REPRODUCTION IN ANIMALS

External and Internal Fertilization

External Fertilization
External fertilization usually occurs in aquatic environments where both eggs and
sperm are released into the water. After the sperm reaches the egg, fertilization
takes place. Most external fertilization happens during the process of spawning
where one or several females release their eggs and the male(s) release sperm in
the same area, at the same time.
The release of the reproductive material may be triggered by water temperature or
the length of daylight. Nearly all fish spawn, as do crustaceans (such as crabs and
shrimp), mollusks (such as oysters), squid, and echinoderms (such as sea urchins
and sea cucumbers). Figure 1a shows salmon spawning in a shallow stream. Frogs,
like those shown in Figure 1b, corals, mayflies, and mosquitoes also spawn.

Figure 1. (a) Salmon reproduce through spawning. (b) During sexual reproduction
in toads, the male grasps the female from behind and externally fertilizes the eggs
as they are deposited. (credit a: Dan Bennett; credit b: “OakleyOriginals”/Flickr)
Pairs of fish that are not broadcast spawners may exhibit courtship behavior. This
allows the female to select a particular male. The trigger for egg and sperm release
(spawning) causes the egg and sperm to be placed in a small area, enhancing the
possibility of fertilization.

External fertilization in an aquatic environment protects the eggs from drying out.
Broadcast spawning can result in a greater mixture of the genes within a group,
leading to higher genetic diversity and a greater chance of species survival in a
hostile environment. For sessile aquatic organisms like sponges, broadcast
spawning is the only mechanism for fertilization and colonization of new
environments. The presence of the fertilized eggs and developing young in the
water provides opportunities for predation resulting in a loss of offspring.
Therefore, millions of eggs must be produced by individuals, and the offspring
produced through this method must mature rapidly. The survival rate of eggs
produced through broadcast spawning is low.

Internal Fertilization
Internal fertilization occurs most often in land-based animals, although some
aquatic animals also use this method. 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. This occurs in most
bony fish, many reptiles, some cartilaginous fish, most amphibians, two mammals,
and all birds. Reptiles and insects produce leathery eggs, while birds and turtles
produce eggs with high concentrations of calcium carbonate in the shell, making
them hard. Chicken eggs are an example of this second type.

In ovoviparity, fertilized eggs are retained in the female, but the embryo obtains
its nourishment from the egg’s yolk and the young are fully developed when they
are hatched. This occurs in some bony fish (like the guppy Lebistes reticulatus),
some sharks, some lizards, some snakes (such as the garter snake Thamnophis
sirtalis), some vipers, and some invertebrate animals (like the Madagascar hissing
cockroach Gromphadorhina portentosa).

In viviparity the young develop within the female, receiving nourishment from the
mother’s blood through a placenta. The offspring develops in the female and is
born alive. This occurs in most mammals, some cartilaginous fish, and a few
reptiles.

Internal fertilization has the advantage of protecting the fertilized egg from
dehydration on land. The embryo is isolated within the female, which limits
predation on the young. Internal fertilization enhances the fertilization of eggs by a
specific male. Fewer offspring are produced through this method, but their survival
rate is higher than that for external fertilization.

Animal Reproductive Structures and Functions

Diversity of Animal Reproductive Anatomy
The reproductive structures of many animals are very similar, even across different
lineages, in a process that begins with two gametes–eggs and sperm–and ends with
a zygote, which is a fertilized egg. In animals ranging from insects to humans,
males produce sperm in testes, and sperm are stored in the epididymis until
ejaculation. Sperm are small, mobile, low-cost cells that occur in high numbers.
Females produce an ovum or egg that matures in the ovary. Eggs are large cells
that require a substantial investment of time and energy to form, are non-mobile,
and are rare relative to sperm numbers. When the eggs are released from the ovary,
they travel to the uterine tubes for fertilization (in animals that reproduce via
internal fertilization) or are released in the aqueous environment (in animals that
reproduce via external fertilization).

For our purposes, all sexual reproducers have females with ovaries that produce
large eggs, which subsequently travel down a uterine tube, and males with testes
that produce small, plenteous sperm, stored in an epididymus. Of course, beyond
this general anatomy, there are some interesting differences.

Different types of animals:

 In some invertebrate species, including many insects and some mollusks and
worms, the female has a spermatheca: a specialized sac which stores sperm
for later use, sometimes up to a year. Fertilization can be timed with
environmental or food conditions that are optimal for offspring survival.

 Non-mammal vertebrates, such as most birds and reptiles, have a cloaca: a

single body opening which functions in the digestive, excretory and
reproductive systems. Mating between birds usually involves positioning the
 cloaca openings opposite each other for transfer of sperm from male to
female. Ducks are a rare exception, where the males have a penis.

 Mammals have separate openings for the systems in the female, and
placental mammals have a uterus for support of developing offspring. The
uterus has two chambers in species that produce large numbers of offspring
at a time, while species that produce one offspring, such as primates, have a
single chamber.
.
Gametogenesis, the production of sperm and eggs, takes place through the process
of meiosis. Meiosis produces haploid cells with half the number of chromosomes
normally found in diploid cells. Hormones are dynamic (changing), so this process
can be trickier to understand than basic anatomy or gametogenesis.
Hormonal changes are the center of the fascinating biology of reproduction. The
human male and female reproductive cycles are both controlled by the interaction
of hormones from the hypothalamus and anterior pituitary with hormones from
reproductive tissues and organs: the hypothalamus sends a gonadotropin-
releasing hormone (GnRH) to the anterior pituitary, and follicle stimulating
hormone (FSH) and luteinizing hormone (LH) are release from the anterior
pituitary into the blood as a result. Although FSH and LH are named after their
functions in female reproduction, they are produced and play important roles in
controlling reproduction in both sexes.

Female Reproductive Anatomy

A number of reproductive structures are exterior to the female’s body. These
include the breasts and the vulva. Internal female reproductive structures
include ovaries, oviducts, the uterus, and the vagina, shown below.
Humans females become capable of reproduction at sexual maturity, which follows
puberty. During puberty, the hypothalamus in the brain signals the pituitary gland
to produce two hormones, follicle-stimulating hormone (FSH) and luteinizing
hormone (LH). In females, FSH and LH stimulate the ovaries to produce the
female sex hormones, estrogen and progesterone. This results in the development
of secondary sex characteristics (such as breasts) and causes the ovaries to begin
producing mature eggs.
This table briefly summarizes the major organs, locations, and functions of
mammalian female reproductive anatomy:

Organ Function

Ovaries Produces and develops eggs

cFallopian tubes Transports egg to uterus, acts as site of fertilization

 Organ Function

(oviducts)

Uterus Supports a developing embryo

Cervix Allows passage between the uterus and the vagina

Receives penis during intercourse, acts as birth

Vagina canal, passes menstrual flow

Breasts Produce and deliver milk

Ovaries are the site of egg development. Egg development occurs in structures
called follicles, which are lined with specialized cells called follicular cells that
surround the egg and promote egg development. During the menstrual cycle, a
batch of follicular cells develops and prepares the eggs for release. At ovulation,
one follicle ruptures and one egg is released, as illustrated below. The ruptured
follicle, which remains in the ovary, is then called the corpus luteum, which
secretes hormones that prevent menstruation until the egg has had time to be
fertilized. If fertilization and implantation in the uterine wall occurs, then the
corpus luteum continues to prevent menstruation; if fertilization does not occur,
then the corpus luteum degenerates and menstruation occurs.
Oocyte maturation within a follicle, followed by ovulation (follicle rupture). The
follicle becomes a corpus luteum after ovulation and degenerates if the egg is not
fertilized.

The oviducts, or fallopian tubes, extend from the uterus to the ovaries, but they
are not in direct physical contact with the ovaries. The ends of the oviducts flare
out into a trumpet-like structure and have a fringe of finger-like projections called
fimbriae. When an egg is released at ovulation, the fimbrae help the egg enter into
the tube and passage to the uterus. Fertilization (the union of sperm and egg)
usually takes place within the oviducts and the developing embryo is moved
toward the uterus for development. It usually takes the egg or embryo a week to
travel through the oviduct.
Female Gametogenesis: Oogenesis.
Oogenesis, the process of producing an egg cell, occurs in the the ovaries. Egg
stem cells, called oogonia, divide by mitosis to produce up to 2 million oocytes (a
precursor to the egg). The process of oogenesis begins while the female is still an
embryo undergoing development: the oocytes start the process of meiosis and then
pause during meiotic prophase I. Because this process occurs during embryonic
development, this means that a female mammal is born with every single egg she
will be able produce during her lifetime already present (in an immature form) in
her ovaries. This situation is very different from males, whose spermatogonia (the
sperm equivalent to oogonia) do not begin producing spermatocytes (the sperm
equivalent to oocytes) until puberty.

The oocyes remain in meiotic prophase I until the onset of puberty, when a series
of events can lead to egg maturation:
1. The anterior pituitary hormones, FSH and LH, cause some of the follicles to
begin developing and oocyte inside the follicle to finish the first meiotic
division.
2. After completing meiosis I, the oocyte pauses again, this time during
metaphase II.
3. Though several follicles are activated during each cycle, only one will
release an oocyte. The released oocyte will begin traveling through the
oviduct, still arrested in meiosis II.
4. If the oocyte is fertilized by a sperm, it will finish meiosis II and undergo
unequal cytokinesis (cell division) to produce a fertilized egg (an embryo)
and another polar body. (If it is not fertilized, the oocyte degrades without
completing meiosis II.)
The process of oogenesis is illustrated below:
Oogenesis begins when the 2n oogonium undergoes mitosis, producing a primary
oocyte. The primary oocytes arrest in prophase I before birth. After puberty,
meiosis of one oocyte per menstrual cycle continues, resulting in a 1n secondary
oocyte that arrests in metaphase II and a polar body. Upon ovulation and sperm
entry, meiosis is completed and fertilization occurs, resulting in a polar body and a
fertilized egg. Image credit: OpenStax Biology.

One final point: when an oocyte undergoes meiosis, it produces only a single egg
(again, this is different from spermatogenesis, which produces four sperm from
each spermatocyte). The oocyte divides unequally, so that almost all of the
cytoplasm goes into only one daughter cell rather than evenly distributed into both.
The smaller cell is called a polar body, and normally dies.
Hormonal control of oogenesis
Oogenesis is controlled by FSH, LH, estrogen, and progesterone.
 FSH stimulates development of egg cells that develop in structures called
follicles, which are located within the ovaries.
 LH also promotes development and maturation of eggs and induction of
ovulation.
 Estrogen is the reproductive hormone in females that assists in
ovulation and regrowing the lining of the uterus; it is also responsible for the
secondary sexual characteristics of females such as breast development.
 Progesterone assists in endometrial re-growth and inhibition of FSH and
LH release.
These hormones together regulate the ovarian and menstrual cycles. The ovarian
cycle governs the preparation of endocrine tissues and release of eggs, while
the menstrual cycle governs the preparation and maintenance of the uterine lining.
These cycles occur concurrently and are coordinated over a 22–32 day cycle, with
an average length of 28 days:
 The first half of the ovarian cycle is the follicular phase. Slowly rising
levels of FSH and LH 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 estrogens. Estrogen levels increase over the course of
the follicular phase as the follicles continue to develop. In the menstrual
cycle, menstrual flow occurs at the beginning of the follicular phase when
estrogen levels are low (when the follicles are only just beginning to
develop); rising levels of estrogen then cause the endometrium to proliferate
(grow), replacing the blood vessels and glands that deteriorated during the
end of the last cycle.
 Ovulation occurs just prior to the middle of the cycle (approximately day
14), when the high level of estrogen produced by the developing follicles
 causes FSH and especially LH to rise rapidly, then fall. The spike in LH
causes ovulation: the follicle which is most mature ruptures and releases its
egg. 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 cells in the follicle undergo physical changes and produce a
structure called a corpus luteum, which produces estrogen and progesterone.
The progesterone facilitates the regrowth of the uterine lining and inhibits
the release of further FSH and LH. The uterus becomes prepared to accept a
fertilized egg, should fertilization occur. The inhibition of FSH and LH by
progesterone prevents any further eggs and follicles from developing. The
level of estrogen produced by the corpus luteum increases to a steady level
for the next few days; estrogen enhances the effects of progesterone.
 It takes about seven days for an egg to travel through the Fallopian tube
from the ovary to the uterus, and it must be fertilized while in the Fallopian
tube:
 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. The figure below
visually compares the ovarian and uterine cycles as well as the
hormone levels controlling these cycles.
 If a fertilized egg implants in the endometrial lining of the uterine
wall, the embryo produces a hormone called human chorionic
gonadotropin (hCG) that maintains the corpus luteum. The ovary
continues to produce progesterone at high levels, and the menstrual
cycle is arrested for the duration of the pregnancy. Because hCG is
unique to pregnancy, it is the hormone detected by pregnancy tests.
The figure below visually compares the ovarian and uterine cycles as well as the
hormone levels controlling these cycles.
Rising and falling hormone levels result in progression of the ovarian and
menstrual cycles. Image credit: modification of work from OpenStax Biology and
OpenStax Anatomy and Physiology; modification of work by Mikael Häggström)
This video provides a great overview of the human female reproductive system,
emphasizing many of the points described above:
Male Reproductive Anatomy
In the male reproductive system, the scrotum houses the testicles or testes
(singular: testis), which produce sperm and some reproductive hormones.
Sperm become are immobile when kept at body temperature; therefore, the
scrotum and penis are external to the body, as illustrated below, so that a proper
temperature is maintained for motility. In land mammals, the pair of testes must be
suspended outside the body at about 2° C lower than body temperature to produce
viable sperm. Infertility can occur in land mammals when the testes do not descend
through the abdominal cavity during fetal development. Though sperm must be
produced and stored at temperatures lower than body temperature in the testes,
sperm are warmed to body temperature when deposited in the female reproductive
tract. The immediate warming of sperm causes them to experience a burst of
swimming activity, but then they begin to lose motility after several hours at body
temperature.
Diagram of male reproductive organs
Image from OpenStax, CC BY 4.0
Sperm are produced in the seminiferous tubules inside the testes. The sperm cell
production is mediated by two different types of cells: “nursemaid” cells
called Sertoli cells which protect the germ cells and promote their development,
and cells of Leydig which produce high levels of testosterone once the male
reaches adolescence and regulate sperm development.
When the sperm have developed flagella and are nearly mature, they leave the
testicles and enter the epididymis, where sperm mature. During ejaculation, 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.
Semen is a mixture of sperm and spermatic duct secretions and fluids from
accessory glands that contribute most of the semen’s volume. The bulk of the
semen comes from the accessory glands associated with the male reproductive
system. These are the seminal vesicles, the prostate gland, and the bulbourethral
gland, all of which are illustrated above.
 The seminal vesicles are a pair of glands that make thick, yellowish, and
alkaline solution. As sperm are only motile in an alkaline environment, a
basic pH is important to reverse the acidity of the vaginal environment. The
solution also contains mucus, fructose (a source of energy for the sperm
cells), a coagulating enzyme, ascorbic acid (vitamin C), and local-acting
hormones called prostaglandins (may help stimulate smooth muscle
contractions in the uterus). The seminal vesicle glands account for 60
percent of the bulk of semen.
 The prostate gland surrounds the urethra, the connection to the urinary
bladder. It has a series of short ducts that directly connect to the urethra. The
gland is a mixture of smooth muscle and glandular tissue. The muscle
provides much of the force needed for ejaculation to occur. The glandular
tissue makes a thin, milky fluid that contains citrate (stimulates sperm
motility), enzymes, and prostate specific antigen (PSA). PSA is a proteolytic
enzyme that helps to liquefy the ejaculate several minutes after release from
the male. Prostate gland secretions account for about 30 percent of the bulk
of semen.
 The bulbourethral gland releases its secretion prior to the release of the
bulk of the semen. The mucous secretions of this gland help lubricate and
neutralize any acid residue in the urethra left over from urine. This usually
accounts for a couple of drops of fluid in the total ejaculate and may contain
a few sperm. Withdrawal of the penis from the vagina before ejaculation to
prevent pregnancy may not work if sperm are present in the bulbourethral
gland secretions.
This table briefly summarizes the major organs, locations, and functions of
mammalian male reproductive anatomy:

Organ Location Function

Scrotum External Carry and support testes

 Penis External Deliver urine, copulating organ

Produce sperm and male

Testes External hormones

Seminal vesicles Internal Contribute to semen production

Prostate gland Internal Contribute to semen production

Bulbourethral
glands Internal Clean urethra at ejaculation

Male Gametogenesis: Spermatogenesis

Spermatogenesis, illustrated below, occurs in the seminiferous tubules in the
testes. Sperm stem cells (called spermatogonia) are present at birth but are inactive
until puberty, when hormones from the anterior pituitary cause the activation of
these cells and the continuous production of sperm. Sperm production continues
into old age. To produce sperm, a cell called a spermatocyte (a precursor to sperm)
undergoes meiosis to produce four haploid spermatids (immature sperm). Once the
spermatid develops a flagellum, (a tail that allows it to swim), it is called a sperm
cell. Four sperm cells result from each spermatocyte that goes through meiosis.
During spermatogenesis, four sperm result from each primary spermatocyte.
Spermatogenesis begins when the 2n (diploid) spermatogonium undergoes mitosis,
producing more spermatagonia. The spermatogonia undergo meiosis I, producing
haploid (1n) secondary spermatocytes, and meiosis II, producing spermatids.
Differentiation of the spermatids results in mature sperm. Image credit: OpenStax
Biology.
Hormonal Control of Spermatogenesis
The information below was adapted from OpenStax Biology 43.4
Just like oogenesis, spermatogenesis is controlled by FSH, LH. Testosterone also
plays a role in spermatogenesis:
 FSH stimulates spermatogenesis in the testes
 LH stimulates testosterone production
 Testosterone further stimulates spermatogenesis. It is also the hormone
responsible for the secondary sexual characteristics that develop in the male
during adolescence, including a deepening of the voice, the growth of facial,
axillary, and pubic hair, and the beginnings of the sex drive.
While this doesn’t occur in a monthly cycle as in females, the hormones do interact
in a feedback cycle which initiates during puberty: In response to signals from the
hypothalamus that begin at the onset of puberty in males, the pituitary gland
produces FSH. FSH enters the testes to begin facilitating spermatogenesis, which
is the production of sperm cells (gametes) by meiosis. LH, made by the pituitary,
also enters the testes to stimulate the production and release of testosterone into
the blood. Testosterone stimulates spermatogenesis as well as the development of
male secondary sex characteristics that include a deepening of the voice, the
growth of facial, axillary, and pubic hair, and the beginnings of the sex drive.
A negative feedback system occurs in the male when sperm counts get too high
(over about 20 million/ml): rising testosterone levels cause Sertoli cells to release
the hormone inhibin, which acts on the hypothalamus and pituitary gland to inhibit
the release of FSH and LH. The inhibition causes spermatogenesis to slow down
until proper levels are again achieved. Once the sperm levels are reduced, the
Sertoli cells stop releasing inhibin, and the sperm count increases.
This video provides a great overview of the anatomy and function of the human
male reproductive system:
How and when are gametes made?
As you’ve just seen in the two videos the production of sperm and eggs takes place
through the process of meiosis, but there are some big differences between the
processes to make eggs versus sperm:
 When gametes start to form: Egg production begins during embryonic
development (before birth), then is arrested during meiosis until puberty;
sperm production does not begin until puberty
 When gametes finish being made: Egg production is not actually completed
until after fertilization (!), while sperm production is complete prior to
ejaculation
 How many gametes are made from a gamete stem cell: Egg production
results in only a single egg from each egg stem cell; sperm production
results in four sperm from each sperm stem cell.
 Rate of production: Once an individual enters puberty, sperm production is
continuous in a “conveyor belt” process; egg production occurs one-at-a-
time at each menstrual cycle.