Sexual reproduction

Sexual reproduction was likely an early evolutionary innovation after the appearance of eukaryotic cells. It appears to have been very successful because most eukaryotes are able to reproduce sexually and, in many animals, it is the only mode of reproduction. And yet, scientists also recognize some real disadvantages to sexual reproduction. On the surface, creating offspring that are genetic clones of the parent appears to be a better system. If the parent organism is successfully occupying a habitat, offspring with the same traits should be similarly successful. There is also the obvious benefit to an organism that can produce offspring whenever circumstances are favorable by asexual budding, fragmentation, or by producing eggs asexually. These methods of reproduction do not require a partner with which to reproduce. Indeed, some organisms that lead a solitary lifestyle have retained the ability to reproduce asexually. In addition, in asexual populations, every individual is capable of reproduction. In sexual populations, the males are not producing the offspring themselves, so hypothetically an asexual population could grow twice as fast.

However, multicellular organisms that exclusively depend on asexual reproduction are exceedingly rare. Why are meiosis and sexual reproductive strategies so common? These are important (and as yet unanswered) questions in biology, even though they have been the focus of much research beginning in the latter half of the 20th century. There are several possible explanations, one of which is that the variation that sexual reproduction creates among offspring is very important to the survival and reproduction of the population. Thus, on average, a sexually reproducing population will leave more descendants than an otherwise similar asexually reproducing population. The only source of variation in asexual organisms is mutation. Mutations that take place during the formation of germ cell lines are also a source of variation in sexually reproducing organisms. However, in contrast to mutation during asexual reproduction, the mutations during sexual reproduction can be continually reshuffled from one generation to the next when different parents combine their unique genomes and the genes are mixed into different combinations by crossovers during prophase I and random assortment at metaphase I.

Sexual reproduction requires the union of two specialized cells, called gametes, each of which contains one set of chromosomes. When gametes unite, they form a zygote, or fertilized egg that contains two sets of chromosomes. (Note: Cells that contain one set of chromosomes are called haploid; cells containing two sets of chromosomes are called diploid.) If the reproductive cycle is to continue for any sexually reproducing species, then the diploid cell must somehow reduce its number of chromosome sets to produce haploid gametes; otherwise, the number of chromosome sets will double with every future round of fertilization. Therefore, sexual reproduction requires a nuclear division that reduces the number of chromosome sets by half.

Most animals and plants and many unicellular organisms are diploid and therefore have two sets of chromosomes. In each somatic cell of the organism (all cells of a multicellular organism except the gametes or reproductive cells), the nucleus contains two copies of each chromosome, called homologous chromosomes. Homologous chromosomes are matched pairs containing the same genes in identical locations along their lengths. Diploid organisms inherit one copy of each homologous chromosome from each genetic contributor.

Meiosis is the nuclear division that forms haploid cells from diploid cells, and it employs many of the same cellular mechanisms as mitosis. However, as you have learned, mitosis produces daughter cells whose nuclei are genetically identical to the original parent nucleus. In mitosis, both the parent and the daughter nuclei are at the same “ploidy level”—diploid in the case of most multicellular animals. Plants use mitosis to grow as sporophytes, and to grow and produce eggs and sperm as gametophytes; so they use mitosis for both haploid and diploid cells (as well as for all other ploidies). In meiosis, the starting nucleus is always diploid and the daughter nuclei that result are haploid. To achieve this reduction in chromosome number, meiosis consists of one round of chromosome replication followed by two rounds of nuclear division. Because many events that occur during each of the division stages are analogous to the events of mitosis, the same stage names are assigned. However, because there are two rounds of division, the major process and the stages are designated with a “I” or a “II.” Thus, meiosis I is the first round of meiotic division and consists of prophase I, prometaphase I, and so on. Likewise, Meiosis II (during which the second round of meiotic division takes place) includes prophase II, prometaphase II, and so on. We discussed meiosis (Figure 26.1) in great detail in BIOL 1403, so we will not spend as much time on it in BIOL 1404.

Life Cycles of Sexually Reproducing Organisms

Fertilization and meiosis alternate in sexual life cycles. What happens between these two events depends on the organism’s “reproductive strategy.” The process of meiosis reduces the chromosome number by half. Fertilization, the joining of two haploid gametes, restores the diploid condition. Some organisms have a multicellular diploid stage that is most obvious and only produce haploid reproductive cells. Animals, including humans, have this type of life cycle. Other organisms, such as fungi, have a multicellular haploid stage that is most obvious. Plants and some algae have alternation of generations, in which they have multicellular diploid and haploid life stages that are apparent to different degrees depending on the group.

Nearly all animals employ a diploid-dominant life-cycle strategy in which the only haploid cells produced by the organism are the gametes. Early in the development of the embryo, specialized diploid cells, called germ cells, are produced within the gonads (such as the testes and ovaries). Germ cells are capable of mitosis to perpetuate the germ cell line and meiosis to produce haploid gametes. Once the haploid gametes are formed, they lose the ability to divide again. There is no multicellular haploid life stage. Fertilization occurs with the fusion of two gametes, usually from different individuals, restoring the diploid state (Figure 26.2).

Most fungi and algae employ a life-cycle type in which the “body” of the organism—the ecologically important part of the life cycle—is haploid. The haploid cells that make up the tissues of the dominant multicellular stage are formed by mitosis. During sexual reproduction, specialized haploid cells from two individuals—designated the (+) and (−) mating types—join to form a diploid zygote. The zygote immediately undergoes meiosis to form four haploid cells called spores. Although these spores are haploid like the “parents,” they contain a new genetic combination from two parents. The spores can remain dormant for various time periods. Eventually, when conditions are favorable, the spores form multicellular haploid structures through many rounds of mitosis (Figure 26.3).

The third life-cycle type, employed by some algae and all plants, is a blend of the haploid-dominant and diploid-dominant extremes. Species with alternation of generations have both haploid and diploid multicellular organisms as part of their life cycle. The haploid multicellular plants are called gametophytes, because they produce gametes from specialized cells. Meiosis is not directly involved in the production of gametes in this case, because the organism that produces the gametes is already haploid. Fertilization between the gametes forms a diploid zygote. The zygote will undergo many rounds of mitosis and give rise to a diploid multicellular plant called a sporophyte. Specialized cells of the sporophyte will undergo meiosis and produce haploid spores. The spores will subsequently develop into the gametophytes (Figure 26.4).

Although all plants utilize some version of the alternation of generations, the relative size of the sporophyte and the gametophyte and the relationship between them vary greatly. In plants such as moss, the gametophyte organism is the free-living plant and the sporophyte is physically dependent on the gametophyte. In other plants, such as ferns, both the gametophyte and sporophyte plants are free-living; however, the sporophyte is much larger. In seed plants, such as magnolia trees and daisies, the gametophyte is composed of only a few cells and, in the case of the female gametophyte, is completely retained within the sporophyte.

Sexual reproduction takes many forms in multicellular organisms. The fact that nearly every multicellular organism on Earth employs sexual reproduction is strong evidence for the benefits of producing offspring with unique gene combinations, though there are other possible benefits as well.

Reading Question #1

Which of the following are benefits of sexual reproduction? Select all that apply.

A. Variation can lead to better adaptations and survival

B. Genetic diversity in the offspring

C. Less susceptible to being wiped out due to disease

D. Doesn’t require a partner

 

Gametogenesis

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

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 in more detail.

Oogenesis

Oogenesis, illustrated in Figure 26.6, 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.

The cell starting meiosis is called a primary oocyte, as shown in Figure 26.6. 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 a number of 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.

Reading Question #2

Which of the following statements accurately describes a key difference between oogenesis and spermatogenesis?

A. Spermatogenesis produces polar bodies while oogenesis does not produce polar bodies.

B. In oogenesis, each meiosis round produces only one egg while in spermatogenesis, each meiosis round produces four sperm.

C. In oogenesis, each meiosis round produces four egg while in spermatogenesis, each meiosis round produces one sperm.

D. Spermatogenesis yields haploid cells while oogenesis yields diploid cells.

 

 

Hormonal Regulation of the Reproductive System

Regulation of the reproductive system is a process that requires the action of hormones from the pituitary gland, theadrenal cortex, and the gonads. During puberty in both males and females, the hypothalamus produces gonadotropin-releasing hormone (GnRH), which stimulates the production and release of follicle-stimulating hormone (FSH) and luteinizing hormone (LH) from the anterior pituitary gland. These hormones regulate the gonads (testes in males and ovaries in females) and therefore are called gonadotropins. In both males and females, FSH stimulates gamete production and LH stimulates production of hormones by the gonads. An increase in gonad hormone levels inhibits GnRH production through a negative feedback loop.

Regulation of the Testicular Reproductive System

In the testes, FSH stimulates the maturation of sperm cells. FSH production is inhibited by the hormone inhibin, which is released by the testes. LH stimulates production of the sex hormones (androgens) by the interstitial cells of the testes and therefore is also called interstitial cell-stimulating hormone.

The most widely known androgen in males is testosterone. Testosterone promotes the production of sperm and a suite of secondary sex characteristics, such as the growth and development of the testes and penis, increased skeletal and muscular growth, enlargement of the larynx, increased growth and redistribution of body hair, and increased sexual drive. The adrenal cortex also produces small amounts of testosterone precursor, although the role of this additional hormone production is not fully understood. Testosterone secretion is regulated by both the hypothalamus and the anterior pituitary gland. The hypothalamus sends releasing hormones that stimulate the release of gonadotropins from the anterior pituitary gland.

Everyday Connection

The Dangers of Synthetic Hormones

Some athletes attempt to boost their performance by using artificial hormones that enhance muscle performance. Anabolic steroids, a form of testosterone, are one of the most widely known performance-enhancing drugs. Steroids are used in sports to help build muscle mass. Other hormones that are used to enhance athletic performance include erythropoietin, which triggers the production of red blood cells, and human growth hormone, which can help in building muscle mass. Most performance enhancing drugs are illegal for nonmedical purposes. They are also banned by national and international governing bodies including the International Olympic Committee, the U.S. Olympic Committee, the National Collegiate Athletic Association, the Major League Baseball, and the National Football League.

The side effects of synthetic hormones are often significant and nonreversible, and in some cases, fatal. Androgens can produce several complications such as liver dysfunctions and liver tumors, prostate gland enlargement, difficulty urinating, premature closure of epiphyseal cartilages, testicular atrophy, infertility, and immune system depression. The physiological strain caused by these substances is often greater than what the body can handle, leading to unpredictable and dangerous effects and linking their use to heart attacks, strokes, and impaired cardiac function.

Regulation of the Ovarian Reproductive System

In the ovaries, FSH stimulates development of egg cells, called ova, which develop in structures called follicles. Follicle cells produce the hormone inhibin, which inhibits FSH production. LH also plays a role in the development of ova, induction of ovulation, and stimulation of estradiol and progesterone production by the ovaries (as well as testosterone production by the testes), as illustrated in Figure 26.8. Estradiol and progesterone are steroid hormones that serve several functions in the human body. Estradiol causes the egg to mature and release during the menstrual cycle, and thickens the uterine lining prior to egg implantation. Estradiol also helps with bone health, nitric oxide production, and brain function. (Low levels of estradiol have been connected to osteoporosis, mood swings, weight gain, and interrupted menstrual cycle. High levels of estradiol correlate with an increased risk of uterine and breast cancer, and cardiovascular disease.) During puberty, estradiol produces a suite of characteristics such as the increased development of breast tissue, redistribution of fat towards hips, legs, and breast, and the maturation of the uterus and vagina. Both estradiol and progesterone regulate the menstrual cycle.

In addition to producing FSH and LH, the anterior portion of the pituitary gland also produces the hormone prolactin (PRL). Prolactin stimulates the production of milk by the mammary glands following childbirth. Prolactin release inhibits the release of GnRH from the hypothalamus, resulting in a loss of FSH and LH release from the anterior pituitary. Prolactin levels are regulated by the hypothalamic hormones prolactin-releasing hormone (PRH) and prolactin-inhibiting hormone (PIH), which is now known to be dopamine. PRH stimulates the release of prolactin and PIH inhibits it.

The posterior pituitary releases the hormone oxytocin, which stimulates uterine contractions during childbirth. (Note: the posterior pituitary secretes only two hormones: antidiuretic hormone (ADH and oxytocin). The uterine smooth muscles are not very sensitive to oxytocin until late in pregnancy when the number of oxytocin receptors in the uterus peaks. Stretching of tissues in the uterus and cervix stimulates oxytocin release during childbirth. Contractions increase in intensity as blood levels of oxytocin rise via a positive feedback mechanism until the birth is complete. Oxytocin also stimulates the contraction of myoepithelial cells around the milk-producing mammary glands. As these cells contract, milk is forced from the secretory alveoli into milk ducts and is ejected from the breasts in milk ejection (“let-down”) reflex. Oxytocin release is stimulated by the suckling of an infant, which triggers the synthesis of oxytocin in the hypothalamus and its release into circulation at the posterior pituitary.

Nearly all eukaryotes undergo sexual reproduction. The variation introduced into the reproductive cells by meiosis provides an important advantage that has made sexual reproduction evolutionarily successful. Meiosis and fertilization alternate in sexual life cycles. The process of meiosis produces unique reproductive cells called gametes, which have half the number of chromosomes as the parent cell. When two haploid gametes fuse, this restores the diploid condition in the new zygote. Thus, most sexually reproducing organisms alternate between haploid and diploid stages. However, the ways in which reproductive cells are produced and the timing between meiosis and fertilization vary greatly.

Menstruation

What Is the Menstrual Cycle?

The menstrual cycle refers to natural changes that occur in the female reproductive system each month during the reproductive years. The cycle is necessary for the production of eggs and the preparation of the uterus for pregnancy. It involves changes in both the ovaries and the uterus and is controlled by pituitary and ovarian hormones. Day 1 of the cycle is the first day of the menstrual period, when bleeding from the uterus begins as the built-up endometrium lining the uterus is shed. The endometrium builds up again during the remainder of the cycle, only to be shed again during the beginning of the next cycle if pregnancy does not occur. In the ovaries, the menstrual cycle includes the development of a follicle, ovulation of a secondary oocyte, and the degeneration of the 

Menarche and Menopause

The female reproductive years are delineated by the start and stop of the menstrual cycle. The first menstrual period usually occurs around 12 or 13 years of age, an event that is known as menarche. There is considerable variation among individuals in the age of menarche. It may occasionally occur as early as eight years of age or as late as 16 years of age and still be considered normal. The average age is generally later in the developing world, and earlier in the developed world. This variation is thought to be largely attributable to nutritional differences.

The cessation of menstrual cycles at the end of a woman’s reproductive years is termed menopause. The average age of menopause is 52 years, but it may occur normally at any age between about 45 and 55 years of age. The age of menopause varies due to a variety of biological and environmental factors. It may occur earlier as a result of certain illnesses or medical treatments.

Variation in the Menstrual Cycle

The length of the menstrual cycle— as well as its phases — may vary considerably, not only among different individuals but also from month to month for a given person. The average length of time between the first day of one menstrual period and the first day of the next menstrual period is 28 days, but it may range from 21 days to 45 days. Cycles are considered regular when a woman’s longest and shortest cycles differ by less than eight days. The menstrual period itself is usually about five days long, but it may vary in length from about two days to seven days.

Ovarian Cycle

The events of the menstrual cycle that take place in the ovaries make up the ovarian cycle. It consists of changes that occur in the follicles of one of the ovaries. The ovarian cycle is divided into the following three phases: the follicular phase, ovulation, and luteal phase. These phases are illustrated in Figure 26.9. 

Follicular Phase

The follicular phase is the first phase of the ovarian cycle. It generally lasts about 12 to 14 days for a 28-day menstrual cycle. During this phase, several ovarian follicles are stimulated to begin maturing, but usually only one — called the Graafian follicle— matures completely so it is ready to release an egg. The other maturing follicles stop growing and disintegrate. Follicular development occurs because of a rise in the blood level of follicle-stimulating hormone (FSH), which is secreted by the pituitary gland. The maturing follicle releases estrogen, the level of which rises throughout the follicular phase. You can see these and other changes in hormone levels that occur during the menstrual cycle in the chart in Figure 26.10.

Ovulation

Uterine Cycle

The events of the menstrual cycle that take place in the uterus make up the uterine cycle. This cycle consists of changes that occur mainly in the endometrium, which is the layer of tissue that lines the uterus. The uterine cycle is divided into the following three phases: menstruation, proliferative phase, and secretory phase. These phases are illustrated in Figure 26.11.

Sexual reproduction in plants

Sexual reproduction takes place with slight variations in different groups of plants. Plants have two distinct stages in their lifecycle: the gametophyte stage and the sporophyte stage. The haploid gametophyte produces the male and female gametes by mitosis in distinct multicellular structures. Fusion of the male and females gametes forms the diploid zygote, which develops into the sporophyte. After reaching maturity, the diploid sporophyte produces spores by meiosis, which in turn divide by mitosis to produce the haploid gametophyte. The new gametophyte produces gametes, and the cycle continues. This is the alternation of generations, and is typical of plant reproduction (Figure 26.9).

The life cycle of higher plants is dominated by the sporophyte stage, with the gametophyte borne on the sporophyte. In ferns, the gametophyte is free-living and very distinct in structure from the diploid sporophyte. In bryophytes, such as mosses, the haploid gametophyte is more developed than the sporophyte.

During the vegetative phase of growth, plants increase in size and produce a shoot system and a root system. As they enter the reproductive phase, some of the branches start to bear flowers. Many flowers are borne singly, whereas some are borne in clusters. The flower is borne on a stalk known as a receptacle. Flower shape, color, and size are unique to each species, and are often used by taxonomists to classify plants.

Sexual Reproduction in Angiosperms

The lifecycle of angiosperms follows the alternation of generations explained previously. The haploid gametophyte alternates with the diploid sporophyte during the sexual reproduction process of angiosperms. Flowers contain the plant’s reproductive structures.

Flower Structure

A typical flower has four main parts—or whorls—known as the calyx, corolla, androecium, and gynoecium (Figure 26.13). The outermost whorl of the flower has green, leafy structures known as sepals. The sepals, collectively called the calyx, help to protect the unopened bud. The second whorl is comprised of petals—usually, brightly colored—collectively called the corolla. The number of sepals and petals varies depending on whether the plant is a monocot or dicot. In monocots, petals usually number three or multiples of three; in dicots, the number of petals is four or five, or multiples of four and five. Together, the calyx and corolla are known as the perianth. The third whorl contains the male reproductive structures and is known as the androecium. The androecium has stamens with anthers that contain the microsporangia. The innermost group of structures in the flower is the gynoecium, or the female reproductive component(s). The carpel is the individual unit of the gynoecium and has a stigma, style, and ovary. A flower may have one or multiple carpels.

If all four whorls (the calyx, corolla, androecium, and gynoecium) are present, the flower is described as complete. If any of the four parts is missing, the flower is known as incomplete. Flowers that contain both an androecium and a gynoecium are called perfect, androgynous or hermaphrodites. There are two types of incomplete flowers: staminate flowers contain only an androecium, and carpellate flowers have only a gynoecium (Figure 26.14).

Adapted from Clark, M.A., Douglas, M., and Choi, J. (2018). Biology 2e. OpenStax. Retrieved from https://openstax.org/books/biology-2e/pages/1-introduction

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