Chapter 18. Animal Reproduction and Development

Fission, also called binary fission, occurs in some invertebrate, multi-celled organisms. It is in some ways analogous to the process of binary fission of single-celled prokaryotic organisms. The term fission is applied to instances in which an organism appears to split itself into two parts and, if necessary, regenerate the missing parts of each new organism. For example, species of turbellarian flatworms commonly called the planarians, such as Dugesia dorotocephala, are able to separate their bodies into head and tail regions and then regenerate the missing half in each of the two new organisms. Sea anemones (Cnidaria), such as species of the genus Anthopleura (Figure 18.2), will divide along the oral-aboral axis, and sea cucumbers (Echinodermata) of the genus Holothuria, will divide into two halves across the oral-aboral axis and regenerate the other half in each of the resulting individuals.

Figure 18.2. 

The Anthopleura artemisia sea anemone can reproduce through fission.

Budding is a form of asexual reproduction that results from the outgrowth of a part of the body leading to a separation of the “bud” from the original organism and the formation of two individuals, one smaller than the other. Budding occurs commonly in some invertebrate animals such as hydras and corals. In hydras, a bud forms that develops into an adult and breaks away from the main body (Figure 18.3).

Figure 18.3. 

(a) Hydra reproduce asexually through budding: a bud forms on the tubular body of an adult hydra, develops a mouth and tentacles, and then detaches from its parent. The new hydra is fully developed and will find its own location for attachment. (b) Some coral, such as the Lophelia pertusa shown here, can reproduce through budding. (credit b: modification of work by Ed Bowlby, NOAA/Olympic Coast NMS; NOAA/OAR/Office of Ocean Exploration)

Concept in Action

View this video to see a hydra budding.

Fragmentation is the breaking of an individual into parts followed by regeneration. If the animal is capable of fragmentation, and the parts are big enough, a separate individual will regrow from each part. Fragmentation may occur through accidental damage, damage from predators, or as a natural form of reproduction. Reproduction through fragmentation is observed in sponges, some cnidarians, turbellarians, echinoderms, and annelids. In some sea stars, a new individual can be regenerated from a broken arm and a piece of the central disc. This sea star (Figure 18.4) is in the process of growing a complete sea star from an arm that has been cut off. Fisheries workers have been known to try to kill the sea stars eating their clam or oyster beds by cutting them in half and throwing them back into the ocean. Unfortunately for the workers, the two parts can each regenerate a new half, resulting in twice as many sea stars to prey upon the oysters and clams.

Figure 18.4. 

(a) Linckia multifora is a species of sea star that can reproduce asexually via fragmentation. In this process, (b) an arm that has been shed grows into a new sea star. (credit a: modifiction of work by Dwayne Meadows, NOAA/NMFS/OPR)

Parthenogenesis is a form of asexual reproduction in which an egg develops into an individual without being fertilized. The resulting offspring can be either haploid or diploid, depending on the process in the species. Parthenogenesis occurs in invertebrates such as water fleas, rotifers, aphids, stick insects, and ants, wasps, and bees. Ants, bees, and wasps use parthenogenesis to produce haploid males (drones). The diploid females (workers and queens) are the result of a fertilized egg.

Some vertebrate animals—such as certain reptiles, amphibians, and fish—also reproduce through parthenogenesis. Parthenogenesis has been observed in species in which the sexes were separated in terrestrial or marine zoos. Two female Komodo dragons, a hammerhead shark, and a blacktop shark have produced parthenogenic young when the females have been isolated from males. It is possible that the asexual reproduction observed occurred in response to unusual circumstances and would normally not occur.

Hermaphroditism occurs in animals in which one individual has both male and female reproductive systems. Invertebrates such as earthworms, slugs, tapeworms, and snails (Figure 18.5) are often hermaphroditic. Hermaphrodites may self-fertilize, but typically they will mate with another of their species, fertilizing each other and both producing offspring. Self-fertilization is more common in animals that have limited mobility or are not motile, such as barnacles and clams. Many species have specific mechanisms in place to prevent self-fertilization, because it is an extreme form of inbreeding and usually produces less fit offspring.

Figure 18.5. 

Many (a) snails are hermaphrodites. When two individuals (b) mate, they can produce up to 100 eggs each. (credit a: modification of work by Assaf Shtilman; credit b: modification of work by “Schristia”/Flickr)

Internal fertilization occurs most often in terrestrial animals, although some aquatic animals also use this method. Internal fertilization may occur by the male directly depositing sperm in the female during mating. It may also occur by the male depositing sperm in the environment, usually in a protective structure, which a female picks up to deposit the sperm in her reproductive tract. There are three ways that offspring are produced following internal fertilization. In oviparity, fertilized eggs are laid outside the female’s body and develop there, receiving nourishment from the yolk that is a part of the egg (Figure 18.7a). This occurs in some bony fish, some reptiles, a few cartilaginous fish, some amphibians, a few mammals, and all birds. Most non-avian reptiles and insects produce leathery eggs, while birds and some turtles produce eggs with high concentrations of calcium carbonate in the shell, making them hard. Chicken eggs are an example of a hard shell. The eggs of the egg-laying mammals such as the platypus and echidna are leathery.

In ovoviparity, fertilized eggs are retained in the female, and the embryo obtains its nourishment from the egg’s yolk. The eggs are retained in the female’s body until they hatch inside of her, or she lays the eggs right before they hatch. This process helps protect the eggs until hatching. This occurs in some bony fish (like the platyfish Xiphophorus maculatus, Figure 18.7b), some sharks, lizards, some snakes (garter snake Thamnophis sirtalis), some vipers, and some invertebrate animals (Madagascar hissing cockroach Gromphadorhina portentosa).

In viviparity the young are born alive. They obtain their nourishment from the female and are born in varying states of maturity. This occurs in most mammals (Figure 18.7c), some cartilaginous fish, and a few reptiles.

Figure 18.7. 

In (a) oviparity, young develop in eggs outside the female body, as with these Harmonia axydridis beetles hatching. Some aquatic animals, like this (b) pregnant Xiphophorus maculatus are ovoviparous, with the egg developing inside the female and nutrition supplied primarily from the yolk. In mammals, nutrition is supported by the placenta, as was the case with this (c) newborn squirrel. (credit b: modification of work by Gourami Watcher; credit c: modification of work by “audreyjm529″/Flickr)

Gastrulation leads to the formation of the three germ layers that give rise during further development to the different organs in the animal body. This process is called organogenesis.

Organs develop from the germ layers through the process of differentiation. During differentiation, the embryonic stem cells express specific sets of genes that will determine their ultimate cell type. For example, some cells in the ectoderm will express the genes specific to skin cells. As a result, these cells will take on the shape and characteristics of epidermal cells. The process of differentiation is regulated by location-specific chemical signals from the cell’s embryonic environment that sets in play a cascade of events that regulates gene expression.

Sperm are immobile at body temperature; therefore, the testes are external to the body so that a correct temperature is maintained for motility. In land mammals, including humans, the pair of testes must be suspended outside the body so the environment of the sperm is about 2 °C lower than body temperature to produce viable sperm. If the testes do not descend through the abdominal cavity during fetal development, the individual has reduced fertility.

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

The penis drains urine from the urinary bladder and is a copulatory organ during intercourse (Figure 18.12; Table 18.1). The penis contains three tubes of erectile tissue that become engorged with blood, making the penis erect, in preparation for intercourse. The organ is inserted into the vagina culminating with an ejaculation. During orgasm, the accessory organs and glands connected to the testes contract and empty the semen (containing sperm) into the urethra and the fluid is expelled from the body by muscular contractions causing ejaculation. After intercourse, the blood drains from the erectile tissue and the penis becomes flaccid.

Semen is a mixture of sperm (about five 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 for motility, a neck that contains the cell’s energy-producing mitochondria, and a head that contains the genetic material (Figure 18.11). An acrosome (acrosomal vesicle) is found at the top of the head of the sperm. This structure contains enzymes that can digest the protective coverings that surround the egg and allow the sperm to fuse with the egg. An ejaculate will contain from two to five milliliters of fluid and from 50–120 million sperm per milliliter.

Figure 18.11. 

As seen in this scanning electron micrograph, human sperm has a flagellum, neck, and head. (credit: scale-bar data from Matt Russell)

Sperm form in the walls of seminiferous tubules that are coiled inside the testes (Figure 18.12; Table 18.1). 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 next to the lumen. The sperm cells are associated with Sertoli cells that nourish and promote the development of the sperm. Other cells present between the walls of the tubules are the interstitial cells of Leydig, which produce testosterone once the male reaches adolescence.

When the sperm have developed flagella they leave the seminiferous tubules and enter the epididymis (Figure 18.12; Table 18.1). This structure lies along the top and posterior of the testes and is the site of sperm maturation. The sperm leave the epididymis and enter the vas 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 (but not the secretions of the accessory glands) from being passed out of the body during ejaculation and preventing fertilization.

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 (Figure 18.12; Table 18.1). The secretions from the accessory glands provide important compounds for the sperm including nutrients, electrolytes, and pH buffering. There are also coagulation factors that affect sperm delivery and motility.

Art Connection

 

Figure 18.12. 

The reproductive structures of the human male are shown.

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

1. The vas deferens carries sperm from the testes to the seminal vesicles.
2. The ejaculatory duct joins the urethra.
3. Both the prostate and the bulbourethral glands produce components of the semen.
4. The prostate gland is located in the testes.

A number of female reproductive structures are exterior to the body. These include the breasts and the vulva, which consists of the mons pubis, clitoris, labia majora, labia minora, and the vestibular glands (Figure 18.13; Table 18.2).

Figure 18.13. 

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)

The breasts consist of mammary glands and fat. 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 (Figure 18.13; Table 18.2). The pair of ovaries is held in place in the abdominal cavity by a system of ligaments. The outermost layer of the ovary is made up of follicles, each consisting of one or more follicular cells that surround, nourish, and protect a single egg. During the menstrual period, a batch of follicular cells develops and prepares their eggs for release. At ovulation, one follicle ruptures and one egg is released. Following ovulation, the follicular tissue that surrounded the ovulated egg stays within the ovary and grows to form a solid mass called the corpus luteum. The corpus luteum secretes additional estrogen and the hormone progesterone that helps maintain the uterine lining during pregnancy. The ovaries also produce hormones, such as estrogen.

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 fimbrae. When an egg is released at ovulation, the fimbrae help the nonmotile egg enter into the tube. The walls of the oviducts have a ciliated epithelium over smooth muscle. The cilia beat, and the smooth muscle contracts, moving the egg toward the uterus. Fertilization usually takes place within the oviduct and the developing embryo is moved toward the uterus. 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, preventing sperm from reaching the egg.

The uterus is a structure about the size of a woman’s fist. The uterus has a thick muscular wall and is lined with an endometrium rich in blood vessels and mucus glands that develop and thicken during the female cycle. Thickening of the endometrium prepares the uterus to receive the fertilized egg or zygote, which will then implant itself in the endometrium. The uterus supports the developing embryo and fetus during gestation. Contractions of the smooth muscle in the uterus aid in forcing the baby through the vagina during labor. If fertilization does not occur, a portion of the lining of the uterus sloughs off during each menstrual period. The endometrium builds up again in preparation for implantation. Part of the uterus, called the cervix, protrudes into the top of the vagina.

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 pathway for the delivery of offspring.

Spermatogenesis occurs in the wall of the seminiferous tubules, with the most primitive cells at the periphery of the tube and the most mature sperm at the lumen of the tube (Figure 18.14). Immediately under the capsule of the tubule are diploid, undifferentiated cells. These stem cells, each called a spermatogonium (pl. spermatogonia), go through mitosis to produce one cell that remains as a stem cell and a second cell called a primary spermatocyte that will undergo meiosis to produce sperm.

The diploid primary spermatocyte goes through meiosis I to produce two haploid cells called secondary spermatocytes. Each secondary spermatocyte divides after meiosis II to produce two cells called spermatids. The spermatids eventually reach the lumen of the tubule and grow a flagellum, becoming sperm cells. Four sperm result from each primary spermatocyte that goes through meiosis.

Figure 18.14. 

During spermatogenesis, four sperm result from each primary spermatocyte. The process also maps onto the physical structure of the wall of the seminiferous tubule, with the spermatogonia on the outer side of the tubule, and the sperm with their developing tails extended into the lumen of the tubule.

Concept in Action

Visit this site to see the process of spermatogenesis.

Oogenesis occurs in the outermost layers of the ovaries. As with sperm production, oogenesis starts with a germ cell. In oogenesis, this germ cell is called an oogonium and forms during the embryological development of the individual. The oogonium undergoes mitosis to produce about one to two million oocytes by the time of birth.

Figure 18.15. 

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

The primary oocytes begin meiosis before birth (Figure 18.15). However, the meiotic division is arrested in its progress in the first prophase stage. At the time of birth, all future eggs are in prophase I. This situation is in contrast with the male reproductive system in which sperm are produced continuously throughout the life of the individual. Starting at adolescence, anterior pituitary hormones cause the development of a few follicles in an ovary each month. This results in a primary oocyte finishing the first meiotic division. The cell divides unequally, with most of the cytoplasm 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. Cell division is again arrested, this time at metaphase II. At ovulation, this secondary oocyte is released and travels toward the uterus through the oviduct. If the secondary oocyte is fertilized, the cell continues through meiosis II, producing a second polar body and haploid egg, which fuses with the haploid sperm to form a fertilized egg (zygote) containing all 46 chromosomes.

At the onset of puberty, the hypothalamus causes the release of FSH and LH into the male system for the first time. FSH enters the testes and stimulates the Sertoli cells located in the walls of the seminiferous tubules to begin promoting spermatogenesis (Figure 18.16). LH also enters the testes and stimulates the interstitial cells of Leydig, located in between the walls of the seminiferous tubules, to make and release testosterone into the testes and the blood.

Testosterone stimulates spermatogenesis. This hormone is also responsible for the secondary sexual characteristics that develop in the male during adolescence. The secondary sex characteristics in males include a deepening of the voice, the growth of facial, axillary, and pubic hair, an increase in muscle bulk, and the beginnings of the sex drive.

Figure 18.16. 

Hormones control sperm production in a negative feedback system.

A negative feedback system occurs in the male with rising levels of testosterone acting on the hypothalamus and anterior pituitary to inhibit the release of GnRH, FSH, and LH. In addition, the Sertoli cells produce the hormone inhibin, which is released into the blood when the sperm count is too high. This inhibits the release of GnRH and FSH, which will cause spermatogenesis to slow down. If the sperm count reaches a low of 20 million/mL, the Sertoli cells cease the release of inhibin, and the sperm count increases.

The ovarian and menstrual cycles are regulated by hormones of the hypothalamus, pituitary, and ovaries (Figure 18.17). The ebb and flow of the hormones causes the ovarian and menstrual cycles to advance. The ovarian and menstrual cycles occur concurrently. The first half of the ovarian cycle is the follicular phase. Slowly rising levels of FSH cause the growth of follicles on the surface of the ovary. This process prepares the egg for ovulation. As the follicles grow, they begin releasing estrogen. The first few days of this cycle coincide with menstruation or the sloughing off of the functional layer of the endometrium in the uterus. After about five days, estrogen levels rise and the menstrual cycle enters the proliferative phase. The endometrium begins to regrow, replacing the blood vessels and glands that deteriorated during the end of the last cycle.

Art Connection

 

Figure 18.17. 

The ovarian and menstrual cycles of female reproduction are regulated by hormones produced by the hypothalamus, pituitary, and ovaries.

Which of the following statements about hormone regulation of the female reproductive cycle is false?

1. LH and FSH are produced in the pituitary, and estrogen and progesterone are produced in the ovaries.
2. Estradiol and progesterone secreted from the corpus luteum cause the endometrium to thicken.
3. Both progesterone and estrogen are produced by the follicles.
4. Secretion of GnRH by the hypothalamus is inhibited by low levels of estrogen but stimulated by high levels of estrogen.

Just prior to the middle of the cycle (approximately day 14), the high level of estrogen causes FSH and especially LH to rise rapidly then fall. The spike in LH causes the most mature follicle to rupture and release its egg. This is ovulation. The follicles that did not rupture degenerate and their eggs are lost. The level of estrogen decreases when the extra follicles degenerate.

Following ovulation, the ovarian cycle enters its luteal phase and the menstrual cycle enters its secretory phase, both of which run from about day 15 to 28. The luteal and secretory phases refer to changes in the ruptured follicle. The cells in the follicle undergo physical changes and produce a structure called a corpus luteum. The corpus luteum produces estrogen and progesterone. The progesterone facilitates the regrowth of the uterine lining and inhibits the release of further FSH and LH. The uterus is being prepared to accept a fertilized egg, should it occur during this cycle. The inhibition of FSH and LH prevents any further eggs and follicles from developing, while the progesterone is elevated. The level of estrogen produced by the corpus luteum increases to a steady level for the next few days.

If no fertilized egg is implanted into the uterus, the corpus luteum degenerates and the levels of estrogen and progesterone decrease. The endometrium begins to degenerate as the progesterone levels drop, initiating the next menstrual cycle. The decrease in progesterone also allows the hypothalamus to send GnRH to the anterior pituitary, releasing FSH and LH and starting the cycles again.