Unit 3:

 Chapter 1. Reproductive Physiology

 3.1.1. Importance of reproductive physiological studies

1. To develop maturation under captive condition

Most of the females can’t mature under captive condition. In general for good
farming practice the organism should be bred under captivity.

2. To enhance sperm count and viability

Although male fishes mature under captive condition but the sperm count is reduced.
The sperm viability enhancement will result better hatching.

3. Control of excess and undesirable spawning

Tilapia like fishes spawn continued in rearing units. Because of this the growth of this
will be minimum after maturation period as most of the energy is directed towards
gonadal growth for the better biomass production the excessive spawning must be
controlled. Production, sterile fish production is the ways to control undesirable
spawning.

4. Seasonality of spawning

Fishes spawn only during the particular season. Indian Major carps (IMC) breed only
during the monsoon seasons. The culture practice is a year round operation. So the
seeds must be available throughout the year. To get year round production supply of
matured fish is required.

5. Synchronous spawning

Once the male responds spawning means at the time the female will not spawn and
vice versa occur is normal as well as captive condition. This affects the spawn
quality. Both male and female respond at the same time for better spawning. To get
a better result study will help how to induce synchronous spawning in fishes.

6. The seed quality improvement

In nature the availability seed is poor. The collected natural seeds are not uniform in
size, have mixing of genetical character and may carry harmful pathogens. To
overcome this a quality seed can be produced under hatchery condition for which
reproductive physiological study will help a lot.

3.1.2. Introduction

The success of any species is ultimately determined by the ability of its members to
reproduce successfully withstanding environmental variations and thus to maintain
viable populations. Fishes are surprisingly diverse and hence present a diverse array
of reproductive strategies and functions.
 3.1.3. Types of reproduction

3 types of reproduction are possible

1. Bisexual reproduction
2. Hermaphroditic reproduction
3. Parthenogenetic reproduction
1. Fishes are generally bisexual and sperms and eggs develop in
separate individuals having either the testes or the ovaries. Eg. Carps.
2. The eggs and the sperms develop in the same gonad and self
fertilization takes place. There are 2 types of hermaphroditism
1. Synchronous or simultaneous hermaphroditism refers to
the condition in which both parts of the ovo-testis mature
at the same time. This gives rise to a fish which is
functionally both male and female. (eg. Coral reef fish
Hamlet, Hypoplectrus unicolor- Atlantic ocean. Fish of
this species usually form pair bonds with a single partner.
Each individual spawns alternatively as male and female.
Shortly prior to spawning the fish ready to assume the
male role gives a courtship call and the fish assuming the
female role responds and releases eggs. Following
fertilization of the eggs the spawning pair breaks apart
and takes cover on the reef. In the next spawning event,
the roles of the 2 fish are reversed with the original male
acting as the female and the original female taking over
the role of the male.
2. Successive hermaphroditism in which the fish undergoes
sex reversal. The sex change varies among
hermaphroditic species with protrandrous- males change
sex to become females. Protogynous- females change
sex to become males.
Sequential or successive hermaphrodites are born with
one sex and change sex sometime during the course of
life. An example of protandry is found in the anemone
fishes. The fishes live with anemones in a symbiotic
relationship; the anemone provides the fish with shelter
and protection from predation, and the fish supply the
anemone with food. Groups of fishes will live with one
anemone, and will not switch anemones. Only the two
largest will mate; the largest female and the second
largest, the male. With the female being the largest, she
can produce the most eggs. When the female dies, the
largest male will change sexes and become the female.
The rest of the fish are immature males. Some sea
basses (Serranidae) are protrandric hermaphrodites,
being males at first, females later. Eg. Sparidae and
Serranidae are true hermaphrodies. 
A classic example of protogyny is found in the wrasses
and parrotfishes. The males in these species form
 harems, with one large male sequestering and defending
a group of smaller females. The male enjoys spectacular
reproductive success, as it has many females to mate
with. The females also enjoy a limited reproductive
success, producing as many eggs as they can, all
fertilized by the one male. The male has the advantage
over the females; it has many females producing eggs for
him to fertilize, whereas the females only have
themselves.
For example some damselfishes (family Pomacentridae)
begin life as males and change into females. In some,
females can revert back to males.
Some seabasses (family Serranidae) change from female
to male, and are capable of reverting back to female.
Most wrasses (family Labridae) are born female, grow
into sexually mature females, and have the potential to
transform into functional males later in life. In many of the
wrasses, sex change correlates with social hierarchy and
social behavior: social structure includes a large
dominant male and many smaller, subordinate females.
Removing the male from the group triggers the largest
female to begin transforming into a male.
3. Parthenogenesis is the development of young without fertilization. This
is also called gynogenesis. Males of other species are required for
sexual reproduction where the sperms only stimulate gynogenetic
development without contributing their chromosomes (it does not take
part in heredity). Thereby the resultant young are always female with
no trace of paternal characters. Eg. Poecilia formosa (Guppies).
(Some species are unisexual. In unisexual species, there is no fusion
of sperm and egg. The sperm is necessary to trigger the egg cell to
develop into an adult, but the sperm cell ultimately degenerates and
does not take any part in heredity. The resulting young always are
females, with no trace of paternal characteristics. Thus, unisexual
species are entirely female. They mate with males of related species to
produce female offspring. Poecilia formosa is an example of a
unisexual species. Always female, P formosa mates with
male P mexicana or P latipinna).

3.1.4. Sex determination

Fish, like all other vertebrates, reproduce sexually and in the majority of species, the
eggs and sperm are formed in separate individuals.

3.1.5. Male reproductive system

In the male fish, the reproductive organ consists of a pair of testes. In the
cartilaginous fishes they are usually elongated ovoid structures located anteriorly in
the body cavity by means of a mesorchium.

In most of the bony fish, the testes are whitish or creamy coloured, elongated,
lobulate organs attached to the dorsal body wall by mesochoria. A vas deferns or
sperm duct arises from the posterior region of each elongated testis and leads to the
urinary papilla which is located between the rectum and the urinary ducts.

3.1.6. Testis and spermatogenesis

Spermatozoa or sperm are formed from the germ cells via a series of
developmental changes known collectively as spermatogenesis or the process
by which sperm cells are formed in the testis is called spermatogenesis or
spermiogenesis.

Spermatogenesis involves an initial proliferation of spermatogonia through

repeated mitotic divisions and growth to form primary spermatocytes. Primary
spermatocytes then undergo reduction division (meiosis) to form secondary
spermatocytes. Division of the secondary spermatocytes results in the
production of spermatids, which then undergo metamorphosis to the motile
spermatozoa. (Fig).

3.1.7. Seasonal changes in testes

Seasonal changes are found to appear in teleost fishes at the time of breeding. The
testicular cycle in bony fishes divide into 5 phase or periods.

1. Resting period: After spawning a rest period comes in between Aug to

sept. This period is called quisant period or phase. In this period, the
testes are thin and yellowish in colour and seminiferous tubules are
small in size but filled with spermatogonia.
2. Preparatory period: This occurs between Oct to Dec. The
morphological appearance is almost similar with resting period but in
this phase volume of testes increases. In this phase spermatogonia
starts to divide.
3. Maturing period: This period occurs between Jan to Mar. In this period
volume and weight of the testes will increase. Testes are more
vascular and opaque and spermatogenesis takes place.
Spermatogonia reduce in number and numerous primary and
secondary spermatocytes are formed. Interstitial leydig cells become
very well developed.
4. Spawning period: Normally it occurs between April to June. During this
phase, the testes increase in volume and weight. They are turgid and
become pink colour. Milt oozes out on pressing the abdomen. The
seminiferous tubules increase in diameter and are full with sperms and
various stage of spermatogenesis takes place.
 5. Post spawning period: It occurs between July to Aug. At this period
testes will be flaccid due to excessive discharge of sperm. Weight of
testes also reduces. Germ cells of this phase are present in the form of
spermatogonia.
6. 3.1.8. Female reproductive system

The female ovaries are paired but in some species one of the ovaries may be greatly
reduced in size and become non functional. In the elasmobranchs the ovaries are
located anteriorly in the body cavity, each ovary being suspended from the dorsal
wall of the body cavity by mesenteric tissue known as the mesovarium. Ova are
released into the body cavity from the ovary and then enter the oviduct via a funnel
like structure.

In many teleosts the ovaries are sac like, hollow paired structures that are
continuous with the oviduct. (The wall of the ovary is thick at the time of non
breeding season, but at time of breeding it becomes very thin and highly vascular.
The ovary wall is made up of 3 layers. 1. Peritonium which is outermost layer.

2. Tunica albuginea middle layer

3. Germinal epithelium innermost layer

3.1.9. Ovarian follicle

The ovary consists of germ cells, oogonia and oocytes and their surrounding follicle
cells and supportive tissues. The ovarian follicles arise from the germinal epithelium
as each germ cell becomes surrounded by a layer of follicle cells early in
development.

With growth, the follicle cells multiply and form a continuous layer known as the
granulose cell layer. At the same time, the outer layers of connective tissues become
organized to form a follicular envelope or thecal cell layer. Thus, the developing
oocyte becomes surrounded by 2 major cell layers, the inner granulose layer and the
outer thecal layer. The granulose and thecal cell layers are separated from each
other by a distinct basement membrane. (Fig)

3.1.10. Oogenesis

Oogenesis is the earliest stages of the development of the female germ cells are
similar to those seen in the initial stages of spermatogenesis. The germ cells first
undergo proliferation via a series of mitotic divisions and then they develop to
become primary oocytes. There is a period of growth and a meiotic reduction
division. The final reduction division with expulsion of the second polar body does
not occur until after fertilization, when the egg has been activated by the sperm. 

The growth of the oocytes varies from species to species with oocyte enlargement
being caused largely by the accumulation of yolk. Yolk accumulation occurs during
the developmental period known as vitellogenesis. The oocyte enlargement may be
considerable for eg, the oocytes of salmonids increasing in diameter from 20-50 µm
to 4-5 mm during the course of development.

3.1.11. Seasonal changes in the female reproductive organ

These complete in 5 phases or periods.

1. Resting period: Normally it occurs between Aug to Sept. The ovaries

are small, thin, thread like, translucent, pale or dirty white in colour with
reduced vascularization. At this phase ovary consists of several
number of oogonia and oocytes in the stage of oocyte I and oocyte II.
2. Preparatory period: This occurs between Oct to Dec. At this phase
ovary becomes slightly larger, thicker, opaque and yellowish in colour.
There is an increase in the weight of ovary. At this stage oocyte II and
oocyte IV are found maximum in number.
3. Maturing period: This period occurs between Jan to Mar. at this phase
ovary increase in the volume and weight about 2/3 to 3/4 th of the body
cavity. Vascularization increases and the blood capillaries become well
developed. Immature oocytes are reduced in number and stage V
oocytes make their appearance in large number. Stage VI and VII
oocytes are also present in reduced numbers.
4. Spawning period: Normally it occurs between April to June. During this
phase, the ovaries show a well marked increase in their volume and
weight. Ovary become turgid and yellow in colour with a large number
of translucent eggs. Ovarian wall is almost transparent. Ova may be
extruded by applying a pressure on the abdomen.
5. Post spawning period: It is also known as spent phase. Normally it
occurs between July to Aug. The ovaries are flaccid, shrunk and sac
like reduced in volume with dull colour. The vascular supply in the
ovary is highly reduced. The ovary shows atretic and discharged
follicles along with stage I and II oocytes.
6. 3.1.12. Sex determination

The reproductive systems and patterns of sexual behavior displayed by fish are
extremely diverse. There are also a number of different genetic systems and sex
determination mechanisms.

As with other animals, the genetic material of the fish is located within the cell
nucleus in the form of chromosomes. The number of chromosomes varies from
species to species, but the chromosome number almost invariably remains constant
within a species. In majority of fish the chromosomes occur in pairs- diploid (2N)
condition. One of the chromosomes of each pair derives from the male parent and
the other from the female. But, there are some exceptions. Some fish eg, salmonids
and catastomids may have evolved via tetraploidy, which involved a doubling of the
original diploid number (2N) of chromosomes to the tetraploid (4N) condition. Some
fish have become triploids, but the occurrence of natural populations of triploid fish
appears to be extremely rare.
 What are genes and chromosomes?

Our bodies are made up of millions of cells. Most cells contain a complete set of
genes. Genes act like a set of instructions, controlling our growth and how our
bodies work. They are responsible for many of our characteristics, such as
our eye colour, blood type or height.

Genes are carried on thread-like structures called chromosomes. Usually, we have

46 chromosomes in most cells. We inherit our chromosomes from our parents, 23
from our mother and 23 from our father, so we have two sets of 23 chromosomes, or
23 pairs. Because the chromosomes are made up of genes we, therefore inherit two
copies of most genes, one copy from each parent. This is the reason why we often
have similar characteristics to our parents. The chromosomes, and therefore the
genes, are made up of a chemical substance called DNA.

The chromosomes numbered 1 to 22 look the same in males and females. These
are called the autosomes. Pair number 23 is different in males and females and they
are called the sex chromosomes. There are two kinds of sex chromosome, one
called the X chromosome and one called the Y chromosome. Females normally
have two X chromosomes (XX). A female inherits one X chromosome from her
mother and one X chromosome from her father. Males normally have an X and a Y
chromosome (XY). A male inherits an X chromosome from his mother and a Y
chromosome from his father. The picture below therefore shows the chromosomes
of a male as the last pair of chromosomes are (XY). 

(A chromosome is an organized structure of DNA and protein that is found in cells .

It is a single piece of coiled DNA containing many genes , regulatory elements and
other nucleotide sequences . Chromosomes also contain DNA-bound proteins,
which serve to package the DNA and control its function).

( Chromosomes are long, stringy aggregates of genes that carry heredity

information. They are composed of DNA and proteins and are located within
the nucleus of our cells . Chromosomes determine everything from hair color
and eye color to sex. Whether you are a male or female depends on the presence or
absence of certain chromosomes.

How many chromosomes does a Fish have?

Gold fish have 94 while carp have 104, a crayfish 200 - Chromosome numbers
varies from species to species 

The XY sex chromosomes are different in shape and size from each other unlike
the autosomes , and are termed allosomes .)

There are 2 basic types of chromosomes- autosomes and sex chromosomes. Sex

chromosomes are those that determine the sex of an individual and the sex
chromosomes are often morphologically different in 2 sexes. Autosomes are the
other pairs of chromosomes and these are morphologically the same in males and
females.

3.1.13. Sex chromosomes

The most common system of sex determination in fish as in other vertebrates is
known as the XY system. The individuals that are homogametic (XX) are female,
while heterogametic (XY) individuals are males.

In the second system, the male is homogametic (ZZ) and the female is
heterogametic (WZ). This system is known as the WZ system.

3.1.14. Autosomes

The sex chromosomes contain genes that determine the gender of an individual. The
remaining chromosomes that are not involved in determining the sex of an individual
are called autosomes.

Any chromosome not considered as a sex chromosome , or is not involved

in sex determination. It occurs in pairs in somatic cells and singly in sex
cells ( gametes ). For instance, a human somatic cell will normally contain 23 pairs
of chromosomes (total=46 chromosomes ). Twenty-two (22) of these pairs will be
autosomes, and only one of them will be a pair of sex chromosomes (the X and Y
chromosomes).

3.1.15. Sexual differentiation

A light press on the belly of most ripe fishes will bring the whitish milt or the eggs into
view near the anus. However, fish also use other features to exhibit the sexes.

The characteristics of sexual difference or sexual dimorphism that enable

identification of the sexes are classed as primary and secondary. Primary sexual
characters are those that are concerned actually with the reproductive process.
Testes and their ducts in the male and ovaries and their ducts in the female
constitute primary sexual characters. The primary sexual characters often require
dissection for their discernment which makes the secondary sexual characters often
more useful. Secondary sexual characters are of 2 kinds- Those which have no
primary relationship with reproduction act

- Those which are definitely accessory to spawning.

3.1.16. Outstanding differences are listed below

1. Body shape is an important secondary character. The female is pot

bellied than the male particularly when she is ripe or near ripe for
spawning because of ovaries.
2. Many fishes develop a “nuptial dress” which is particularly clearly
expressed in the salmonids. During the reproductive period the change
is first of all expressed in the colouration which becomes much brighter
and in which colours appear that were not present before. Particularly
in the males, the skeleton of the jaws become bent and large teeth
 appear on them. In the cyprinoids, the nuptial dress is expressed in the
appearance of a brighter colouration and of peculiar tubercles on the
horny epithelium of the head and scales of the body. Very great
changes occur in the colouration of many gobies. (Fig.).
3. Both the paired and the unpaired fins are slightly larger in males than in
females (Fig.).
4. In some fishes, the lower lobe of the caudal fin is greatly extended
Eg. Sword tail.
5. In the males of several species, the anal fin becomes enlarged into an
intromittant, copulatory organ designated as gonopodium.
Eg. Gambusia affinis (Mosquito fish), guppy.
6. The pelvic fins of sharks are modified as claspers or myxopterygia.
7. Colouration also serves sexual distinction which is termed sexual
dichromatism. In general, males are brighter in colour than females.
8. Head characteristics also serve to distinguish the sexes among fishes.
In the chimeras, the male develops a spiny stout, retractile knob, the
frontal clasper on the upper part of the head which has similar
characteristics of forehead brooders.
9. Deep sea angler fish, the male is minute and is parasitic attached to a
larger female (Fig).
10. 3.1.17. Maturation and spawning

Testes and ovaries become mature after undergoing through well marked histo-
morphological alterations and increase in size. This is called maturation and the final
expulsion of sperms or ova from the body into the surrounding water is called
“spawning” followed by fertilization. Each species spawns during a specified period
under favourable conditions. The part of the year during which the gonads attain full
maturity and spawning occurs in the population is called breeding season of the
species. After spawning the residual germ cells undergo a brief rest and enter the
maturation cycle again to produce gametes for the next season.

Teleosts vary in the rhythm of maturation of oocytes. Some species spawn only
once, others twice, while still others may spawn several times during a year.

According to Qasim (1973), many Indian marine fishes appear to be continuous

breeders and have a long spawning period extending upto 7-9 months. Maturation
and spawning period varies in different species and even in the same species
collected from different places. Temperature, Salinity and photo period and duration
are considered important factors controlling maturation of gonads. Most of the Indian
freshwater teleosts spawn after the onset of monsoon. The asian seabass lates
callcarifer spawns during june to october, while the grey mullet mugil sephalus
spawns during mansoon seasons from october to june. 

3.1.18. Ovulation and fertilization

The oocyte mature projects into the lumen of the ovary. The follicle ruptures and the
egg escapes from the aperture. This is called ovulation or it refers to the expulsion
of the mature oocyte from its follicle into the ovarian cavity. The forces causing
rupture of the follicle are not clearly known and the muscle fibres of the ovarian wall
may be responsible for it. Fish ovaries may be of three types: gymnovarian,
secondary gymnovarian and cystovarian. In the first type, the oocytes are released
directly into the coelomic cavity, which then travel through the ostium and enter the
oviduct and are eliminated. In the secondary gymnovarian ovaries, ova are shed into
the coelom and then they go directly into the oviduct. In the third type, the oocytes
are directly deposited into the exterior by traveling through the oviduct. Depending
on a female fish’s species, they may have one of these three types of ovaries.

Gymnovaries are the primitive condition found in lungfishes, sturgeons, and bowfins.
Cystovaries are the condition that characterizes most of the teleosts, where the
ovary lumen has continuity with the oviduct. Secondary gymnovaries are found in
salmonids and a few other teleosts.

(ostium An opening, derived from the Latin word for a door or opening).

Fertilization is usually external in water and sperms and ova are discharged close to
each other in water. Sperms become very active soon after they are released in
water, but survive only for a few minutes during which fertilization takes place.

The released egg is protected by a fairly tough chorion or egg case. Within this the
cytoplasm and yolk are contained by a vitelline membrane. Often one or more oil
globules are present. The chorion has only one opening, the micropyle for the
entrance of sperm. Usually the micropyle closes after entry of one sperm to prevent
multiple fertilization. They are collectively referred to as egg activation. The vitelline
membrane separates from the chorion creating a perivitelline space. Some
elasmobranchs have polyspermy in which a number of spermatozoa may enter the
micropyle but only one fuses with the nucleus of the egg and the rest are being
resorbed and perhaps used as an additional nutrient. After fertilization the chorion
hardens. The egg goes through a process of cleavage and morphogenesis as the
cells divide form layers and then organs

3.1.19. Follicular atresia

All the oocytes developing in the ovary of a fish do not reach full maturity and several
of them degenerates. These are called atretic oocytes or pre-ovulation corpora lutea
and are seen in a section of the ovary during pre-spawning and after spawning
period. A number of oocytes are thus lost from the ovary due to degeneration of the
oocytes at various stages of maturation. This influences the fecundity or the
reproductive potential of the fish. The process of atresia has been seen in a number
of Indian teleosts such as Mystus seenghala, Wallago attu, Channa punctatus, Tor
sp, Clarius batrachus, H. fossilis etc. Both mature and young oocytes becomes
atretic and the follicular cells surrounding them are unable to produce healthy mature
ova.

3.1.20. Causes

Various external and internal factors are believed to be responsible for the
degeneration of oocytes. These are hormones, temperature, photoperiod,
overcrowding, nutritional state of the breeders, physic-chemical nature of water, etc.
Studies show that imbalance of the hormones and steroids causes degeneration of
oocytes. Various environmental factors such as light, temperature, pH, salinity etc
must be favourable for healthy growth of oocytes. Hence, any disturbance in these
factors may cause atresia.

(In the female, gamete development also involves the manufacture and incorporation
of yolk sac into the developing oocytes (vitellogenesis). During the early part of
gonadal development the oogonia becomes associated with a number of pre-
follicular cells and it is from these that the ovarian follicle is derived. In the salmonids
for eg, the oocytes increase in diameter from about 50 µm to 500-1000 µm prior to
the start of vitellogenesis. The incorporation of yolk that occurs during the
vitellogenic phase leads to the diameter of the oocyte increasing to around 5000 µm
(5 mm). thus, there is an enormous increase in the oocyte volume during
vitellogenesis. The majority of this increase is due to the incorporation of
exogenously synthesized yolk into the cytoplasm of the oocyte.

The yolk is derived from vitellogenin, a lipophospho-protein- ca complex with a

molecular weight of approximately 440000. Vitellogenin is synthesized in
the liver and is then released into the blood system. The vitellogenin is transported to
the ovary in the blood from which is set aside by the oocyte. Following the uptake
into the oocyte the vitellogenin is spilt into 2 major components- phosvitin and
lipovitellin which forms the yolk stores).

3.1.21. Modes of reproduction

1. Live bearers/ internal bearers

Most sharks give birth to live young, but some release eggs that hatch later. The
gestation period (the time the embryo spends inside the female, developing) can be
up to almost two years long (the spiny dogfish shark may have the longest shark
gestation period).

Baby sharks (called pups) are born with a full set of teeth and are fully ready to take
care of themselves. They quickly swim away, even from their mothers who might eat
them. Litter size ranges from one or two pups (for a great white shark) to over 100
(for a large blue shark and the whale shark).

i. Ovi-ovoviviparous

This is considered to be the first evolutionary step towards live bearing. In some
sharks and rays after internal fertilization and after a number of weeks on incubation
large few embryos are laid each covered with a horny capsule. Among teleosts, they
lay their embryos immediately after fertilization.

ii. Ovo-viviparous or Aplacental Viviparity

This is best known from squaliform sharks and from the coelacanth (Latimeria). In
these fishes, development is internal. The eggs hatch and the babies develop inside
the female's body but there is no placenta to nourish the pups. The large active
young are born after the yolk sac has been absorbed or having their young
developed inside them without direct maternal nourishment .

The eggs develop in the oviduct of elasmobranchs and the ovary of teleosts. The
gestation period can last from a day or 2 in small tropical teleosts to 1 or 2 years in
sharks. The pups eat any unfertilized eggs and each other (they are oviphagous).
Very few pups in a litter survive until birth due to this form of sibling cannibalism. Eg.
for teleosts are Scorpaneidae, Cottidae, Hexagrammidae.

iii. Viviparous

To be precise the female keeps her developing eggs in the final section of the
oviduct, called the womb or uterus and the babies are fed by a placenta which
transfers nourishment from the mother to the babies (via an umbilical cord which is
connected to the baby shark behind the between the pectoral fins). The placenta
helps transfer of nutrients and oxygen from the mother's bloodstream and transfers
waste products from the baby to the mother for elimination. F ollowing a long
gestation period gives birth to fully developed, extremely independent pups, in other
words, sharks give birth to living offspring. The long gestation period inside the
mother's body effectively protects the young from enemies. The number of pups in a
litter ranges from 2-20 or more.

Eg. Hammerhead sharks (Sphyrinidae), carcharhnidae, Embiotocidae, Poecillidae.

2. oviparous, or "egg-laying"

Lay undeveloped eggs, external fertilization (90% of bony fish), internal fertilization
(some sharks and rays).

In fishes, oviparity is most common. The adult can produce many offspring, which
they broadcast into the plankton column. When the offspring settle out of the
plankton, they may be in totally new environments, allowing for a great area in which
the young may survive. This mode also comes with its disadvantages; when born,
the fish must first go through a larval stage for growth before they transform into the
adult stage. In this larval stage, they must fend for themselves in obtaining food and
avoiding predation. They may not find a suitable environment when they settle out of
the plankton column. The survival of individual eggs is very low, so millions of eggs
must be produced in order for the parent to successfully produce offspring. The other
modes have their advantages, the eggs are much less prone to predation when
carried within the mother, and the young are born fully advanced and ready to deal
with the environment as miniature adults. These advantages come with a price-tag
also; the adult must supply nutrients to its offspring and can only produce a few eggs
at a time. The young are limited to the environment that their parents were in, and if
this environment is deteriorating, they are stuck with it.

T he sharks deposit eggs in the ocean which will hatch later if they are not eaten by
predators. The eggs are not guarded by either parent. Shark eggs (sometimes called
"mermaid's purses") are covered by a tough, leathery membrane. Their shape
ranges from pouch-like to screw-shaped (like the California horn shark and the Port
Jackson shark). Some eggs (like those of cat sharks) have tendrils that attach the
egg to objects on the sea bed. The egg has a yolk that feeds the embryo, very much
like a chicken egg. Oviparous sharks include the Zebra shark, the cat sharks, swell
shark, the necklace carpet shark, some Epaulette sharks, and the Horn shark.

3.1.22. Seasonal regulation of reproduction/ environmental influence

on the reproductive cycle

The general, cues that trigger gamete growth and development appear to be
environmental in nature. These environmental cues are often linked to the annual
cycles of daylength and temperature variations. For eg, Cyprinus carpio and many
other cyprinids, gonadal maturation process is thought to be triggered by a
combination of increasing daylength and the commencement of the rise in water
temperature. The spawning season comes to an end during the summer. It is
thought that the cessation of spawning activity may in part be due to inhibitory effects
of high summer water temperatures on gonadal differentiation and maturation.
Spawning does not however, resume as temperature begins to fall with the approach
of winter. This indicates that temperature is not the only environmental factor
exerting a controlling influence over reproduction. Thus, it is thought that both
photoperiod and temperature are major environmental cues responsible for the
mediation of the reproductive cycle in several species of fish that inhibit temperate
latitudes.

In the tropics, there may be only very minor seasonal changes in photoperiod and
water temperature. Consequently, these 2 environmental factors are unlikely to act
as major cues influencing the reproductive cycle, the timing and duration of the
spawning season. In some tropical species, there does not appear to be any distinct
spawning season and members of the population can be found to be engaged in
reproductive activities at almost any time of the year. In other species, spawning
activities do seem to vary on a seasonal basis with factors such as variations in
rainfall possibly acting as the environmental cues triggering changes in gonadal
development and maturation.

3.1.23. Hormonal control of reproduction

Hormones of pituitary gland play a major role in reproduction. Pituitary release

gonadotropin hormones which control growth of testes and ovary in fishes. Pituitary
also show many effect on behavior indirectly by their participation in the development
of secondary sexual characters in fishes.

But the hormones of the gonads and adenohypophysis of pituitary (anterior pituitary)
play a major role in controlling reproductive processes. Some evidence proves that
neurohypophysial and thyroid hormones are also involved in the regulation of certain
components of reproductive behavior in fishes. Other hormones also affect
reproductive behavior indirectly by their effects on metabolism and growth.

(The primary hormones that regulate sex and reproduction are the sex steroids such
as androgens, estrogens and progesterone which have masculinizing, feminizing
and gestational effects, respectively. These hormones are secreted from the gonads
called target glands which are in turn under the pituitary control. The pituitary gland
manufactures many hormones and their mechanism of action varies considerably.
The gonadotropins are secreted from the anterior pituitary (adenohypophysis) and
affect their target glands- the gonads).

3.1.24. Shellfish

The reproduction of crustaceans also varies. Some change sexes in the course of
their lives; others are even hatched with both sexual organs.

3.1.25. Separate Sexes

The sexes are separate in most crustacea and usually can be differentiated from
each other by secondary sex characters. Chief among these characters are the size
and shape of the body, appendages, or both, and placement of the genital apertures.
For these crustaceans, reproduction functions similarly to many of the fish species.
The female produces eggs in the ovaries that are either released in the water or
attached to the female waiting for fertilization. The male then fertilizes the eggs.
Lobsters perform serial monogamy, meaning the males are engaged with only one
female at a time, but will repeat the fertilization with many females. The female
sheds her outer shell and moves into the male's lair for approximately two weeks.

3.1.26. Hermaphroditism

A hermaphrodite is an animal or plant with both male and female reproductive

organs. In crustaceans such as the Notostraca, or tadpole shrimp, shrimp are
hatched containing both male and female sexual characteristics. The ovaries contain
sperm producing lobes and eggs.
In other species, such as the cirripede, males and hermaphrodites exist
simultaneously. The males, if present, are much smaller than the hermaphrodites
and called dwarfs. They attach themselves to the hermaphrodite to assist in the
fertilization of the eggs.

3.1.27. Sex Change

In some shrimp species, it is common for the shrimp to change sexes as they age.
The Aesop Shrimp (Pandalus montagui) start as males and change to female after
13 months. They are found in the Atlantic Ocean. The Oregon Pill bug
(Gnorimosphaeroma oregonense) is one of the only crustaceans to hatch as females
and after maturity, become males. It is advantageous for males to mate when at a
larger size to fend off competition from other males. The mature males mate with the
less developed females.

3.1.28. A sexual Reproduction

Some branchiopods such as fairy shrimp and water fleas, ostracods such as mussel
shrimp and isopods such as wood lice are parthenogenic, meaning they are asexual
and therefore produce eggs that do not need any fertilization. They hatch on their
own. Marmokrebs are a species of crayfish that are all female and reproduce
asexually. The female creates a clutch of eggs attached to her abdomen with
between 20 and 200 eggs that later hatch without sperm fertilization.

Penaeids are dioecious and the external structures of the genital system are the
major dimorphic features. The male has two pairs of modified abdominal
appendages (Pleopods) on the first and second abdominal segments
(the petasma and appendix masculina) that deliver sperm to the female's external
receptacle (the thelycum) located between the bases of the fifth walking legs
(pereopods). The petasma, appendix masculine and thelycum are located on the
ventral surface.

The petasma is formed by the endopodites of the first pair of pleopods which are
modified as interlocking structures for spermatophore transfer. The appendix
masculina are on the endopodites of the second pair of pleopods and serve to
separate the petasma into two component halves. The thelycum may be "open" or
"closed", depending on the species. "Closed" thelyca are those where the
spermatophore is placed by a male in the groove below the plates while the female
is in the soft exoskeleton stage following molting. The spermatophore is stored for
some time before spawning. "Open" thelyca are not enclosed by plates, and the
spermatophore must be placed on it by a male when the female's exoskeleton is
hard; usually within hours of spawning. The presence of a spermatophore on the
female is evidence that she has successfully mated. Open thelyca are found in some
shrimp species endemic to the Western Hemisphere, such as P. stylirostris and P.
vannamei; while closed thelyca are characteristic of most Asian species, such as P.
monodon, P. chinensis, P. indicus and P. merguiensis.

Internal organs of the male reproductive system include a paired testes, vas

deferens and terminal ampoules for spermatophore storage. The female
reproductive system includes paired (but partially fused) ovaries that extend from the
mid-thorax to the posterior end of the abdomen, and oviducts terminating adjacent to
a single thelycum.

(Dioecious species are those whose individual members can usually produce only

one type of gamete ; each individual organism belonging to a dioecious species is
distinctly female or male (the word deriving from the Greek for "two households").
The majority of animal species (for example, all mammals and most reptiles ) are
dioecious).

Crustaceans produce from eggs, which have been fertilized by sperm in much the
same manner as other animals. The eggs are produced in the ovaries in the female
and passed to the outside through oviducts. The sperms are produced in tubular
testes in the male. After the eggs have been fertilized, they begin development and
then hatch.

The eggs of most crustaceans are carried attached to the female until hatched.
Some females develop brood pouches in which the young are retained for a time. A
nutrient secretion which sustains the young until they are released is produced in
some species having a brood chamber. Penaeid shrimp and a few of the lower
crustacea deposit their eggs in the medium in which they live, in some cases
attaching them to aquatic vegetation.
When the eggs hatch, this can take several days to several weeks depending on the
species, the young larvae are detached. From this point, they are on their own and
must feed, grow, swim and survive. After a series of transformations, the larva
becomes a miniature adult.

3.1.29. Life histories vary from the simple to the complex within the
different groups of Crustacea

The life history of the three species of penaeid shrimps is generally similar, although
the time of spawning varies with each species. Mating takes place in shallow
offshore waters and involves the transfer of spermatophores from the male to the
female shrimp. Actual spawning takes place in deeper offshore waters. The eggs are
released and fertilized externally in the water. During spawning the last 3 pairs of
periopods are held tightly together and are flapped vigorously. The unfertilized eggs
are extruded from the paired ovipores located at the base of the third periopods.
Simultaneously stored sperm are released from the thelycum. Both are ejected with
considerable force where the eggs appear as a greenish, and the sperm as a
whitish, cloud exiting from beneath the females as she swims. The sperm are non-
motile and must come into contact by passive collision with the surface of the egg at
the moment of ejection. It is believed that the vigorous flapping of the female’s
pereiopods creates an intense vortex that increases the likelihood of sperm and egg
coming into contact. A female may lay between one half to one million eggs at a
single spawning. Within 24 hours, the tiny eggs hatch into a microscopic larva known
as a nauplius. The nauplius stage is followed by the protozoeal, mysis, and
postlarval stages (Figure 1). Passage from the nauplius to the postlarval stage takes
several weeks. All of the developmental stages are found in the offshore plankton.
The larvae are capable of little horizontal, directional movement and are unable to
swim independently of the water currents. They are totally reliant upon favorable
currents to transport them to inshore waters. The migration from offshore waters to
coastal bays occurs during the last planktonic stage and shrimp enter estuarine
nursery grounds as postlarvae. Once they move into brackish waters, the postlarvae
abandon their planktonic way of life and become part of the benthic community.

Postlarval and juvenile shrimp occupy the shallow, brackish waters of the Sound
where they feed and grow. Growth of the young is rapid when waters are warm
(above 20.0C). Young shrimp remain in the estuary until they approach maturity.
Adult shrimp migrate offshore to spawn, and the cycle is repeated.

3.1.30. Breeding Biology of Penaeus monodon

The ovary lies dorsal to the gut and extends from the cephalothorax (head and
thorax region) along the entire length of the tail. The determination of ovarian
development reveals the shadow of the ovary in the tail region and is scored from 1
to 5.

Figure 1.2 The view observed by hatchery operators when female broodstock are
graded for ovarian development by torchlight. The wide saddle of ovarian tissue
directly behind the carapace (Stage IV) is indicative of an immediate pre-spawning
female. A female scored as a Stage IV during the day is most likely to spawn that
night.

The intensity of the ovarian shadow is due to the different density of the ovary and
the pigmentation of the egg mass. Although the majority of the ovary is found within
the cephalothorax area, the intense pigmentation of the shell in this region prevents
the visualization of any ovarian outline. In immediate post-spawning females a vague
shadow may be seen which is the area the previously enlarged ovary occupied. In
some instances a dark shadow is seen with intermittent areas of vague shadow
outline. This indicates that a partial spawning has occurred and that only a proportion
of the egg mass was ovulated (spawned).

In penaeid prawns the ovaries are paired, but partially fused in the cephalothoracic
region, and consist of a number of lateral lobes. In an undeveloped state, the ovary
either does not cast any shadow or a thin opaque line is seen along the length of the
tail, and is scored as Stage 1. At this point the ovary is comprised of a connective
tissue capsule surrounding a soft vascular area containing future eggs, called
oogonia, and accessory cells, also called follicle or nurse cells. The internal wall of
the ovary capsule is lined with epithelial cells (called the germinal epithelium)
containing oogonia. Once the female is sexually mature, the germinal epithelium will
produce oogonia by mitosis division throughout the reproductive life of the female.
The eggs develop from oogonia in an area known as the zone of proliferation. As the
oogonia develop they increase in size and enter the first stage of meiotic division and
henceforth are irreversibly destined to become haploid, with only one set of maternal
chromosomes. At this point, although the developing eggs are increasing in size they
are not as yet producing yolk, and are known as previtellogenic oocytes. At this
stage the ovary can be visualized with a light beam as a large centrally located
opaque rope-like structure, and classified as Stage 2.

Typically, it is at Stage 2 that inhibitory hormones in the eyestalk, arising from what is
called the X-organ-sinus-gland (XO-SG) neurosecretory complex, prevent further
ovarian growth, especially if nutritional or environmental conditions – such as those
while females are held in captivity in tanks – are deemed unfavourable by the
female.

This block on ovarian development can be removed by eyestalk ablation. However,

since the prawn is bilaterally symmetric, ie with two eyes, eyestalk ablation
only results in the removal of one of the two XO-SG complexes producing an
inhibitory substance. Nevertheless, in most cases this is sufficient to induce further
ovarian development and spawning, albeit under sub-optimal conditions.

As the oocytes develop further they migrate out towards the margins of the ovarian
lobes in preparation for ovulation. During this migration, follicle cells are attached to
the periphery of each oocyte. It is believed that the follicle cells produce the yolk that
is internalised in the oocytes in a process called vitellogenesis. As vitellogenesis
proceeds, oocytes mature synchronously as yolk accumulates and develop a
characteristic dark green colour as a result of deposition of carotenoid pigments. It is
the carotenoid pigmentation that mainly causes the dark ovarian shadow during
illumination of the female by torchlight. The female is now in Stage 3. By the end of
vitellogenesis, the eggs develop cortical granules filled with a jelly-like substance
destined to form part of the egg shell membrane after ovulation. At this time the
shadow cast by the ovary is large, resulting in a very distinct dark thick region
extending the length of the abdomen, with an enlarged bulbous region directly
behind the carapace, called the saddle. The saddle may not be as apparent in some
broodstock, such as those that have made several spawnings after eyestalk ablation
or in captive reared broodstock. The female is now in a pre-spawning state and is
scored as a Stage 4.

The culmination of ovarian development is marked by release of the fully mature

oocytes (eggs) into the oviduct at ovulation. Ovulation is typically followed within a
few minutes by oviposition, ie the release of eggs and stored sperm into the water –
the actual spawning event itself (see sections on mating and spawning). Within the
egg the nuclear membrane disappears in readiness for fusion between the two
(haploid) pronuclei, one in the egg originating from the female and the second, which
comes from the sperm at the time of fertilization.

The complete ovary extends from the head to the tail. The majority of the ovarian
mass is within the cephalothorax region which cannot be observed by torchlight.

A female can spawn several times within a single molt cycle. A new population of
primary oogonia is recruited for each spawning event. If the female does spawn
repeatedly within a molt cycle the same reserve of sperm within the thelycum, from
the initial mating after molt, is utilized to fertilise the eggs. Typically, successive
spawnings have progressively decreasing fertility rates as the sperm mass is
exhausted. If the female does not spawn within a molt cycle, the developed ovarian
mass is reabsorbed a day or two before the next molt. After each molt, the ovary is
fully regressed and the ovary must mature a new.

3.1.31. Decapod Crabs - Reproduction and Development

Decapod crabs are built like a folded-up lobster. The abdomen, which is equivalent
to a tail, is folded up tightly underneath its body to form an abdominal flap.

When you turn a crab over and look underneath, you can tell a crab's sex by looking
at its abdominal flap. A male crab has a small triangular flap, while a female crab
has a broad oval-shaped abdominal flap.

When a male and female crab mate, many female decapod crabs can store the male
sperm until her eggs are ready to be released. When the eggs are released, the
stored sperm flows over them and they become fertilised.

(The reproduction habits of crabs are interesting as well. The male delivers his
sperm to the female for egg fertilization via the gonophores located on the legs. They
then find a comfortable place for the female to lay her brood and watch over them
until they hatch. The first two sets of appendages are used for sperm transfer and
therefore the gonophores are located more towards the midline of the body.)

The female crab holds the fertilised eggs in a big spongy mass between its
abdominal flap and the body. The eggs are cemented to the pleopods, which are
small legs, creating the "berried" appearance.
To keep the eggs healthy, the female crab continually "waves" water over the eggs
with the pleopods. When the eggs hatch into zoea larvae, they drift away in the
ocean currents as plankton. As the juvenile crab grows in size, it goes through a
series of moults, each larval stage changing form and function as it grows in a
process called metamorphosis.

At each moult more segments are added to the end (posterior), and the feathered
limbs are replaced by the clawed limbs. The megalops stage more closely
resembles the adult decapod crab

Molluscs

Mollusks are primarily of separate sexes and the reproductive organs (gonads) are

simple. Reproduction via an unfertilized gamete (parthenogenesis) is also found
among gastropods of the subclass Prosobranchia. Most reproduction, however, is by
sexual means. Eggs and sperm are released into the water by members of some
(primitive) species, and fertilization occurs there. In prosobranch gastropods, water
currents may cause a simple internal fertilization within the mantle cavity, or males
may fertilize eggs internally using a muscular penis. Both male and female
reproductive organs may be present in one individual (hermaphroditism) in some
species, and various groups’ exhibit different adaptations to this body form. For
example, in hermaphroditic bivalves and prosobranch gastropods, male and female
gonads are functional at separate times and in rhythmic and consecutive patterns
(successive hermaphroditism). Conversely, male and female gonads are functional
at the same time (simultaneous hermaphroditism) in solenogasters and many other
gastropods.

Fertilization by transfer of capsules containing sperm (spermatophores) typically

occurs in cephalopods and some gastropods. In cephalopods, transfer of
spermatophores is usually combined with copulation by a modified arm,
or hectocotylus. Copulation in solenogasters, often by means of a special genital
cone, may be supported by copulatory stylets. Various penis formations, in part with
copulatory stylets, or darts, are widely found in gastropods.

Eggs are deposited singly or in groups, generally on some hard surface and often
within jelly masses or leathery capsules. Squids of the suborder Oegopsida and
some gastropods have eggs that are suspended in the water. Fertilized eggs
commonly undergo spiral cleavage, as in annelids and a number of other
“protostome” phyla. The eggs of cephalopods, on the other hand, possess a large
amount of yolk, which displaces the dividing cells and causes a characteristic type of
development.

Many mollusks develop into free-swimming larvae; these larvae are either feeding
(planktotrophic) or nonfeeding (lecithotrophic). The larva in primitive bivalves is
a pericalymma (test cell) larva in which the embryo is protected below a covering
(test) of cells provided with one to four girdles of cilia, at the apex of which is a
sensory plate of ciliated cells. After the developing juvenile has grown out apically of
the test (which then is lost), the animal settles and develops into an adult. The test in
other lecithotrophic larvae is restricted to a preoral girdle of ciliated cells (the
prototrochus) and is called the trochophore larva. Trochophores are encountered in
the development of many marine annelid species (phylum Annelida). In more
advanced mollusks (such as in marine gastropods and bivalves), the trochophore
larva develops into a veliger larva. In these generally planktotrophic larvae, the girdle
of ciliated cells widens to form a velum that entraps food and also propels the
microscopic mollusk through the water. As the larva continues to develop, the shell,
mantle cavity, tentacles, and foot appear. After a specific amount of time, which
varies according to species and environmental conditions, the larva loses the velum
and metamorphoses into an adult. A substantial change in shell morphology usually
marks the transition to adult form.

Secondary (newly evolved) larvae have developed among some freshwater bivalves
and some cephalopods. Maternal protection of the developing eggs (brood) is not
unexceptional behaviour in solenogasters, bivalves, and certain gastropod adults.
Direct development without a larval stage or the bearing of live young from a yolky
egg, or both, are typical in cephalopods and most nonmarine (and many marine)
gastropods. Many species go through two breeding seasons per year, whereas in
some cephalopod species mating or egg laying appears to be rapidly followed by
death effected by hormones.

3.1.32. CLASS BIVALVIA (PELECYPODA) - (Bi-valv i-a) (pely-cy-poda)

The majority of bivalves are dioecious (two sexes). Their two gonads are very closely

situated next to each other and they encompass the intestinal loops. The gonoducts
are very simple as there is no copulation amongst bivalves.

In the Protobranchs and Filibranchs, the gonoducts opens directly into the nephridia
and provide for the exit of sperm and eggs.

In the Eumellibranchs, the gonoducts opens directly into the mantle cavity very close
to the nephridiopore.

A few bivalves such as the Cockles (Cardiidae), Poromyidae, a few of the oysters

and scallops (Pectinidae), some of the fresh water
clams Sphaeriidae and Unionidae are hermaphroditic (one sex).

In most of the bivalves, sperm and eggs are released into the surrounding water
where fertilization occurs. The eggs and sperm, which were deposited into the
suprabrachial chamber, are swept out along with the exhalent current.

In a few of the bivalves, such as the common oyster Ostrea edulis L., fertilization

occurs within the suprabranchial chamber itself when sperm is drawn in along with
the inhalent current. The fertilized eggs then develop in the gill filaments.

In some of the freshwater hermaphrodites, self - fertilization may actually occur in the
genital ducts before the eggs are deposited into the suprabranchial chamber. The
eggs then travel into the water tubes of the gill and there they develop into larvae.

3.1.33. CLASS CEPHALOPODA - (ceph-a-lo-poda)

Cephalopods usually are usually dioecious (two sexes), and fertilization is internal.
Courtship, which in some species can be quite elaborate, is often a precursor to
copulation. Gonads are located in the posterior of the body.

In some species, males can be distinguished by the modified sucker discs found at
the tips of his longer two tentacles. The male uses these long arms to remove a
sperm packet from his mantle cavity and to then insert this packet into the female’s
mantle cavity. This arm is autonomous and sometimes can get broken off while
mating and is retained within the female’s cavity. If this happens, the male simply
grows a new one: no problem!!

Within two months of mating, the female octopus will attach long strands of clustered
eggs (resembling a cluster of grapes) to the ceiling of her lair. While the eggs are
incubating, the female will gently caress them to keep them clean and free of
bacteria. She also keeps a steady flow of fresh, oxygenated water flowing over her
precious eggs. When they are ready to hatch, her caresses become more rigorous,
this helps the young to escape from their egg sacs.

The pelagic Cephalopod family Argonautidae, commonly knows as the paper

nautilus, have a remarkable adaptation for egg deposition. The two dorsal arms of
the female are greatly expanded at their tip to form a membrane. The expanded
portion of each arm secretes one half of a beautiful calcareous bivalved shell. She
then deposits her eggs directly into this case. The shell acts as both a brood
chamber and a retreat for the female. The posterior of the female usually remains in
this shell. The male, which is much smaller than the female, does not have such a
shell and is often found in cohabitation within the same shell, more or less as a
freeloader.

The gonads, ovaries and testes, of all vertebrates are similar in structure and
function. The gonads are usually paired (although they may be fused or single in
some groups). The testes in all vertebrates produce millions of sperm at a time. As
we shall see, the major differences in male reproductive systems involve the
mechanisms for transferring those millions of sperm to the ova.

Ovaries produce ova surrounded by fluid filled sacs called follicles. These follicles
burst releasing ova into the coelom and then the oviduct. The number of ova
produced at any one time varies considerably depending upon the habitat of the
animal and the amount of parental care the young will receive. Thus fish, like the
cod, which release unprotected eggs into the ocean and do not care for their young
may produce and release more than a million eggs at once. Other fish and
amphibians which do not broadcast their ova as far afield or which give some
measure of parental protection to the eggs may produce thousands of eggs at once.
Birds and mammals, which invest considerable parental energy in protecting their
offspring both before and after birth, produce only a few eggs (ova) at one time.

Once ova are released from the ovary, they are transported through the oviduct
toward the exterior. In the case of fish and amphibians, which release shell-less
eggs, the oviduct is essentially an unspecialized tube leading indirectly to the
exterior. Birds and reptiles produce shelled eggs; the shell is critical in protecting the
egg from desiccation in the terrestrial environment. The oviducts in these animals
are modified, being in close association with a shell-forming gland and capable of
considerable distension to accommodate the enormous, inflexible eggs that result.

3.1.34. Embryonic Development in Fish

Embryonic development starts with eggs

The initial stages of embryonic development are almost identical for both lower and
higher level invertebrates. Fish embryonic development consists of seven stages
leading to hatching. These stages are the zygote period, cleavage period, blastula
period, gastrula period, segmentation period, pharyngula period, and finally hatching.

Inception

Fish embryonic development begins after inception with the zygote period. During
this time the embryo develops one cell that looks like a half bubble. This lasts on
average for 45 minutes with a cell .7 mm in diameter. The chorion, also known as the
egg shell, will swell and lift away from the fertilized egg.

Cleavage Period

The next stage of embryonic development allows the fish to rapidly accumulate new
cells. As cleavage occurs, the cells will divide at 15-minute intervals. Depending on
the complexity of the fish, this cycle will last 45 minutes to 2 1/2 hours.

Gastrula and Pharyngula Periods

Gastrula period will last anywhere from 5 1/4 to 10 hours. During this stage organs
will begin to become visible, and the developing tail becomes more prominent. The
embryonic structure also elongates. Next the pharyngula period occurs, where the
fish will begin to exhibit fully formed organs. This can last anywhere from 24 to 48
hours. It is during this period that the distinction between upper and lower vertebrate
embryonic development can be made.

Hatching

During the next 48 to 72 hours the external indicators of the fish develop. Gills, jaw
and pectoral fins grow at an accelerated rate. Once development is complete the tiny
fish is ready to hatch. Over the next several days the fish larvae will emerge from
their eggs at various rates, ready to hide and grow in their new environment.

3.1.35. Hormones involved in reproduction and their sources

transaction of external signal

The External environment where the fish is living plays vital role in inducing the
animal to go for maturation. Photoperiod (the duration / or length of day time) plays
important role in maturation of animals. The pineal organ through the photoreceptor
cells and other environmental culs produce electrical pulse which includes the
production of melatonin from Nacetyl serotomin in the presence of hydroxyindole–O-
methyl transferare an enzyme. This organ in fishes mediates photoperiod control of
gonadal activity, which involves the pineal hormone melat and the regulation of
diurnal cycle of gonada release. Illustration

Spring spawner normally breed during the lengthier photoperiod where as the winter
spawner breed during only shorter photoperiod.

Mechanism involved in maturation and ovulation

Egg development and Endocrine regulation

Steroids in the male :

 steroid production sites

 intestitial cells in testis – leydig cells
 sertoli cells or cyst cells
 seminal vescicles
 spermatogonia
Produced steroids

 Androgen
 Progestins
 Steroid conjugates
Antrogen

Testosterone, 11 oxyhydrogenated androgen 11 keto testosterone

1. Testosterone increase spermatogenesis after spermination amount

decrease
2. 11 Ketotestosterone increase spermiation
3. Secondary serval character regulation
4. forms pheromones – glucuromide
Progestins

Produced in the testis 17 µ hysony 20 ß dihydro progesterons – peak occur after

fertilization and before the onset of spermiation. It involves in spermiation process by
conrolling ionic composition in the seminial fluid and involves in spawn behaviour of
male and female.

Steroid conjugates

 Glucuronides
 Maximum during breeding season and found during sperroiogenesis
Glucuronidation is the process to eliminate steroids by converting into water soluble
and thus more readily excretable products function as sex pheromones.

Endocrine regulation of spermatogenesis and secondary sexual

characteristics

Gonadal regression
Attesia of garmetes occur if the normal levels of hormones are disturbed. Altesia is
involved in the removal of unused gametes after the spawning season; any may be
related to the change in hormone level after spawning.

Factors inhabiting reporduction