Learning Objectives

In this section, you will explore the following questions:

What are the features of the human male and female reproductive anatomies?
What are the events associated with the human sexual response?
What are the differences and similarities between spermatogenesis and oogenesis?

Connection for AP® Courses

Connection for AP^® Courses

Much of the information in this section is not within the scope for AP^®. You do not need to match a list of the different anatomical features of the male and female reproductive systems with their functions, but you likely will find the material informative and relevant. The reproductive structures that evolved in land animals, including humans, allow males and females to mate, fertilize internally, and support the growth and development of offspring. The section provides an opportunity to review the process of meiosis that we first explored in a previous chapter.

Information presented and the examples highlighted in the section support concepts outlined in Big Idea 3 of the AP^® Biology Curriculum Framework. The AP^® Learning Objectives listed in the Curriculum Framework provide a transparent foundation for the AP^® Biology course, an inquiry-based laboratory experience, instructional activities, and AP^® exam questions. A learning objective merges required content with one or more of the seven science practices.

Big Idea 3	Living systems store, retrieve, transmit and respond to information essential to life processes.
Enduring Understanding 3.A	Heritable information provides for continuity of life.
Essential Knowledge	3.A.2 In eukaryotes, heritable information is passed to the next generation via processes that include the cell cycle and mitosis or meiosis plus fertilization.
Science Practice	6.2 The student can construct explanations of phenomena based on evidence produced through scientific practices.
Learning Objective	3.9 The student is able to construct an explanation, using visual representations or narratives, as to how DNA in chromosomes is transmitted to the next generation via mitosis, or meiosis followed by fertilization.
Essential Knowledge	3.A.2 In eukaryotes, heritable information is passed to the next generation via processes that include the cell cycle and mitosis or meiosis plus fertilization.
Science Practice	7.1 The student can connect phenomena and models across spatial and temporal scales.
Learning Objective	3.10 The student is able to represent the connection between meiosis and increased genetic diversity necessary for evolution.

Human Reproductive Anatomy

The reproductive tissues of male and female humans develop similarly in utero until a low level of the hormone testosterone is released from male gonads. Testosterone causes the undeveloped tissues to differentiate into male sexual organs. When testosterone is absent, the tissues develop into female sexual tissues. Primitive gonads become testes or ovaries. Tissues that produce a penis in males produce a clitoris in females. The tissue that will become the scrotum in a male becomes the labia in a female; that is, they are homologous structures.

Male Reproductive Anatomy

In the male reproductive system, the scrotum houses the testicles or testes (singular: testis), including providing passage for blood vessels, nerves, and muscles related to testicular function. The testes are a pair of male reproductive organs that produce sperm and some reproductive hormones. Each testis is approximately 2.5 by 3.8 cm (1.5 by 1 in) in size and divided into wedge-shaped lobules by connective tissue called septa. Coiled in each wedge are seminiferous tubules that produce sperm.

Sperm are immobile at body temperature; therefore, the scrotum and penis are external to the body, as illustrated in Figure 34.10 so that a proper temperature is maintained for motility. In land mammals, the pair of testes must be suspended outside the body at about 2 °C lower than body temperature to produce viable sperm. Infertility can occur in land mammals when the testes do not descend through the abdominal cavity during fetal development.

Visual Connection

Illustration shows a cross section of the penis and testes. The penis widens at the end, into the glans, which is surrounded by the foreskin. The urethra is an opening that runs through the middle of the penis to the bladder. The tissue surrounding the urethra is the Corpus spongiosum, and above the Corpus spongiosum is the Corpus cavernosum. The testes, located immediately behind the penis, are covered by the scrotum. Seminiferous tubules are located in the testes. The epididymis partly surrounds the sac

Figure 34.10 The reproductive structures of the human male are shown.

Which of the following statements about the male reproductive system is FALSE?

The vas deferens carries sperm from the testes to the penis.
Sperm mature in seminiferous tubules in the testes.
Both the prostate and the bulbourethral glands produce parts of the semen.
The prostate gland is located in the testes.

Sperm mature in seminiferous tubules that are coiled inside the testes, as illustrated in Figure 34.10. The walls of the seminiferous tubules are made up of the developing sperm cells, with the least developed sperm at the periphery of the tubule and the fully developed sperm in the lumen. The sperm cells are mixed with nursemaid cells called Sertoli cells, which protect the germ cells and promote their development. Other cells mixed in the wall of the tubules are the interstitial cells of Leydig. These cells produce high levels of testosterone once the male reaches adolescence.

When the sperm have developed flagella and are nearly mature, they leave the testicles and enter the epididymis, shown in Figure 34.10. This structure resembles a comma and lies along the top and posterior portion of the testes; it is the site of sperm maturation. The sperm leave the epididymis and enter the vas deferens (or ductus deferens), which carries the sperm, behind the bladder, and forms the ejaculatory duct with the duct from the seminal vesicles. During a vasectomy, a section of the vas deferens is removed, preventing sperm from being passed out of the body during ejaculation and preventing fertilization.

Semen is a mixture of sperm and spermatic duct secretions (about 10 percent of the total) and fluids from accessory glands that contribute most of the semen’s volume. Sperm are haploid cells, consisting of a flagellum as a tail, a neck that contains the cell’s energy-producing mitochondria, and a head that contains the genetic material. Figure 34.11 shows a micrograph of human sperm as well as a diagram of the parts of the sperm. An acrosome is found at the top of the head of the sperm. This structure has lysosomal enzymes that can digest the protective coverings that surround the egg to help the sperm penetrate and fertilize the egg. An ejaculate will contain from two to five milliliters of fluid with 50 to 120 million sperm per milliliter.

Micrograph shows human sperm, which have an oval head about 3 microns across and a very long flagellum. Illustration shows that the head is surrounded by the acrosome. The part of the tail closest to the head, called the neck, is thicker than the rest.

Figure 34.11 Human sperm, visualized using scanning electron microscopy, have a flagellum, neck, and head. (credit b: modification of work by Mariana Ruiz Villareal; scale-bar data from Matt Russell)

The bulk of the semen comes from the accessory glands associated with the male reproductive system. These are the seminal vesicles, the prostate gland, and the bulbourethral gland, all of which are illustrated in Figure 34.10. The seminal vesicles are a pair of glands that lie along the posterior border of the urinary bladder. The glands make a solution that is thick, yellowish, and alkaline. As sperm are only motile in an alkaline environment, a basic pH is important to reverse the acidity of the vaginal environment. The solution also contains mucus, fructose (a sperm mitochondrial nutrient), a coagulating enzyme, ascorbic acid, and local-acting hormones called prostaglandins. The seminal vesicle glands account for 60 percent of the bulk of semen.

The penis, illustrated in Figure 34.10, is an organ that drains urine from the renal bladder and functions as a copulatory organ during intercourse. The penis contains three tubes of erectile tissue running through the length of the organ. These consist of a pair of tubes on the dorsal side, called the corpus cavernosum, and a single tube of tissue on the ventral side, called the corpus spongiosum. This tissue will become engorged with blood, becoming erect and hard, in preparation for intercourse. The organ is inserted into the vagina culminating with an ejaculation. During intercourse, the smooth muscle sphincters at the opening to the renal bladder close and prevent urine from entering the penis. An orgasm is a two-stage process: first, glands and accessory organs connected to the testes contract, then semen (containing sperm) is expelled through the urethra during ejaculation. After intercourse, the blood drains from the erectile tissue and the penis becomes flaccid.

The walnut-shaped prostate gland surrounds the urethra, the connection to the urinary bladder. It has a series of short ducts that directly connect to the urethra. The gland is a mixture of smooth muscle and glandular tissue. The muscle provides much of the force needed for ejaculation to occur. The glandular tissue makes a thin, milky fluid that contains citrate (a nutrient), enzymes, and prostate specific antigen (PSA). PSA is a proteolytic enzyme that helps to liquefy the ejaculate several minutes after release from the male. Prostate gland secretions account for about 30 percent of the bulk of semen.

The bulbourethral gland, or Cowper’s gland, releases its secretion before the release of the bulk of the semen. It neutralizes any acid residue in the urethra left over from urine. This usually accounts for a couple of drops of fluid in the total ejaculate and may contain a few sperm. Withdrawal of the penis from the vagina before ejaculation to prevent pregnancy may not work if sperm are present in the bulbourethral gland secretions. The location and functions of the male reproductive organs are summarized in Table 34.1.

Male Reproductive Anatomy
Organ	Location	Function
Scrotum	External	Carry and support testes
Penis	External	Deliver urine, copulating organ
Testes	Internal	Produce sperm and male hormones
Seminal Vesicles	Internal	Contribute to semen production
Prostate Gland	Internal	Contribute to semen production
Bulbourethral Glands	Internal	Clean urethra at ejaculation

Table 34.1

Female Reproductive Anatomy

Several reproductive structures are exterior to the female’s body. These include the breasts and the vulva, which consists of the mons pubis, clitoris, labia majora, labia minora, and the vestibular glands, all illustrated in Figure 34.12. The location and functions of the female reproductive organs are summarized in Table 34.2. The vulva is an area associated with the vestibule which includes the structures found in the inguinal (groin) area of women. The mons pubis is a round, fatty area that overlies the pubic symphysis. The clitoris is a structure with erectile tissue that contains a large number of sensory nerves and serves as a source of stimulation during intercourse. The labia majora are a pair of elongated folds of tissue that run posterior from the mons pubis and enclose the other components of the vulva. The labia majora derive from the same tissue that produces the scrotum in a male. The labia minora are thin folds of tissue centrally located within the labia majora. These labia protect the openings to the vagina and urethra. The mons pubis and the anterior portion of the labia majora become covered with hair during adolescence; the labia minora is hairless. The greater vestibular glands are found at the sides of the vaginal opening and provide lubrication during intercourse.

Side and front views of female reproductive organs are shown. The vagina is wide at the bottom, and narrows into the cervix. Above the cervix is the uterus, which is shaped like a triangle pointing down. Fallopian tubes extend from the top sides of the uterus. The Fallopian tubes curve back in toward the uterus, and end in fingerlike appendages called fimbriae. The ovaries are located between the fimbriae and the uterus. The urethra is located in front of the vagina, and the rectum is located behind. The

Figure 34.12 The reproductive structures of the human female are shown. (credit a: modification of work by Gray's Anatomy; credit b: modification of work by CDC)

Female Reproductive Anatomy
Organ	Location	Function
Clitoris	External	Sensory organ
Mons pubis	External	Fatty area overlying pubic bone
Labia majora	External	Covers labia minora
Labia minora	External	Covers vestibule
Greater vestibular glands	External	Secrete mucus; lubricate vagina
Breast	External	Produce and deliver milk
Ovaries	Internal	Carry and develop eggs
Oviducts (Fallopian tubes)	Internal	Transport egg to uterus
Uterus	Internal	Support developing embryo
Vagina	Internal	Common tube for intercourse, birth canal, passing menstrual flow

Table 34.2

The breasts consist of mammary glands and fat. The size of the breast is determined by the amount of fat deposited behind the gland. Each gland consists of 15 to 25 lobes that have ducts that empty at the nipple and that supply the nursing child with nutrient- and antibody-rich milk to aid development and protect the child.

Internal female reproductive structures include ovaries, oviducts, the uterus, and the vagina, shown in Figure 34.12. The pair of ovaries is held in place in the abdominal cavity by a system of ligaments. Ovaries consist of a medulla and cortex: the medulla has nerves and blood vessels to supply the cortex with nutrients and remove waste. The outer layers of cells of the cortex are the functional parts of the ovaries. The cortex is made up of follicular cells that surround eggs that develop during fetal development in utero. During the menstrual period, a batch of follicular cells develops and prepares the eggs for release. At ovulation, one follicle ruptures and one egg is released, as illustrated in Figure 34.13a.

Illustration A shows a cross section of a human ovary, which is oval with a stem-like structure at one end that anchors it to the uterus. The central part of the ovary is the medulla, and the outer part is the cortex. Follicles exist in the cortex. Small, immature follicles are located near this stem-like structure. As a follicle matures, it grows and moves toward the edge of the ovary opposite the stem, it ruptures, releasing the egg. The follicle is now called a corpus luteum. The corpus luteum matures

Figure 34.13 Oocytes develop in (a) follicles, located in the ovary. At the beginning of the menstrual cycle, the follicle matures. At ovulation, the follicle ruptures, releasing the egg. The follicle becomes a corpus luteum, which eventually degenerates. The (b) follicle in this light micrograph has an oocyte at its center. (credit a: modification of work by NIH; scale-bar data from Matt Russell)

The oviducts, or fallopian tubes, extend from the uterus in the lower abdominal cavity to the ovaries, but they are not in contact with the ovaries. The lateral ends of the oviducts flare out into a trumpet-like structure and have a fringe of finger-like projections called fimbriae, illustrated in Figure 34.12b. When an egg is released at ovulation, the fimbriae help the non-motile egg enter into the tube and passage to the uterus. The walls of the oviducts are ciliated and are made up mostly of smooth muscle. The cilia beat toward the middle, and the smooth muscle contracts in the same direction, moving the egg toward the uterus. Fertilization usually takes place within the oviducts and the developing embryo is moved toward the uterus for development. It usually takes the egg or embryo a week to travel through the oviduct. Sterilization in women is called a tubal ligation; it is analogous to a vasectomy in males in that the oviducts are severed and sealed.

The uterus is a structure about the size of a woman’s fist. It is lined with an endometrium rich in blood vessels and mucus glands. The uterus supports the developing embryo and fetus during gestation. The thickest portion of the wall of the uterus is made of smooth muscle. Contractions of the smooth muscle in the uterus aid in passing the baby through the vagina during labor. A portion of the lining of the uterus sloughs off during each menstrual period, and then builds up again in preparation for an implantation. Part of the uterus, called the cervix, protrudes into the top of the vagina. The cervix functions as the birth canal.

The vagina is a muscular tube that serves several purposes. It allows menstrual flow to leave the body. It is the receptacle for the penis during intercourse and the vessel for the delivery of offspring. It is lined by stratified squamous epithelial cells to protect the underlying tissue.

Sexual Response during Intercourse

The sexual response in humans is both psychological and physiological. Both sexes experience sexual arousal through psychological and physical stimulation. There are four phases of the sexual response. During phase one, called excitement, vasodilation leads to vasocongestion in erectile tissues in both men and women. The nipples, clitoris, labia, and penis engorge with blood and become enlarged. Vaginal secretions are released to lubricate the vagina to facilitate intercourse. During the second phase, called the plateau, stimulation continues, the outer third of the vaginal wall enlarges with blood, and breathing and heart rate increase.

During phase three, or orgasm, rhythmic, involuntary contractions of muscles occur in both sexes. In the male, the reproductive accessory glands and tubules constrict placing semen in the urethra, then the urethra contracts expelling the semen through the penis. In women, the uterus and vaginal muscles contract in waves that may last slightly less than a second each. During phase four, or resolution, the processes described in the first three phases reverse themselves and return to their normal state. Men experience a refractory period in which they cannot maintain an erection or ejaculate for a period of time ranging from minutes to hours.

Gametogenesis (Spermatogenesis and Oogenesis)

Gametogenesis, the production of sperm and eggs, takes place through the process of meiosis. During meiosis, two cell divisions separate the paired chromosomes in the nucleus and then separate the chromatids that were made during an earlier stage of the cell’s life cycle. Meiosis produces haploid cells with half of each pair of chromosomes normally found in diploid cells. The production of sperm is called spermatogenesis and the production of eggs is called oogenesis.

Spermatogenesis

Figure 34.14 During spermatogenesis, four sperm result from each primary spermatocyte.

Spermatogenesis, illustrated in Figure 34.14, occurs in the wall of the seminiferous tubules (Figure 34.10), with stem cells at the periphery of the tube and the spermatozoa at the lumen of the tube. Immediately under the capsule of the tubule are diploid, undifferentiated cells. These stem cells, called spermatogonia (singular: spermatagonium), go through mitosis with one offspring going on to differentiate into a sperm cell and the other giving rise to the next generation of sperm.

Meiosis starts with a cell called a primary spermatocyte. At the end of the first meiotic division, a haploid cell is produced called a secondary spermatocyte. This cell is haploid and must go through another meiotic cell division. The cell produced at the end of meiosis is called a spermatid and when it reaches the lumen of the tubule and grows a flagellum, it is called a sperm cell. Four sperm result from each primary spermatocyte that goes through meiosis.

Stem cells are deposited during gestation and are present at birth through the beginning of adolescence, but in an inactive state. During adolescence, gonadotropic hormones from the anterior pituitary cause the activation of these cells and the production of viable sperm. This continues into old age.

Link to Learning

Visit this site to see the process of spermatogenesis.

During spermatogenesis, which cells undergo meiosis to ultimately produce haploid gametes?

spermatogonia
primary spermatocytes
secondary spermatocytes
spermatids

Oogenesis

Oogenesis, illustrated in Figure 34.15, occurs in the outermost layers of the ovaries. As with sperm production, oogenesis starts with a germ cell, called an oogonium (plural: oogonia), but this cell undergoes mitosis to increase in number, eventually resulting in up to about one to two million cells in the embryo.

Figure 34.15 The process of oogenesis occurs in the ovary’s outermost layer.

The cell starting meiosis is called a primary oocyte, as shown in Figure 34.15. This cell will start the first meiotic division and be arrested in its progress in the first prophase stage. At the time of birth, all future eggs are in the prophase stage. At adolescence, anterior pituitary hormones cause the development of many follicles in an ovary. This results in the primary oocyte finishing the first meiotic division. The cell divides unequally, with most of the cellular material and organelles going to one cell, called a secondary oocyte, and only one set of chromosomes and a small amount of cytoplasm going to the other cell. This second cell is called a polar body and usually dies. A secondary meiotic arrest occurs, this time at the metaphase II stage. At ovulation, this secondary oocyte will be released and travel toward the uterus through the oviduct. If the secondary oocyte is fertilized, the cell continues through the meiosis II, producing a second polar body and a fertilized egg containing all 46 chromosomes of a human being, half of them coming from the sperm.

Egg production begins before birth, is arrested during meiosis until puberty, and then individual cells continue through at each menstrual cycle. One egg is produced from each meiotic process, with the extra chromosomes and chromatids going into polar bodies that degenerate and are reabsorbed by the body.

Science Practice Connection for AP® Courses

Think About It

Compare spermatogenesis and oogenesis as to the timing of the processes and the number and type of cells finally produced.

Disclaimer

This section may include links to websites that contain links to articles on unrelated topics. See the preface for more information.

34.3 Human Reproductive Anatomy and Gametogenesis

Learning Objectives