smooth
muscle contracts in response to stimuli, such as touch, cold, and sexual arousal, the nipple becomes
erect.

Physiology of Female Reproduction

Puberty in Females

The initial change that results in puberty is most likely maturation of the hypothalamus. In girls, puberty,
which typically begins between ages 11 and 13 and is largely completed by age 16, is marked by the
first episode of menstrual bleeding, which is called menarche. During puberty, the vagina, uterus,
uterine tubes, and external genitalia begin to enlarge. Adipose tissue is deposited in the breasts and
around the hips, causing them to enlarge and assume an adult form. In addition, pubic and axillary hair
grows. The development of sexual drive is also associated with puberty.

The changes associated with puberty primarily result from the increasing rate of estrogen and
progesterone secretion by the ovaries. Before puberty, estrogen and progesterone are secreted in very
small amounts. At puberty, the cyclical adult pattern of hormone secretion is gradually established.

Before puberty, the rate of GnRH secretion from the hypo-thalamus and the rate of LH and FSH
secretion from the anterior pituitary are very low. Estrogen and progesterone from the ovaries have a
strong negative-feedback effect on the hypothalamus and pituitary. After the onset of puberty, the
hypothalamus and anterior pituitary secrete larger amounts of GnRH, LH, and FSH. Estrogen and
progesterone have less of a negative-feedback effect on the hypothalamus and pituitary, and a
sustained increase in estrogen concentration has a positive-feedback effect. The normal cyclical pattern
of reproductive hormone secretion that occurs during the menstrual cycle becomes established.

Menstrual Cycle

The term menstrual cycle refers to the series of changes that occur in sexually mature, nonpregnant
females and that culminate in menses. Menses a period of mild hemorrhage, during which part of the
endometrium is sloughed and expelled from the uterus. Typically, the menstrual cycle is about 28 days
long, although it can be as short as 18 days or as long as 40 days. The menstrual cycle results from the
cyclical changes that occur in the endometrium of the uterus. These changes, in turn, result from the
cyclical changes that occur in the ovary and are controlled by the secretions of FSH and LH from the
anterior pituitary gland.
The first day of menstrual bleeding (menses), when the endometrium sloughs off, is considered day 1
of the menstrual cycle. Sloughing of the endometrium is inhibited by progesterone but stimulated by
estrogen. Menses typically lasts 4 or 5 days and can be accompanied by strong uterine contractions,
called menstrual cramps, that are extremely uncomfortable in some women. Menstrual cramps are the
result of forceful myometrial contractions that occur before and during menstruation. The cramps can
result from excessive secretion of prostaglandins. As the endometrium of the uterus sloughs off, it
becomes inflamed, and prostaglandins are produced as part of the inflammation. Many women can
alleviate painful cramps by taking medications, such as aspirin-like drugs, that inhibit prostaglandin
biosynthesis just before the onset of menstruation. These medications, however, are not effective in
treating all painful menstruation, especially when the cause of the pain, such as that experienced by
women who have tumors of the myometrium, is not due to the inflammatory response.

Ovulation occurs on about day 14 of the menstrual cycle, although the timing of ovulation varies from
individual to individual and can vary within an individual from one menstrual cycle to the next. To avoid
or optimize contraception, it is critical to predict ovulation; however, there is no single reliable method
that can predict its exact timing. The simplest method of predicting ovulation is looking for a drop in
basal body temperature preceding ovulation, but it is the least reliable method.

Between the end of menses and ovulation is the proliferative phase, which refers to proliferation of
the endometrium. During the proliferative phase, the secondary follicles in the ovary mature; as they
do so, they secrete increasing amounts of estrogen. Estrogen acts on the uterus and causes the
epithelial cells of the endometrium to divide rapidly. The endometrium thickens, and spiral glands form.

The sustained increase of estrogen secreted by the developing follicles stimulates GnRH secretion from
the hypothalamus. GnRH, in turn, triggers FSH and LH secretion from the anterior pituitary gland. FSH
stimulates estrogen secretion at an increasing rate from the developing follicles. This positive-feedback
loop produces a series of larger and larger surges of FSH and LH secretion. Ovulation occurs in response
to the large increases in LH levels that normally occur on about day 14 of the menstrual cycle. This large
increase in LH is also responsible for the development of the corpus luteum.

Following ovulation, the corpus luteum begins to secrete progesterone and smaller amounts of
estrogen. Progesterone acts on the uterus, causing the cells of the endometrium to become larger and
to secrete a small amount of fluid. Together, progesterone and estrogen act on the hypothalamus and
anterior pituitary gland to inhibit GnRH, LH, and FSH secretion. Thus, LH and FSH levels decline after
ovulation.

Between ovulation and the next menses is the secretory phase of the menstrual cycle, called this
because of the small amount of fluid secreted by the cells of the endometrium. During the secretory
phase, the lining of the uterus reaches its greatest degree of development.

If fertilization occurs, the zygote undergoes several cell divisions to produce a collection of cells called
the blastocyst. The blastocyst passes through the uterine tube and arrives in the uterus by 7 or 8 days
after ovulation. The endometrium is prepared to receive the blastocyst, which becomes implanted in
the endometrium, where it continues to develop. If the secondary oocyte is not fertilized, the
endometrium sloughs away as a result of declining blood progesterone levels. Unless the secondary
oocyte is fertilized, the corpus luteum begins to produce less progesterone by day 24 or 25 of the
menstrual cycle.

By day 28, the declining progesterone causes the endometrium to slough away to begin menses and
the next menstrual cycle. The declining progesterone secretion results in a small increase in FSH
secretion at the beginning of the next menses, which triggers more follicles to mature.

An ectopic pregnancy results if implantation occurs anywhere other than in the uterine cavity. The
most common site of ectopic pregnancy is the uterine tube. Implantation in the uterine tube is
eventually fatal to the fetus and can cause the tube to rupture. In some rare cases, implantation occurs
in the mesenteries of the abdominal cavity; the fetus can develop normally but must be delivered by
caesarean section. However, maternal mortality rates for abdominal pregnancies are significantly higher
than for fallopian tube ectopic pregnancies.
Menopause

When a woman is 40–50 years old, the menstrual cycles become less regular, and ovulation does not
occur consistently during each cycle. Eventually, the cycles stop completely. The cessation of menstrual
cycles is called menopause and the whole time period from the onset of irregular cycles to their
complete cessation is called the female climacteric.

The major cause of menopause is age-related changes in the ovaries. The number of follicles remaining
in the ovaries of menopausal women is small. In addition, the follicles that remain become less sensitive
to stimulation by FSH and LH, and therefore fewer mature follicles and corpora lutea are produced.

Older women experience gradual changes in response to the reduced amount of estrogen and
progesterone produced by the ovaries (table 19.4). For example, during the climacteric, some women
experience sudden episodes of uncomfortable sweating (hot flashes), fatigue, anxiety, temporary
decrease in libido, and occasionally emotional disturbances. Many of these symptoms can be treated
successfully with hormone replacement therapy (HRT), which usually consists of small amounts of
estrogen and progester-one. HRT has been linked to a slightly increased risk of developing breast
cancer, uterine cancer, heart attack, stroke, or blood clots. On the positive side, HRT slows the decrease
in bone density that can become severe in some women after menopause, and it decreases the risk of
developing colorectal cancer.

Female sexual Behavior and the Female Sex Act

Sexual drive in females, like sexual drive in males, is dependent on hormones. Testosterone-like
hormones, and possibly estrogen, affect brain cells (especially in the area of the hypothalamus) and
influence sexual behavior. Testosterone-like hormones are produced primarily in the adrenal cortex.
Psychological factors also play a role in sexual behavior. The sensory and motor neural pathways
involved in control-ling female sexual responses are similar to those found in the male.

During sexual excitement, erectile tissue within the clitoris and around the vaginal opening becomes
engorged with blood. The mucous glands within the vestibule, especially the greater vestibular glands,
secrete small amounts of mucus. Larger amounts of mucus-like fluid are also extruded into the vagina
through its wall. These secretions provide lubrication to allow easy entry and movement of the penis in
the vagina during intercourse. Tactile stimulation of the female’s genitals during sexual intercourse and
psychological stimuli normally trigger an orgasm, or climax. The vaginal and uterine smooth muscle,
as well as the surrounding skeletal muscles, con-tract rhythmically, and muscle tension increases
throughout much of the body. After the sex act, there is a period of resolution, which is characterized
by an overall sense of satisfaction and relaxation. Females are sometimes receptive to further immediate
stimulation, however, and can experience successive orgasms. Orgasm is not necessary for fertilization
to occur. Ovulation results from hormonal stimuli and is not dependent on the female sex act.