Fish Reproduction and

Development

Coevolved traits for producing another

generation that will produce another
generation...and another...and another...
 Reproduction

Reproduction is the biological process by which new

"offspring" or individual organisms are produced from
their "parents". Reproduction is a fundamental feature of
all known life; each individual organism exists as the
result of reproduction

The known methods of reproduction are broadly grouped

into two main types:

a) Sexual (Bisexual, Hermaphrodite)

b) Asexual (Parthenogenetic)
 Type of Reproduction
1) Bisexual or heterosexual- most of the fish (Gonochoristic (single sex, fixed
at maturity)
2) Hermaphrodite- Perca, Stezostedon, Micropterus

Simultaneous hermaphrodites function as male and female at same time (23

families; ex. Anguilliformes, eels; Atheriniformes, killifish)

Sequential hermaphrodites start life as one sex, change sex after maturity
Protandrous: male first, female later
Protogynous: female first, male later (most common, Wrasses)
n

3) Parthenogenetic (asexual)- (Amazon molly Poecilia formosa)

n

In parthenogenetic reproduction, unfertilized eggs develop

into embryos.
Gynogenetic – sperm needed for egg development, but mating without
fertilization, result is daughters are genetic clones of mothers
Hybridogenetic - egg development with fertilization by males of other
species, but male genes discarded at next generation
 Reproduction Strategies
Based on reproductive strategies adopted by different fishes, they are categorised into
three groups:

1. Non guarders
2. Guarders
3. Bearers

Non Guarders (they do not guard their egg and larvae) are of two types

i) Open substrate spawners (lay eggs in open places)

ii) Brood Hiders (lay eggs in hidden places)

Guarders (Male or Female or both the parents guard the eggs and larvae after laying)

i) Substrate choosers
ii) Nest spawners

Bearers

i) External Bearers
ii) Internal Bearers
 Mating System
– Promiscuous - both sexes with multiple partners -
most common
– Polygamy-Either male or female will have
multiple partners in a given breeding season
– Polygynous - males with multiple mates (cichlids)
– Polyandry - females with multiple mates – few
(Anglerfish, males “parasitize” females
– Monogamy - mating pair remains together over
time, long gestation of young (some cichlids,
seahorses, pipefish)
 Method of fertilization
– most fishes use external fertilization
• less time and energy in courtship, pair bonding
• increases number of potential mates
• greater fecundity

– internal fertilization in few groups:

• sharks, rays, skates, ratfishes (Chondrichthyes)
• guppies, mollies, etc. - Poeciliidae, Goodeidae
• surfperches - Embiotocidae
 Method of fertilization cont……
– internal fertilization requires
• lengthy courtship, preparation for mating
• intromittent organ
– claspers (pelvic fins) in Chondrichthyes
– modified anal fin in poeciliids, goodeids
– modified genital papilla in embiotocids
• male structure for storing sperm (seminal vesicle)

– buccal fertilization
Sperm swallowing (eg. Callichthyid catfish (Corydoras)
 Reproduction Methods
1) Oviparous- egg layerer
Producing eggs that develop outside the maternal body (many bony fishes)

2) Live bearer
a) Viviparous
fertilization is internal and during gestation, there is maternal-embryonic
transfer of nutrients. Anablepidae, Jenynsiidae, Goodeidae, Poeciliidae
b) Ovoviviparous
Fertilization is internal, but during gestation there is no maternal-embryonic
exchange of nutrients and developing embryo sustain on yolk reserve of
oocyte. Scorpaenidae, Cottidae, Hexagrammidae
In some ovoviviparous fishes the embryo develops in the egg while the egg is still
within its follicular covering within the ovary, and ovulation (or release of the egg)
and birth occur at the same time. In other ovoviviparous forms the eggs are released
from the protective follicles into the cavity of the hollow ovary, where development
continues. In some viviparous fishes the walls of the egg follicle are in intimate
contact with the embryo, supplying it with nourishment. In the viviparous sharks, a
part of the oviduct, or egg channel, is developed into a uterus, where the modified
yolk sacs of the young are closely joined to pockets within the uterus.
 Sexual Dimorphism
Most of the fishes exhibit sexual dimorphism or secondary sexual characters by which
sex can be distinguished from each other. In few fishes secondary sexual characters
are discernible throughout the life span and show some structural differences like
1. Show morphological peculiarities which facilitate fertilization of ova , as copulatory
organ in male.
2. Structural peculiarities that are not related to fertilization but are meant for
courtship and fighting with other males.

While in some fishes secondary sexual characters are discernible during breeding
season and external morphological differences pertain to the following features
1. Size of fish
2. Length/shape/texture of fins
3. Coloration
4. Genital papilla
5. Ovipositor
6. Shape of mouth
Function of secondary sexual characters
a) Recognition of opposite sex, b) helping in the act of copulation such
as sexual embrace, c) transfer of spermatozoa from male to female,
d) facilitating parental care
 Reproductive morphology

• Cartilaginous fishes:
– male: testes -> Leydig’s gland -> seminal vesicle ->
cloaca -> claspers
– female: ovary -> ostium tubae -> oviduct -> shell
gland -> [uterus] -> cloaca

• Most Bony Fishes:

– male: testes -> vas deferens -> urogenital pore
– female: ovary -> oviduct -> urogenital pore
 Reproductive organ

The fishes have one pair of bilateral gonad. Generally

symmetrical.

Suspended from dorsal portion of body cavity by mesenteries

in close association of kidneys. Mesentery is richly supported
with blood vessels and nerve fibres.

In male mesentery is called mesorchium

In female mesentery is called mesovarium

 Male Reproductive Organ
Male reproductive organ Composed of
i) Testis
It is paired, elongated, flattened structure present on either side of
ventral to kidney in the abdominal cavity. Attached to the body wall by
means mesorchium

ii) Sperm duct or vas deferens

iii) Genital pore or cloaca

 Testis
In most cases, testes are a pair of elongated structures
composed of numerous seminiferous tubules embedded in the
stroma.

The testis consists of thin-walled tubules or lobules that contain

germ cells - the spermatogonia - which are endodermal in origin.
Germ cells divide in clusters enclosed by a cyst.

Primary spermatogonia - which are present throughout the year,

divide mitotically to give rise to secondary spermatogonia which
get transformed into primary spermatocytes.

They divide by meiosis and give rise to spermatids from which

spermatozoa are formed.

The seminiferous tubules are packed with spermatozoa in the

pre-spawning and spawning periods,
 Development of sperm
or spermatozoa from
primary germ cell or
spermatogonium is
called spermatogenesis

Spermatids undergo
metamorphosis to
produce sperm and this
process is called
spermiogenesis
 Structure of sperm

a) Head piece

b) Middle piece

c) Tail

In fish sperm, acrosomes are absent

 Reproductive cycle (Testicular)
Most of the fishes breed in one season every year (seasonal spawners) and some
breed through out the year (year round spanwers). Seasonal breeders exhibit
rhythmic changes in the structure and physiology of ovary and testes in different
seasons. These changes in testis are demarcated into five phases

1. Resting phase: Testis remain in immature state, Seminiferous tubules are solid
being filled with spermatogonial cells
2. Preparatory phase: During this phase cell proliferation and 1st and 2nd meiotic
division occur as a result primary spermatocytes, secondary spematocytes and
spermatids are produced (spermatogenesis).
3. Mature Phase: during this spermatids undergo further development and
metamorphosis and develop into mature spermatozoa or sperm. The preocces of
development of spermatozoa from spermatids is called spermiogenesis.
4. Spermiation Phase: During courtship and mating process, male eject milt
(spermatozoa in seminal fluid) out of its body through genital aperture to fertilized the
eggs released by female. The seminal fluid is mostly secreted by the cells lining the
vas deferens (sperm duct) and provide nourishment to the mature sperm. Both
spermatogenesis and seminal fluid secretion are under control of gonadotropin of
pituitary gland and male hormone testosterone.
5. Post spermiation phase: during this phase testis is characterized by the presence
of evacuated seminiferous tubules.
Maturity stages of testes
 Female Reproductive Organ
Female reproductive organ Composed of
i) Ovaries
It is paired, elongated sac like structure present on either side of
ventral to kidney in the abdominal cavity. Attached to the body wall
by means mesovarium

ii) Oviduct
iii) Genital pore or cloaca
Types of ovary:
1.Cystoarian: mature oocytes released in to
ovoceal that is continued as oviduct to open into
exterior (Lepidosteus and Teleost fishes)

2. Gymnoarian: mature oocytes released into

body or coelomic cavity from where they passout
through genital aperature (Chondrichthyes, Dipnoi,
Chondrostei and Amia)
 Ovary

The ovary in most teleost fishes is a hollow sac-like organ into

which extend numerous ovigerous folds (lamellae) lined by
germinal epithelium.

The germ cells, oogonia (originate from germinal epithelium)

multiply mitotically and get transformed into non-yolky primary
oocytes

Primary oocytes, covered generally by two layers of follicle cells,

an outer thecal layer and an inner granulosa layer, undergo
vitellogenesis when yolk is deposited in the ooplasm.

Analysis of yolky eggs indicates that the egg-yolk is composed of

lipovitellin and phosvitin, the former is rich in lipid and poor in
phosphorus and the latter is rich in phosphorus and poor in lipid
Process of development of
mature egg from
primordial germ cells
called oogenesis
Stages in the maturation of oocytes
 Reproductive Cycle (Ovarian)
Parallel to testicular cycle, the primary reproductive organ of female i.e. ovary also
undergoes rhythmic change which is divided into five phases as follows:

1. Resting Phase: the ovary is in immature state containing nests of oogonial cells
stage 1 oocytes at different phases of growth and few stage II oocytes. During this
stage the first meiotic division of nucleus is initiated and the same is arrested at
pachytene stage.
2. Preparatory phase: oocytes are in stage III. There is beginning of vitellogenesis
in the oocyte cytoplasm. Oocyte envelops get differentiated with the thecal and
granulosa layers assuming the steroidogenic function. Cells of granulosa layer start
producing female hormone, estradiol.
3. Maturing or Prespawning phase: Characterized by intensive process of
vitellogenesis by which ooplasm of an oocyte is loaded with yolk granules and in stage
IV. 1st meiotic division completed and second meiotic division occures. The size
of the ovary and GSI will be maximum at this phase.
4. Spawning Phase: characterized by gravid ovary containing ripe oocyts. During
spawning follicle of fully ripe oocytes rupture as a result oocytes released into ovocoel
(cystovarian ovary) or body cavity (gymnovarian ovary) from where they pass out
through genital pore into water.
5. Post spawning phase: the ovary exhibits a collapsed appearance as evaculated
follicles are seen after the release of eggs.
Seasonal Changes in the ovary of teleost
 Vitellogenesis

Accumulation of yolk substances in eggs called vitellogenesis

At the beginning there is no yolk. Blood of mature female contain a
calcium binding lipophospoprotein called vitellogenin which is
synthesized in the liver under the stimulation of female sex
steroids. It passes from the plasma to oocyte and undergoes
limited proteolysis give rise to yolk protein lipovitellin (Lv),
phosvitin (Pv), β’-component (β’-c), and C-terminal peptide.

Yolk protein deposited as three yolk substances:

Yolk vesicles -contain glycoprotein

yolk globules-lipoprotein
oil droplets- Glycerides

After completion of vitellogenesis, movement of germinal vesicles to

animal pole, Fusion of yolk granules and grouping of oil droplets
occurs
Vitellogenesis
 Gonadosomatic Index (GSI)

The Gonatosomatic index of a species can be calculated by

the following formula

GSI= Weight of the gonads/weight of the fishX100

Gonadosomatic index of a species has been widely used to

indicate the maturity and periodicity of the spawning of the
fish. The GSI increases with the maturation of the fish and is
maximum during the peak period of maturity. It decrease
abruptly after spawning.
 Fecundity
Fecundity is a measure of the reproductive capacity of a female fish and can be
defined as the number of ova that are likely to be laid by a fish during the
spawning season. It varies from species to species and different individual of same
species may also exhibit variation depending upon size, age, nutritional status,
environmental factors and genetic composition.
The fecundity of a species can be measured by any one of the following

(i) Volumetric Method: The mature ovaries are taken out of the abdomen and total
volume is determined. Now small pieces of the ovary are taken as random
samples from anterior, middle and posterior parts. The volume of each sample is
determined and number of ova in each is counted under a lens. The total number
of ova in the total volume of the ovaries is then calculated.

(ii) Gravimetric Method: mature ovaries are preserved in 10% formalin for estimating
fecundity. Weight of the ovaries is determined. And three samples of 100 mg each
are taken at random from anterior, middle and posterior parts. The the number of
ova in each sample are counted under a binocular microscope. Tthe total volume
of the ova are calculated as follows

Fecundity=average number of ova from three samples of 100 mg each X total

weight of ovary/100
 Hormonal
Control of
Reproduction
 Embryonic Development
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.
Zygote period: Soon after fertilization, the micropyle is closed thereby preventing
the entry of any other sperm into oocyte. The perivetelline space gets filled with
water and colloid, swelling of oocytes takes place. This phenomenon is called
water hardening. During water hardening the animal pole develops a small hillock
of protoplasm on yolk mass known as egg blastodisc.
Cleavage period: Cleavage furrow is limited to cytoplasm of animal and such
type of cleavage is called meroblastic cleavage. As successive cleavage continue,
a group of cells are formed and it is called morula.
Blastula period: After further division the cell arrange in to the from of a layer
called blastoderm. Each cell of blastoderm is called blastomere. Cleavage also
result in the formation of another type of cells called periblast which lie between
blastoderm and yolk. The embryo develops from the blastoderm. A space develop
between the blastoderm and periblast which is called segmental cavity or
blastocoel. At this stage embryo is called blastula. At blastula stage presumptive
germ layers (ectoderm, endoderm and mesoderm) can be noted.
 Embryonic Development continued…….
Gastrula period: The process of formation of gastrula is called gastrulation. At this
time the under rim of blastodisc thickens to form a merginal ridge and inner layer of
this is called germ ring. The germ ring is thickest at the caudal end, which is called
embryonic shield. Gastrulation occurs by three process invagination, involution and
epiboly. An invagination appears at the future endodermal region of blastula . The
portion that is invaginated grow below the blastoderm and this portion is called
hypoblast. The cells of endoderm and mesoderm multiply and migrate inward over
the lip of blastopore and this process is called involution. During this process the
cells of the blastoderm also continue to grow over the yolk which is called epiboly.
As a result yolk mass is covered by a layer of cells called epiblast and future
ectodermal cells are involve in this process. The periblast also grows over the yolk
forming the inner layer covering the yolk below epiblast. Inner priblast and outer
epiblast layer enclose the yolk which is called yolk sac. With the formation of yolk
sac gastrulation is completed.

Segmentation period: The primary organ rudiments appear in the anterior part of
the embryo. Anterior part of the neural plate from where the brain develops appears
first. Notochord gets separated from mesoderm. Mesoderm laying on the both side
of notochord gets segmented and various organs of body are formed. This result in
a small embryo with cylindrical and bilaterally symmetrical body.
 Embryonic Development continued…….

Pharyngula period: In this stage embryo will begin to exhibit fully formed
organs Ectoderm give rise to epidermis and its derivative like enamel of
teeth, olfactory epithilium, lens of eye, inner ear, brain, spinal cord, retina
and optic nerve. Mesoderm divide into three parts the dorsal epimere,
middle mesomere and ventral hypomere. Epimere becomes divided in to
somites, each somites having three parts such as outer dermatome,
middle myotome and inner sclerotome. Dermatome give rise to
connective tissue and muscles of dermis of skin, myotome give rise to
muscle of trunk, appendicular skeleton, fins and their muscles. Vertebral
column is derived from sclerotome. Kidney, gonads and their ducts are
derivatives of mesomere. Hypomere develops to somatic and splanchnic
layers of mesoderm and enclose coelomic cavity. Mesenchyme cells
derived from splanchnic layer develops to involuntary muscles and
connective tissue of gut, head and blood vessels. Mesenchyme of head
region gives rise to skeleton and muscle of head and outer layer of eye.
At the end of this stage embryo become active and exhibit continuous
twitching movement which show that embryonic development id
completed.
 Embryonic Development continued…….

Hatching: There are two mechanism involve in hatching 1)

mechanical and 2) enzymatic. In mechanical hatching egg
envelops are broken down by mechanical action such as
pressure excreted from within or mastication by embryo itself. In
enzymatic hatching, emergence of young occurs after
dissolution or softening of egg envelope by the enzyme secreted
by embryo. This enzyme is called hatching enzyme. These
enzymes are secreted into perivitelline fluids at the time of
hatching from hatching gland cells located in the epidermis of
the embryo.
 Reproduction in
Freshwater Prawn
 Sexual Dimorphism
The sexes are separate and there is well marked sexual dimorphism in fresh
water prawn Macrobrachium rosenbergii
Male Female

•Larger, 2nd pereopod much larger •Smaller, 2nd pereopod thinner and
and thicker. shorter.

•Cephalothorax larger, abdomen •Cephalothorax smaller, abdomen

narrower. broader.

•Appendix masculina and appendix •Appendix masculina is absent.

interna at endopodite of 2nd
pleopod.
•Genital pores at the base of 3rd walking
•Genital pores at the base of 5th leg.
walking leg.
•Pleura of 1st, 2nd, 3rd abdominal
•Pleura of abdomen is shorter. segment are longer and forms a brood
chamber

•Reproductive setae are absent. •Reproductive setae present.

•Distance/gap between last pair •Distance/gap between last pair walking

walking leg is very less in male. leg is more in female
•Ventral side of the abdominal •No lump or hard point at the ventral
segment has a lump- a hard point- at side of 1st abdominal segment.
the centre which can be felt with
finger.
•Reproductive setae are absent on •Reproductive setae appear on the
the ventral side of thorax and ventral side of thorax and pleopods of
pleopods of mature male. mature females such as ovipositing and
ovigarous setae. Ovipositinng setae
appears on coxa of last 3 pairs of
preopods and posteriors margin of
sperm receptacle area and the pleopods.
It helps in guiding and propelling eggs
during spawning. Ovigarous setae serve
to anchors the eggs to pleopods.

•No sperm receptacle in the thoracic •There is an sperm receptacle on the

sterna between last three pairs of between last three pairs of preopods.
preopods.
 Peak Breeding seasons

Hoogly estuary – March to May

Kerala backwater- Oct and Nov

AndhraPradesh- Aug to Oct

 Reproductive System

Male Female
1) A pair of Testis located on mid 1) A pair of ovaries located
dorsal side of cephalothorax below dorsal to the stomach and hepato-
carapace pancreas in the cephalothorax
cavity and Each ovary consist of
2) Vas deferens has 4 regions anterior lobe, lateral lobe and
abdominal lobe
•Proximal region
•Medial region 2) Oviduct which open at the
base of the coxa of 3rd pereopod
• Distal region and
4 stages of ovary
•Terminal ampoule which open at the 1. Immature
base of the coxa of 5th pereiopod 2. Early maturing
3. Maturing
4. Ripe
 Different stages of ovary
Stage colour Oocytes size Ovary size &
(mm) position
Immature transparent 0.064 to 0.128 Vary and present in
Spherical posterior most
region of carapace
cavity
Early maturing Yellowish due to 0.191 to 0.447 Occupy ¼ to ½ of
light deposition of total carapace cavity
yolk
Maturing Light orange in 0.319 to 0.547 ¾ of carapace cavity
color due to
heavier deposition
of yolk
Ripe Dark orange Ova 0.4468 to 0.7761 Occupy entire
become opaque carapace cavity
due to heavy yolk
 Mating

• Take place between hard shelled males and mature soft

shelled females which have just completed their premating
moult.
• Time between moulting and mating varies from 1.2 to 21.8
hrs (avg. 9.1 hrs).
• Male gonopore brought close to sperm receptacle of female
on thoracic region and transfer the sperm to receptacle as
spermatophore. Spermatophore adheres to the sperm
receptacle because of the glutinous nature of attachment
matrix
• After mating female may spend several hours in
prespawning preening behavior.
 Spawning
• Spawning occurs after few hrs of mating.
• Eggs laid by female are fertilized by sperm attached to
underside of thorax.
• The fertilized eggs are transferred to the ventral side of
abdomen (brood chamber) by ovipository setae and attched to
the ovigerous setae of 1st 4 pleopods and kept in place by a
membrane.
• Movements of pleopods helps in aerating the eggs.
• Adult females carrying eggs on the ventral side of abdomen is
called berried female.
• Fecundity varies from 20000-100000 eggs. However, their
first broods, (i.e. those which are produced within their first
year of life), are often not more than 5 000 to 20 000.
 Incubation

•Initially eggs are slightly elliptical (0.6-0.7 mm in long axis)

and bright orange in color

•Embryonic development proceeds.

• Incubation period avg. 19 days.

• Turns grey black 2-3 days before hatching.

• Freshly hatched larvae are called zoea.

 Macrobrachium rosenbergii,
segmentation and
embryonic development.
Times refer to period since
fertilization.
(A) 7 h - completion of second
nuclear division. (B) 8 h 45 min -
third nuclear division nearly
completed, appearance of 4
cleavage furrows. (C) 8 h 55 min -
third nuclear division completed,
tips of the 4 cleavage furrows have
met at 2 points from which the
median furrow is developing. (D)
9h - complete formation of 4
quadrants (blastomeres). (E) 14 h -
32 nuclei. (F) 24 h - completion of
segmentation. (G) 6 days-formation
of caudal papilla. (H) 7 days-
formation of optic vesicle. (I) 9
days-eye pigment developed. (J) 14
days -larva fully formed. (K) 19
days-larva ready to hatch.
 Development of Larvae
Newly hatched zoea about 1.92mm in length. It has the zoeal
characteristics such as a body distinguishable into cephalothorax and
abdomen.The segmented body is without pleopods. Larvae swim upside
down by using their thoracic appendages and are positively attracted to
light.
By stage XI they are about 7.7 mm long. Newly metamorphosed post
larvae (PL) are also about 7.7 mm long and are characterized by the fact
that they move and swim in the same way as adult prawns. They are
generally translucent and have a light orange-pink head area.
Larvae require salinity from 11-13 ppt for survival and growth.
Larval passes through 11 zoeal stages within 23-32 days
depending on temperature, food, and water quality to become PL.
 Identifying Characters of larval stages
Larval Age Total length Distinguishing features
stage (days) (mm)
I 1 1.92 Sessile eyes
II 2 1.99 Stalked eyes
III 3 2.14 Uropod present
IV 4-6 2.5 Two dorsal rostral teeth, uropod biramous
with setae
V 5-8 2.8 Telson narrow, elongated
VI 7-10 3.75 Pleopod buds appear
VII 11-17 4.06 Pleopods biramous and bare
VIII 14-19 4.68 Pleopods with setae
IX 15-22 6.07 Endopods of pleopods with appendix interna
X 17-24 7.05 3-4 dorsal rostral teeth
XI 19-29 7.73 Teeth on half of upper dorsal margin
PL 23-27 7.69 Teeth on upper and lower margin of
rostrum, adult behaviour
 Life cycle

Male +
female=
mating

Fresh
growth spawning
water

Brackish
water
Larval
development=
juvenile pass through
11 zoeal
stages