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0 REPRODUCTION
Reproduction is a biological process by which an organism gives rise to an
offspring that is biologically similar to the organism. It involves the segregation
of a portion of the parent organism by sexual or asexual process which
eventually grows and differentiates into a new individual. The process enables
and ensures the continuity of species from generation to generation. It is the
mechanism by which life continues to exist on earth.

Reproductive cycles
Reproductive cycles and patterns vary extensively among mammals. Most
animals exhibit cycles in reproductive activity, usually related to changing
seasons. This allows animals to conserve resources and reproduce when more
energy is available and when environmental conditions favour the survival of
offspring. Reproductive cycles are controlled by a combination of
environmental and hormonal cues. Environmental cues may include seasonal
temperature, rainfall, day length, and lunar cycles.

Asexual and sexual reproduction

Reproduction in animals can be either asexual or sexual they may alternate
between the two modes, depending on environmental conditions. E.g. Daphnia
reproduce by parthenogenesis under favorable conditions and sexually during
times of environmental stress. Asexual reproduction involves the formation of
individuals whose genes come from a single parent and there is no fusion of
sperm and egg. Sexual reproduction on the other hand is the formation of
offspring by the fusion of haploid gametes to form a diploid zygote. Oviparity
is a mode of reproduction in which animals lay eggs. It involves the oviposition
of an embryo within a shelled egg requiring some period of embryonic
development outside of the female reproductive tract. E,g. birds and some
reptiles are oviparous. Viviparity is the production of live young ones. It
involves retaining fertilized eggs in or on a parent’s body until the embryo
develops into a complete organism and birthed live. The embryo has a
connecting tissue with the mother through which it is fed and sustained till birth.
E,g, Humans, dogs, cattle are viviparous. Ovoviviparity is a mode of

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reproduction in which animals lay eggs and keep them inside the mother’s body
until they are ready to hatch. The embryo depend on the yolk in the egg not the
mother for nutrition. E.g. Some fishes, snakes, insects.

Asexual reproduction
There are different mechanisms of asexual reproduction enable animals to
produce identical offspring rapidly.
i) Fission - In which a parent separates into two or more approximately equal-
sized individuals.
ii) Budding – This is also common among invertebrates. This is a form of
asexual reproduction in which new individuals split off from existing ones.
iii) Fragmentation – In this process, the body breaks into several pieces, some or
all of which develop into complete adults. Reproducing in this way involves the
regeneration of lost body parts. Many animals can also replace new appendages
by regeneration.
iv) Parthenogenesis is the process by which an unfertilized egg develops
without being fertilized. Parthenogenesis plays a role in the social organization
of some bees, wasps, and ants. Male honeybees (drones) are haploid, and female
honeybees (queens and workers) are diploid. Several genera of fishes,
amphibians, and lizards reproduce by a form of parthenogenesis that produces
diploid zygotes.

Advantages of sexual reproduction

Asexual reproduction has a number of advantages.
i) It allows isolated animals to reproduce without needing to find a mate.
ii) It can create numerous offspring in a short period of time.
iii) In stable environments, it allows for the perpetuation of successful
genotypes.

Sexual reproduction
In sexual reproduction, the female gamete, the unfertilized egg, or ovum, is
usually large and nonmotile while the male gamete which is the sperm, is
usually small and motile. Sexual reproduction increases genetic variation among

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offspring by generating unique combinations of genes inherited from two
parents. In sexual reproduction, there are some instances in which one
individual functions as both a male and a female. They have both the male and
female gametes. This is called HERMAPHRODITISM and the organisms are
referred to as HERMAPHRODITES. Some hermaphrodites can self-fertilize,
but most mate with another member of the same species. In such matings, each
individual receives and donates sperm, and this results in twice as many
offspring as would be produced if only one set of eggs were fertilized. In
sequential hermaphroditism, an individual reverses its sex during its lifetime. In
some species, the sequential hermaphrodite is female first. In other species, the
sequential hermaphrodite is male first.

5.1 The reproductive system

Reproductive systems produce gametes and make them available to gametes of
the opposite sex. The least complex reproductive systems lack gonads which are
the organs that produce gametes in most animals. E.g. Polychaete worms lack
gonads. Their eggs and sperm develop from undifferentiated cells lining the
coelom. As the gametes mature, they are released from the body wall and fill the
coelom. In some species, the body splits open to release the gametes, killing the
parent. Most insects have separate sexes with complex reproductive systems. In
many species, the female reproductive system includes a spermatheca, a sac in
which sperm may be stored for a year or more.
The reproductive systems of vertebrates are similar., however, there are some
variations.
i) In many nonmammalian vertebrates, the digestive, excretory, and
reproductive systems share a common opening to the outside, the cloaca.
ii) Mammals have separate openings for the digestive and reproductive systems.
iii) Female mammals also have separate openings for the excretory and
reproductive systems.
Iv The uterus of most vertebrates is partly or completely divided into two
chambers.
v) Male reproductive systems differ mainly in copulatory organs.

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vi) Many mammalian vertebrates do not have a well-developed penis and
simply turn the cloaca inside out to ejaculate.

Human reproductive organs

The reproductive organs in humans produce and transport gametes. Human
reproduction involves intricate anatomy and complex behavior.
The male’s external reproductive organs consist of the scrotum and penis. The
internal reproductive organs consist of gonads that produce sperm and
hormones, accessory glands that secrete products essential to sperm movement,
and ducts to carry the sperm and glandular secretions. The male gonads, or
testes, consist of highly coiled tubes surrounded by layers of connective tissue.
The tubes are called seminiferous tubules, where sperm are produced. Leydig
cells scattered between the seminiferous tubules produce testosterone and other
androgens. The scrotum, a fold in the body wall, holds the testes outside the
body cavity at a temperature about 2°C below that of the abdomen. This keeps
testicular temperature cooler than that in the body cavity. The testes develop in
the body cavity and descend into the scrotum just before birth. From the
seminiferous tubules of the testes, the sperm pass through the coiled tubules of
the epididymis. As they pass through this duct, sperm become motile and gain
the ability to fertilize an egg. Ejaculation propels sperm from the epididymis to
the vas deferens. The vas deferens run from the scrotum and behind the urinary
bladder. Each vas deferens joins with a duct from the seminal vesicle to form an
ejaculatory duct. The ejaculatory ducts open into the urethra. The urethra drains
both the excretory and reproductive systems. Accessory sex glands add
secretions to semen. A pair of seminal vesicles contributes about 60% of total
semen volume. Seminal fluid is thick, yellowish, and alkaline. It contains
mucus, fructose, a coagulating enzyme, ascorbic acid, and prostaglandins. The
prostate gland secretes directly into the urethra. Prostatic fluid is thin and milky.
This fluid contains anticoagulant enzymes and citrate.

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 Fig 5.1 The male reproductive organ

The bulbourethral glands are a pair of small glands along the urethra below the
prostate. Prior to ejaculation, they secrete clear mucus that neutralizes any
acidic urine remaining in the urethra. Bulbourethral fluid also carries some
sperm released before ejaculation. This is one of the reasons the withdrawal
method of birth control has a high failure rate. A male usually ejaculates about
2–5 mL of semen, with each milliliter containing about 50–130 million sperm.
Once in the female reproductive tract, prostaglandins in semen thin the mucus at
the opening of the uterus and stimulate uterine contractions that help move the
semen. When ejaculated, semen coagulates, making it easier for uterine
contractions to move it along. Anticoagulants then liquefy the semen, and the
sperm begin swimming. The alkalinity of semen helps neutralize the acidic
environment of the vagina, protecting the sperm and increasing their motility.
The human penis is composed of three layers of spongy erectile tissue. During

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sexual arousal, the erectile tissue fills with blood from arteries. The resultant
increased pressure seals off the veins that drain the penis, causing it to engorge
with blood. The engorgement of the penis with blood causes an erection, which
is essential for the insertion of the penis into the vagina.
The penis of some mammals possesses a baculum, a bone that helps stiffen the
penis. The sensitive head, or glans penis, is covered by thinner skin. The glans
is covered by the foreskin, or prepuce, which may be removed by circumcision.

Fig 5.2 Female reproductive organ

The reproductive anatomy of the human female includes external and internal
reproductive structures. External reproductive structures consist of two sets of
labia surrounding the clitoris and vaginal opening. Internal reproductive organs
consist of a pair of gonads and a system of ducts and chambers. The role of the
ducts and chambers is to conduct the gametes and house the embryo and foetus.
The ovaries, the female gonads, lie in the abdominal cavity, attached to the
uterus by a mesentery. Each ovary is enclosed in a tough protective capsule and
contains many follicles. Each follicle consists of one egg cell surrounded by one
or more layers of follicle cells. A female is born with about 400,000 follicles.

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Only several hundred of these will release eggs during a female’s reproductive
years. Follicles produce the primary female sex hormones, estrogens. Usually
one follicle matures and releases its egg during each menstrual cycle in the
process of ovulation. After ovulation, the remaining follicular tissue develops
into the corpus luteum. The corpus luteum secretes additional estrogens and
progesterone, hormones that help maintain the uterine lining during pregnancy.
If pregnancy does not occur, the corpus luteum disintegrates and a new follicle
matures during the next cycle. At ovulation, the egg is released into the
abdominal cavity near the opening of the oviduct. The cilia-lined funnel-like
opening of the oviduct draws in the egg. Cilia convey the egg through the
oviduct to the uterus. The highly vascularized inner lining of the uterus is called
the endometrium. The neck of the uterus, the cervix, opens into the vagina.
The vagina is a thin-walled chamber that forms the birth canal and is the
repository for sperm during copulation. It opens to the outside at the vulva, the
collective term for the external female genitalia. The vaginal opening is partially
covered by a thin sheet of tissue called the hymen. The vaginal and urethral
openings are located within a recess called the vestibule. The vestibule is
surrounded by a pair of slender folds called the labia minora. The labia majora
enclose and protect the labia minora and vestibule. The clitoris is found at the
front edge of the vestibule. During sexual arousal, the clitoris, vagina, and labia
engorge with blood and enlarge. During sexual arousal, Bartholin’s glands
secrete mucus into the vestibule, providing lubrication and facilitating
intercourse.
Mammary glands are present in both males and females but normally function
only in females. They are not a component of the human reproductive system
but are important to mammalian reproduction. Within the glands, small sacs of
epithelial tissue secrete milk, which drains into a series of ducts opening at the
nipple. Adipose tissue forms the main mass of the mammary gland of a
nonlactating mammal. The low estrogen level in males prevents the
development of the sensory apparatus and fat deposits, so that male breasts
remain small, with nipples unconnected to the ducts.

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5.2 Spermatogenesis/ oogenesis
Gametogenesis is based on meiosis. Spermatogenesis and oogenesis both
involve meiosis but differ in three significant ways.

Spermatogenesis
Spermatogenesis is the production of mature sperm cells from spermatogonia.
Spermatogenesis is a continuous and prolific process in the adult male. Each
ejaculation contains 100–650 million sperm. Spermatogenesis occurs in
seminiferous tubules. Primordial germ cells of the embryonic testes differentiate
into spermatogonia, the stem cells that give rise to sperm. As spermatogonia
differentiate into spermatocytes and then into spermatids, meiosis reduces the
chromosome number from diploid to haploid. As spermatogenesis progresses,
the developing sperm cells move from the wall to the lumen of a seminiferous
tubule and then to the epididymis, where they become motile. The structure of
sperm fits its function. A head containing the haploid nucleus is tipped with an
acrosome, which contains enzymes that help the sperm penetrate to the egg.
Behind the head are a large number of mitochondria (or a single large one) that
provide ATP to power the flagellum.

Oogenesis
Oogenesis is the production of ova from oogonia. Oogenesis differs from
spermatogenesis in three major ways.
i) At birth an ovary may contain all of the primary oocytes it will ever have,
while sperm are produced from spermatogonia throughout a man’s life.
ii) Unequal cytokinesis during meiosis results in the formation of a single large
secondary oocyte and three small polar bodies. The secondary oocyte becomes
the ovum, while the polar bodies degenerate. In spermatogenesis, all four
products of meiosis become mature sperm.
iii) Oogenesis has long resting periods. Spermatogenesis produces mature sperm
from spermatogonia in an uninterrupted sequence.
Oogenesis begins in the female embryo with differentiation of primordial germ
cells into oogonia, ovary-specific stem cells. An oogonium multiplies by mitosis
and begins meiosis, but the process stops at prophase I. The primary oocytes

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remain quiescent within small follicles until puberty. Beginning at puberty,
follicle-stimulating hormone (FSH) stimulates a follicle to grow and induces its
primary oocyte to complete meiosis I and start meiosis II. It is arrested at
metaphase II as a secondary oocyte. The secondary oocyte is released when the
follicle breaks open at ovulation. Meiosis is completed when a sperm penetrates
the oocyte. Oogenesis is completed, producing an ovum. The haploid nuclei of
the sperm and ovum fuse in fertilization. The ruptured follicle develops into the
corpus luteum. If the released oocyte is not fertilized, the corpus luteum
degenerates.

Fertilzation
Fertilization depends on mechanisms that help sperm meet eggs of the same
species. The mechanisms of fertilization, the union of sperm and egg, play an
important part in sexual reproduction. In external fertilization, eggs are released
by the female into a wet environment, where they are fertilized by the male. A
moist habitat is almost always required for external fertilization to prevent
gametes from drying out and to allow the sperm to swim to the eggs. In external
fertilization, timing is crucial to ensure that mature sperm encounter ripe eggs.
Environmental cues such as temperature or day length may cause gamete
release by the whole population. Individuals may engage in courtship behavior
that leads to fertilization of the eggs of one female by one male.
In internal fertilization, sperm are deposited in or near the female reproductive
tract, and fertilization occurs within the tract. Internal fertilization is an
adaptation to terrestrial life that enables sperm to reach an egg in a dry
environment. Internal fertilization requires sophisticated reproductive systems,
including copulatory organs that deliver sperm and receptacles for their storage
and transport to ripe eggs. Mating animals may use pheromones which are
chemical signals released by one organism that influence the behavior or
physiology of other individuals of the same species. Pheromones are small,
volatile, or water-soluble molecules that disperse into the environment. Like
hormones, pheromones are active in minute amounts. Many pheromones act as
male attractants. In internal fertilization, All species produce more offspring
than can survive to reproduce. Internal fertilization usually involves the

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production of fewer zygotes than external fertilization. However, the survival
rate is higher for internal fertilization. Major types of protection mechanisms for
internal fertilization include tough eggshells, development of the embryo within
the reproductive tract of the mother, and parental care of the eggs and offspring.
Marsupial mammals retain their embryos for only a short period in the uterus.
The embryos crawl out and complete foetal development attached to a
mammary gland in the mother’s pouch. The embryos of eutherian mammals
develop entirely within the uterus, nourished through the placenta. Parental care
of offspring can occur regardless of whether fertilization is external or internal.

5.3 Hormeones regulating the reproductive system: menstrual cycle and

birth.
The main reproductive hormones are Estrogen, testosterone and progesterone.
They are produced in the ovaries in females and in the testes in males and are
responsible for pregnancy, puberty, menstruation, sperm production and all
other aspects of sexuality and fertility.
Estrogen – Promotes the maturation and release of an ovum in every menstrual
cycle. It is also produced by the placenta during pregnancy,
Progesterone -Supports menstruation and pregnancy. The major function is to
prepare the lining of the uterus (endometrium) for a fertilized egg to implant
and grow. If pregnancy does not occur, the endometrium sheds during the
menstrual period. However if pregnancy occurs. Progesterone increases to
support the pregnancy and soppresses the production of estrogen.
The hormones controlling the female reproductive system include
Gonadotropin-releasing hormone (GnRH), which stimulates the pituitary gland
to secrete small amounts of Follicle-stimulating hormone (FSH) and Luteinizing
hormone (LH). They are all produced in the brain. Human chorionic
gonadotropin (HCG).
GnRH is produced in the hypothalamus in the brain. When it circulates in the
blood, it causes the release of FSH and LH from the pituitary gland in another
part of the brain. FSH is produced during the first half of the menstrual cycle. It
stimulates development of the maturing ovarian follicle and controls ovum
production in the female and sperm production in the male. LH stimulates the

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ovaries to produce oestrogen and progesterone. It triggers the release of the
mature ovum from the ovary called Ovulation. LH also promotes the
development of the corpus lutem. The corpus is a collection of cells that form in
the ovaryafter the ovarian follicle releases a mature ovum during ovulation. Its
function is to make the uterus suitable for a foetus to grow.

Humans and many other primates have menstrual cycles whereby if pregnancy
does not occur, the endometrium is shed through the cervix and vagina in
menstruation. Other mammals have oestrous cycles whereby if pregnancy does
not occur, the uterus reabsorbs the endometrium. Oestrous cycles are associated
with more pronounced behavioral cycles than are menstrual cycles. In the
oestrus cycle, the period of sexual activity is the only time the condition of the
vagina permits mating. However, human females may be sexually receptive
throughout their menstrual cycle. The term menstrual cycle refers specifically to
the changes that occur in the uterus, and is also called the uterine cycle. It is
caused by cyclic events that occur in the ovaries, the ovarian cycle.

The cycle begins with the release from the hypothalamus of Gonadotropin-
releasing hormone (GnRH, which stimulates the pituitary gland to secrete small
amounts of Follicle-stimulating hormone (FSH) and Luteinizing hormone (LH).
FSH stimulates follicle growth, aided by LH, and the cells of the growing
follicles start to make estrogen. There is a slow rise in estrogen secreted during
the follicular phase, the part of the ovarian cycle in which follicles are growing
and oocytes maturing.
The low level of estrogen inhibits secretion of the pituitary hormones, keeping
FSH and LH levels low. The levels of FSH and LH shoot up when the secretion
of estrogen by the growing follicle rises sharply. The high concentration of
estrogen stimulates the secretion of gonadotropins by acting on the
hypothalamus to increase its output of GnRH. This stimulates the secretion of
FSH and LH. LH secretion is especially high, because the high concentration of
estrogen increases the sensitivity of LH-releasing cells in the pituitary to GnRH.
LH induces the final maturation of the follicle and ovulation. The follicle and
adjacent wall of the ovary rupture, releasing the secondary oocyte. Following

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ovulation, during the luteal phase of the ovarian cycle, LH stimulates the
transformation of the follicle into the corpus luteum, a glandular structure.
Under the continued stimulation by LH during this phase, the corpus luteum
secretes progesterone and estrogen. As the levels of these hormones rise, they
exert negative feedback on the hypothalamus and pituitary, inhibiting the
secretion of LH and FSH. Near the end of the luteal phase, the corpus luteum
disintegrates, causing concentrations of estrogen and progesterone to decline.
The pituitary and hypothalamus are liberated from the inhibitory effects of these
hormones. The pituitary begins to secrete enough FSH to stimulate the growth
of new follicles in the ovary, initiating the next ovarian cycle. The follicular
phase of the ovarian cycle is coordinated with the proliferative phase of the
menstrual cycle. Secretion of estrogens during the follicular phase stimulates
endometrial thickening. The estrogen and progesterone of the luteal phase
stimulate development and maintenance of the endometrium, including the
enlargement of arteries and the growth of endometrial glands. The glands
secrete a nutrient fluid that can sustain an early embryo before it implants in the
uterine lining. Thus, the luteal phase of the ovarian cycle is coordinated with the
secretory phase of the uterine cycle. The rapid drop in ovarian hormones as the
corpus luteum disintegrates causes spasms in the uterine lining, depriving it of
blood. The upper two-thirds of the endometrium disintegrates, resulting in
menstruation, or the menstrual flow phase of the uterine cycle, and the
beginning of a new cycle.
During menstruation, new ovarian follicles begin to grow. Estrogen is also
responsible for female secondary sex characteristics, including deposition of fat
in the breasts and hips, increased water retention, and stimulation of breast
development. It also influences sexual behavior. Menopause, the cessation of
ovarian and menstrual cycles, usually occurs between ages 46 and 54. During
these years, the ovaries lose their responsiveness to FSH and LH, and
menopause results from a decline in estrogen production by the ovary.

In humans and other mammals, a complex interplay of hormones regulates

gametogenesis. In females, the secretion of hormones and the reproductive

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events they regulate are cyclic. Hormonal control of the female cycle is
complex.
The principal sex hormones in the male are the androgens. These are steroid
hormones produced mainly by the Leydig cells of the testes which are
interstitial cells near the seminiferous tubules. Testosterone, the most important
male androgen, and other androgens are responsible for the primary and
secondary male sex characteristics. Primary sex characteristics are associated
with the development of the vas deferens and other ducts, development of the
external reproductive structures, and sperm production. Secondary sex
characteristics are features not directly related to the reproductive system,
including deepening of the voice, distribution of facial and pubic hair, and
muscle growth. Androgens also affect behavior. In addition to specific sexual
behaviors and sex drive, androgens increase general aggressiveness. They are
responsible for vocal behavior, like singing in birds and calling by frogs.
Hormones from the anterior pituitary gland and hypothalamus control androgen
secretion and sperm production by the testes.
In humans and other placental mammals, an embryo grows into a newborn in
the mother’s uterus. In placental mammals, pregnancy or gestation is the
condition of carrying one or more embryos. A human pregnancy averages 266
days. Many rodents have gestation periods of 21 days. Cows have a gestation of
27 days, and elephant gestation lasts 600 days.
Fertilization or conception occurs in the oviduct. Twenty-four hours later,
cleavage begins. Three to four days after fertilization, the embryo reaches the
uterus as a ball of cells. By one week past fertilization, the blastocyst forms as a
sphere of cells containing a cavity. After a few more days, the blastocyst
implants in the endometrium. The embryo secretes hormones to signal its
presence and control the mother’s reproductive system. Human chorionic
gonadotropin (HCG) acts like pituitary LH to maintain secretion of
progesterone and estrogens by the corpus luteum for the first few weeks of
pregnancy. Some HCG is excreted in the urine, where it is detected by
pregnancy tests.
Human gestation is divided into three trimesters of three months each. For the
first 2–4 weeks of development, the embryo obtains nutrients from the

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endometrium. The outer layer of the blastocyst, called the trophoblast invades
the endometrium, eventually helping to form the placenta. The placenta allows
diffusion of material between maternal and embryonic circulations, providing
nutrients, exchanging respiratory gases, and disposing of metabolic wastes for
the embryo. Blood from the embryo travels to the placenta and returns via the
umbilical vein. Organogenesis occurs during the first trimester. By the end of
week four, the heart is beating. By the end of week eight, all the major
structures of the adult are present in rudimentary form. The rapidity of
development makes this a time when the embryo is especially sensitive to
environmental insults such as radiation or drugs. High levels of progesterone
initiate changes in the maternal reproductive system. These include
- Increased mucus in the cervix to form a protective plug.
- Growth of the maternal part of the placenta, enlargement of the uterus.
- Cessation of ovarian and menstrual cycling.
- The breasts enlarge rapidly and may be very tender.
During the second trimester, the foetus grows rapidly to 30 cm and is very
active. The mother may feel movements during the early part of the second
trimester. Hormonal levels stabilize as HCG declines, the corpus luteum
deteriorates, and the placenta takes over the secretion of progesterone, which
maintains the pregnancy.
During the third trimester, the foetus grows rapidly to about 3–4 kg in weight
and 50 cm in length. Foetal activity may decrease as the foetus fills the space
available to it. Maternal abdominal organs become compressed and displaced,
leading to frequent urination, digestive blockages, and back strain.
A complex interplay of local regulators (prostaglandins) and hormones
(estrogen and oxytocin) induces and regulates labor. The mechanism that
triggers labor is not fully understood. In one possible model, high levels of
estrogen induce the formation of oxytocin receptors on the uterus. Oxytocin,
produced by the foetus and the mother’s posterior pituitary, stimulates powerful
contractions by the smooth muscles of the uterus. Oxytocin also stimulates the
placenta to secrete prostaglandins, which enhance the contractions. The physical
and emotional stress associated with the contractions stimulate the release of

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more oxytocin and prostaglandins, a positive feedback system that underlies the
process of labor.
Birth, or parturition, is brought about by strong, rhythmic uterine contractions.
The process of labor has three stages.
i) The first stage is the opening and thinning of the cervix, ending in complete
dilation.
ii) The second stage is the expulsion of the baby as a result of strong uterine
contractions.
i) The third stage is the expulsion of the placenta.

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