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11.4 Sexual Reproduction

GAMETOGENESIS

Gametogenesis describes the formation of both the male and female gametes,
spermatogenesis is the production of sperm in the male testes and oogenesis is the production
of ova within the ovaries.

Spermatogenesis and oogenesis depend on mitosis and meiosis.

 Male and females meiosis differ in terms of cytoplasm distribution.


 Male: end result is four identical spermatids, which differ in genetic
information.
 Female: ovum receives an unequal share of the cytoplasm and remains
functional while the polar bodies are pockets of genetic information with
minimal cytoplasm and therefore non-functional.

 Timing of meiosis is also different for males and females.

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Spermatogenesis:
 Spermatogenesis occurs continuously from puberty where spermatogonia divide into
spermatozoa in the testes. The seminiferous tubules produce haploid cells. It involves
several steps including mitosis and meiosis. It takes about 74 days in humans. 300
million sperm cells are produced per day.

Meiosis forms the most significant part of process of gametogenesis. Gametogenesis for the
formation of sperms is termed spermatogenesis, while that of ova is called oogenesis. Both
spermatogenesis and oogenesis comprise similar phases of sequential changes viz.,

(i) multiplication phase,

(ii) growth phase and

(iii) maturation phase.

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(a) Multiplication Phase:

At sexual maturity, the undifferentiated primordial germ cells divide several times by mitosis
to produce a large number of spermatogonia (Gr. sperma = seeds, gonos- generation).

(b) Growth Phase:

Each spermatogonium actively grows to a larger primary spermatocyte by obtaining


nourishment from the nursing cells.

(c) Maturation Phase:

Each primary spermatocyte undergoes two successive divisions, called maturation divisions.
The first maturation division is reductional or meiotic. Hence, the primary spermatocyte
divides into two haploid daughter cells called secondary spermatocytes. Both secondary
spermatocytes now undergo second maturation division which is an ordinary mitotic division
to form, four haploid spermatids, by each primary
spermatocyte.

(ii) Formation of Spermatozoa from Spermatids


(Spermatogenesis):

The transformation of spermatids into spermatozoa is


called spermiogenesis. The spermatozoa are later on
known as sperms. Thus four sperms are formed from
one spermatogonium. After spermiogenesis sperm
heads become embedded in the Sertoli cells and are
finally released from the seminiferous tubules by the
process called spermiation.

Significance of Spermatogenesis:

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(i) During spermatogenesis, one spermatogonium produces four sperms, (ii) Sperms have half
the number of chromosomes. After fertilization, the diploid chromosome number is restored
in the zygote. It maintains the chromosome number of the species, (iii) During meiosis I
crossing over takes place which brings about variation, (iv) Spermatogenesis occurs in various
organisms. Thus it supports the evidence of the basic relationship of the organisms.

Oogenesis
The ovary is where gametogenesis and folliculogenesis occur (the growth of the follicle,
which is the structure containing the oocyte).

 Oogenesis occurs by meiosis, however females do not make millions of ova since a
female does not carry millions of fetuses. No more than one ovum is produced per
month therefore females do not have constant mitosis of their germ cells occurring.

 Oogenesis begins during fetal life. During the first trimester of pregnancy the oogonia
(primordial germ cells) of the female fetus divides continuously, mitotically.
 Each oogonia has 46 chromosomes.

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 The oogonia reach a population of 7 million. These oogonia begin the first meiotic
division but stop in prophase I of meiosis I as a primary oocyte and lie within the
ovaries surrounded by other cells, follicular cells (together they comprise the primary
follicle).

 Prior to birth there is a huge reduction in primary oocytes to only 2 million.

 Meiosis does not continue again until puberty, when menarche begins. At puberty only
400,000 oocytes remain in the ovaries. A monthly surge of LH causes the primary
oocyte to continue meiosis from where it left off (prophase I). Every month, one
primary oocyte is hormonally stimulated to begin to enlarge and complete meiosis.
 The end result is a secondary oocyte and a polar body.

 The primary oocyte finishes the first meiotic division by dividing its genetic material
appropriately for meiosis I and undergoing cytokinesis (separation of daughter cells
into two separate cells via a cleavage furrow).
 In oogenesis cytokinesis does not occur down the middle of the parent cell but
unevenly. Only one of the daughter cells can become the ovum, as only one
ovum is released per month. Therefore the one cell that becomes the ovum out
of the four daughter cells receives all the nutrients. The polar bodies die
quickly.

 The ovulated secondary oocyte begins meiosis II but stops at metaphase II.
 Secondary oocyte contains 23 chromosomes, is released from the
ovary.
 Travels down the uterine tube.
 If it does not encounter a spermatozoan, it never undergoes the second
meiotic division but will go through atresia.
 If it encounters a spermatozoan it rapidly completes Anaphase II and
Telophase II to form the ovum and another polar body.
 The ovum is then ready to fuse with the spermatozoan.

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 Only 400 oocytes are used over the reproductive life span of the female.

Oogenesis

(a) Multiplication phase:

In the foetal development, certain cells in the germinal epithelium of the ovary of the foetus
are larger than others. These cells divide by mitosis, producing a couple of million egg mother
cells or oogonia in each ovary of the foetus. No more oogonia are formed or added after birth.
The oogonia multiply by mitotic divisions forming the primary oocytes.

(b) Growth phase:

This phase of the primary oocyte is very long. It may extend over many years. The oogonium
grows into a large primary oocytes. Each primary oocyte then gets surrounded by a layer of
granulosa cells to form primary follicle. A large number of these follicles degenerate during

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the period from birth to puberty. So at puberty only 60,000- 80,000 primary follicles are left
in each ovary. The fluid filled cavity of the follicle is called antrum.

(c) Maturation phase:

Like a primary spermatocyte, each primary oocyte undergoes two maturation divisions, first
meiotic and the second meiotic. The results of maturation divisions in oogenesis are, however,
very different from those in spermatogenesis. In the first, meiotic division, the primary oocyte
divides into two very unequal haploid daughter cells— a large secondary oocyte and a very
small first polar body.

In the second maturation division, the first polar body may divide to form two second polar
bodies. The secondary oocyte again divides into unequal daughter cells, a large ootid and a
very small second polar body. The ootid grows into a functional haploid ovum. Thus from one
oogonium, one ovum and three polar bodies are formed. The ovum, is the actual female
gamete. The polar bodies take no part in reproduction and, hence, soon degenerate.

In human beings, ovum is released from the ovary in the secondary oocyte stage. The
maturation of secondary oocyte is completed in the mother’s oviduct (Fallopian tube) usually
after the sperm has entered the secondary oocyte for fertilization.

In humans (and most vertebrates), the first polar body does not undergo meiosis II, whereas
the secondary oocyte proceeds as far as the metaphase stage of meiosis II. However, it then
stops advancing any further; it awaits the arrival of sperm for completion of meiosis II.

Entry of the sperm restarts the cell cycle breaking down MPF (M-phase promoting factor) and
turning on APC (Anaphase promoting complex). Completion of meiosis II converts the
Compare the processes of spermatogenesis and oogenesis.[8]
secondary oocyte into a fertilized ovum (egg) or zygote (and also a second polar body).

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Hormonal Control of Oogenesis:

GnRH secreted by the hypothalamus stimulates the anterior lobe of pituitary gland to secrete
LH and FSH. FSH stimulates the growth of Graafian follicles and also the development of
egg/oocyte within the follicle to complete the meiosis I to form secondary oocyte. FSH also
stimulates the formation of oestrogens.

LH induces the rupture of the mature Graafian follicle and thereby the release of secondary
oocyte. Thus LH causes ovulation. In brief ovulation in human beings may be defined as the
release of the secondary oocyte from the Graafian follicle. The remaining part of the Graafian
follicle is stimulated by LH to develop into corpus luteum (“yellow body”). The rising level
of progesterone inhibits the release of GnRH, which in turn, inhibits production of FSH, LH
and progesterone.

Significance of Oogenesis:

(i) One oogonium produces one ovum and three polar bodies.

(ii) Polar bodies have small amount of cytoplasm. It helps to retain sufficient amount of
cytoplasm in the ovum which is essential for the development of early embryo. Formation of
polar bodies maintains half number of chromosomes in the ovum.

(iii) During meiosis first crossing over takes place which brings about variation.

(iv) Oogenesis occurs in various organisms. Therefore, it supports the evidence of basic
relationship of the organisms.

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Fertilization involves mechanisms that prevent polyspermy.

Of the 300,000,000 human sperm ejaculated during coitus, only about 200 reach the site of
fertilization in the oviduct. There is evidence that chemical signals released by the follicle
cells that surround the ovulated egg attract the sperm to the egg, but the nature of the
chemoattractant molecules is unknown. Once it finds an egg, the sperm must first migrate
through the layer of follicle cells and then bind to and cross the egg coat—the zona pellucida.
Finally, the sperm must bind to and fuse with the egg plasma membrane. To become
competent to accomplish these tasks, ejaculated mammalian sperm must normally be
modified by conditions in the female reproductive tract, a process called capacitation, which
requires about 5–6 hours in humans. Capacitation is triggered by bicarbonate ions (HCO 3–) in
the vagina, which enter the sperm and directly activate an enzyme in the cytosol.

Capacitation alters the lipid and glycoprotein composition of the sperm plasma membrane,
increases sperm metabolism and motility, and markedly decreases the membrane potential
(that is, the membrane potential moves to a more negative value so that the membrane
becomes hyperpolarized).

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Once a capacitated sperm has penetrated the layer of follicle cells, it binds to the zona
pellucida. The zona usually acts as a barrier to fertilization across species, and removing it
often eliminates this barrier.

1. The acrosome reaction

The zona pellucida of mammalian eggs is composed mainly of three glycoproteins, all of
which are produced exclusively by the growing oocyte.

On binding to the zona, the sperm is induced to undergo the acrosome reaction, in which the
contents of the acrosome are released by exocytosis.

The acrosome reaction is required for fertilization. It exposes various hydrolytic enzymes that
help the sperm tunnel through the zona pellucida, and it exposes other proteins on the sperm
surface that bind to the glycoprotein and thereby help the sperm maintain its tight binding to
the zona while burrowing through it.

2. Fusion of plasma membrane sperm and egg

The acrosome reaction exposes proteins in the sperm plasma membrane that mediate the
binding and fusion of this membrane with that of the egg. Although many sperm can bind to
an egg, normally only one fuses with the egg plasma membrane and injects its nucleus and
other organelles into the egg cytoplasm. This is called fertilization.

3. Cortical reaction and prevention of polyspermy

If more than one sperm fuses—a condition called polyspermy—multipolar or extra mitotic
spindles are formed, resulting in faulty segregation of chromosomes during cell division;
nondiploid cells are produced, and development usually stops.

Two mechanisms can operate to ensure that only one sperm fertilizes the egg. In many cases,
a rapid depolarization of the egg plasma membrane, which is caused by the fusion of the first
sperm, prevents further sperm from fusing and thereby acts as a fast primary block to
polyspermy. But the membrane potential returns to normal soon after fertilization, so that a

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second mechanism is required to ensure a longer-term, secondary block to polyspermy. This is


provided by the egg cortical reaction.

When the sperm fuses with the egg plasma membrane, it causes a local increase in cytosolic
Ca2+, which spreads through the cell in a wave. In some mammalian eggs, the initial increase
in Ca2+ is followed by prolonged Ca2+ oscillations. There is evidence that the Ca 2+ wave or
oscillations are induced by a protein that is introduced into the egg by the sperm, but the
nature of the protein is unknown.

The Ca2+ wave or oscillations activate the egg to begin development, and they initiate the
cortical reaction, in which the cortical granules release their contents by exocytosis.

The contents of the cortical granules include various enzymes that are released by the cortical
reaction and change the structure of the zona pellucida. The altered zona becomes “hardened,”
so that sperm no longer bind to it, and it therefore provides a block to polyspermy. Among the
changes that occur in the zona is the breakdown of the glycoprotein.

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Fertilization in animals can be internal or external.

Once released, egg and sperm alike are destined to die within minutes or hours unless they
find each other and fuse in the process of fertilization. Through fertilization, the egg and
sperm are saved: the egg is activated to begin its developmental program, and the haploid
nuclei of the two gametes come together to form the genome of a new diploid organism. The
mechanism of fertilization has been most intensively studied in marine invertebrates,
especially sea urchins. In these organisms fertilization occurs in sea water, into which huge
numbers of both sperm and eggs are released. Such external fertilization has been more
accessible to study than the internal fertilization of mammals, which normally occurs in the
female reproductive tract after mating.

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Implantation of the blastocyst in the endometrium is essential for


the continuation of pregnancy.
After fertilization in humans, the fertilized ovum divides by mitosis to form 2 diploid nuclei
and the cytoplasm of the fertilized egg cell divides equally to form two celled embryo.

The two cells undergo mitosis and are also capable of replicating their DNA. Further division
gives the embryo the shape of a hollow ball called the blastocyst.

At 7 days, the blastocyst consists about 125 cells and it has reached the uterus. The zona
pellucida now breaks down. Having used all the food reserves in the egg cell, the blastocyst
sink into the endometrium to obtain more nutrients in the process called implantation.

Finger-like projections are developed by the outer lining of the blastocyst to penetrate the
uterus lining. These projections allow exchange of materials between the fetus and the
mother.

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HCG stimulates the ovary to secrete progesterone during early pregnancy.

Human chorionic gonadotropin (hCG)

In pregnancy hCG is critical since it stimulates the corpus luteum, and this maintains
progesterone secretion. Pregnancy depends on the maintenance of the endometrium, which
depends on the continued production of progesterone and oestrogen.

These hormones prevent the degeneration of the uterus lining which is required to support the
developing fetus.

During early pregnancy, the embryo produces hCG hormone. This hormone stimulates the
corpus luteum in the ovary to continue to secrete progesterone and estrogen. These hormones
stimulate the continued development of the uterus wall, which supplies the embryo with
everything it needs.

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The placenta facilitates the exchange of materials between the mother and
fetus.

The placenta is literally the “tree of life.” The derivation of the word placenta comes from
Latin for cake (placenta), from Greek for flat, slab-and from German for mother cake all
referring to the round, flat appearance of the human placenta.

Functionally, the placenta is a highly complex machine:


(1) it acts like a lung in the exchange of oxygen and CO2;
(2) it works as a digestive system, absorbing all necessary nutrients for fetal development and
growth;
(3) it functions as a kidney to remove wastes;
(4) it behaves as an immune barrier that protects the growing fetus from antigen attack from
the maternal system.

The placenta is also an important endocrine organ producing many hormones and growth
factors that regulate the course of pregnancy, support and promote fetal growth, and initiate
parturition. However, all these tasks depend on normal vascular development within the
placenta itself.

Explain the structure and function of the placenta.

[8]

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Estrogen and progesterone are secreted by the placenta once it has formed.
In addition to its role in transporting molecules between mother and fetus, the placenta is a
major endocrine organ. It turns out that the placenta synthesizes a huge and diverse number of
hormones and cytokines that have major influences on ovarian, uterine, mammary and fetal
physiology.

Estrogens: The placenta produces several distinct estrogens. In women, the major estrogen
produced by the placenta is estriol, and the equine placenta synthesizes a unique group of
estrogens not seen in other animals. Depending on the species, placental estrogens are derived
from either fetal androgens, placental progestins, or other steroid precursors.

With few exceptions, the concentration of estrogens in maternal blood rises to maximal
toward the end of gestation. Two of the principle effects of placental estrogens are:

 Stimulate growth of the myometrium and antagonize the myometrial-suppressing


activity of progesterone. In many species, the high levels of estrogen in late gestation
induces myometrial oxytocin receptors, thereby preparing the uterus for parturition.
 Stimulate mammary gland development. Estrogens are one in a battery of hormones
necessary for both ductal and alveolar growth in the mammary gland.

Progestins: Progestins are molecules that bind to the progesterone receptor. Progesterone
itself is often called the hormone of pregnancy because of the critical role it plays in
supporting the endometrium and hence on survival of the conceptus.

The placentae of all mammals examined produce progestins, although the quantity varies
significantly. In some species (women, horses, sheep, cats), sufficient progestin is secreted by
the placenta that the ovaries or corpora lutea can be removed after establishment of the
placenta and the pregnancy will continue. In other animals (cattle, pigs, goats, dogs), luteal
progesterone is necessary throughout gestation because the placenta does not produce
sufficient amounts.

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Progestins, including progesterone, have two major roles during pregnancy:

 Support of the endometrium to provide an environment conducive to fetal survival. If


the endometrium is deprived of progestins, the pregnancy will inevitably be
terminated.
 Suppression of contractility in uterine smooth muscle, which, if unchecked, would
clearly be a disaster. This is often called the "progesterone block" on the myometrium.
Toward the end of gestation, this myometrial-quieting effect is antagonized by rising
levels of estrogens, thereby facilitating parturition(birth).

Progesterone and other progestins also potently inhibit secretion of the pituitary
gonadotropins luteinizing hormone and follicle stimulating hormone. This effect almost
always prevents ovulation from occurring during pregnancy.

Application: The average 38-week pregnancy in humans can be positioned on


a graph showing the correlation between animal size and the development of
the young at birth for other mammals.
The gestation period of humans is 266 days, 8 days short of nine months. Many such
biological characteristics have an explanation in terms evolutionary survival value. Even
some seemingly cultural characteristics of human society have such a biological explanation.
The gestation periods of other animal reveals that the length of the gestation period is largely
a matter of the animal specie's size. Asian elephants have a gestation period of 645 days and
African elephants 640 days. Dogs and other canines have a gestation period of about 60,
which is also the period for cats. Even within the species of apes and monkeys gestation
period seem to be a matter of size. For Rhesus monkeys it is 164 days and baboons 187 days.
For small animals such as rabbits the period is about 33 days and for mice about 20 days.

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NOS: Assessing risks and benefits associated with scientific research—the risks to
human male fertility were not adequately assessed before steroids related to
progesterone and estrogen were released into the environment as a result of the use of
the female contraceptive pill.

The past several years have seen a steady drumbeat of news reports, blog posts, and scientific
studies which have raised concerns about the presence of estrogenic compounds (natural
estrogens and synthetic chemicals that mimic natural estrogen) in waterways and drinking
water, and potential harm to human health or aquatic life. Frequently, one particular synthetic
estrogen has been singled out for purportedly detrimental effects on the environment: ethynyl-
estradiol, or EE2, a synthetic estrogen used in birth control pills, patches, rings, and
injectables. Journalists from along the political spectrum and anti-contraception advocates
alike have seized on the idea of the-Pill-as-environmental-pollutant, and it has been difficult
to separate environmental health concern from sensational coverage or politically motivated
rhetoric.

At the same time, anecdotal evidence suggests environmental footprint or “greenness” is


increasingly one of the factors that many women consider in choosing among birth control
methods, along with effectiveness, safety, convenience, cost, and acceptability. This raises the
possibility that reports about the environmental impact of hormonal contraception could

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influence a woman's choice of this method, and underscores the need to understand the
current science and provide women with unbiased information that allows them to make
informed choices.

The effect of estrogenic compounds in the water supply from industry, agriculture, and other
sources raises concerns about human health and deserves scrutiny. Estrogenic compounds are
part of a larger category of chemicals known as endocrine-disruptors (EDCs), chemicals that
can alter the hormonal and homeostatic systems enabling an organism—like a human being or
other animal—to communicate with and respond to its environment

Birth is mediated by positive feedback involving estrogen and oxytocin.


Birth takes place at the end of the third trimester, 38 weeks from contraception. Hormones in
the female body induce placenta cells of the mother to produce hormone estrogen that cause
the smooth muscle of the uterine wall to contract. The pressure of the fetus head against the
cervix triggers the hypothalamus to release the hormone oxytocin from the pituitary gland.
Working together, oxytocin and estrogen stimulate waves of contraction in the walls of the
uterus. This is known as the positive feedback loop.

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The contraction occurs each hour increasing to every 2-3 minutes. Eventually strong
contractions aided by the mother’s pushing, expel the fetus through the vagina. The fetus is
now a newborn.

Oxytocin has been best known for its roles in female reproduction. It is released in large
amounts during labor, and after stimulation of the nipples. It is a facilitator for childbirth and
breastfeeding. One of the oldest applications of oxytocin as a proper drug is as a therapeutic
agent during labor and delivery. It is a stimulant widely employed to induce or augment labor,
especially at term, when adequate oxytocin receptors are present.

Outline the hormonal control of the process of birth. [4]

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