Chapter 3.
Biology of fish
3.1 Fish reproduction and growth
3.1.1. Fish reproduction
Parthenogenetic- in this type of reproduction all the
offspring’s are female individuals and there is no male
individual which is produced.
Hermaphroditism both male and female sex in a single
individual. Two types:-
a) Synchronous: Both functional testes and ovaries are found
in the same individuals at the same time, as a result self
fertilization can occurs.
b)Sequential: Individual fish with one sex in there life time
shift sex in another time.
- Protogynous: first fish have female sex, and then they develop
male sex.
- Protandros: first the fish is male, then it become females.
Bisexuality: live-bearing
Several groups of fishes are live-bearing (i.e., giving birth to free-
living larvae, rather than laying eggs).
A prerequisite to this pattern is mating, copulation, and internal
fertilization.
Bisexuality:
a) Viviparous fishes are those that give birth to live young,
meaning the eggs develop inside the mother's body and
receive nourishment from her, rather than laying eggs.
have internal fertilization and are characterized by the embryo
developing in close contact with the nourishing maternal tissue-
no egg membrane covers the embryo.
b) Ovoviviparous characterized by having internal
fertilization.
Ovoviviparous fish are those that retain their eggs within the
body until they hatch, resulting in live birth, but without
placental attachment, meaning the developing embryos are
nourished by the egg yolk, not the mother's body.
c) Oviparous fishes “egg-laying” fishes. Males and females
swim close together so that the eggs are shed into a cloud of
sperm.
The eggs are then fertilized and develop in the environment
outside the female.
An egg membrane is present, and the embryonic stage is
nourished entirely by the yolk.
The most common type of fish reproduction.
These are Oreochromis (mouth brooder) the female tilapia
carries the egg in their mouth until hatching. in other word
survival of young or egg is guaranteed by the care of female
Nile tilapia.
Some fishes such as genus tilapia are substrate spawner,
they guarantee the survival of their population by hiding the
egg or fry in substrate (cave or in the shell).
But some fishes spawn numerous eggs and produce a large
number of offspring.
• E.g The female Nile perch can produce up to 16 million
eggs at one spawning time.
• Here, survival in the case of Nile perch is not guaranteed by
parental care but it is taking place by laying large number of
eggs and no parental care.
Fecundity is the total number of eggs in the ovaries that will
mature during a particular spawning season.
Spawning: release of eggs and sperm from fish
Sex determination before dissection
Tilapia
• Sex can be determined in few no of fish species
from morphological observations of the fish
• The male has two body openings,
– anus – release face
– Urethra- discharge milt (sperm) and urine
• The female has three body openings:
– Anus- discharge faces
– Urethra- discharge urine
• hardly to see using naked eye) &
– genital - opening of the oviduct (eggs are released)
• Mature Nile tilapia distinguished by their coloration
– Males under the jaw –reddish
– Female grayish and yellow in color
Sex Determination After dissection
After dissecting the fish, sex organ can be
identified
Testis-smaller in size
Smooth, always whitish in color
Ovaries- larger in size
have granular of grain, different color
E.g.
Nile tilapia yellowish,
Gara whitish
The location of testis and ovaries are situated
above the visceral organs and under kidney
Life cycle of fish
Most fish have five stages in their life cycle. The stages in
the life cycle will go in the following order.
Stage 1: Egg
Every fish will start its life as an egg. The female fish will lay
lots of eggs in a safe place. When it is still an egg, the fish will
start to grow its organs and a tail.
Stage 2: Larvae
When the eggs are ready to hatch, they become known as
larvae. This is the second stage in the life cycle of a fish.
Larvae have their own food called a yolk sac. This is joined to
their body. The yolk sac has everything that the larva will need
to help it to grow.
Stage 3: Fry
When it has eaten all the yolk sac, the larvae change to fry.
Fry are very young fish. They are very small but will grow
quickly. Fry will start to eat by themselves and this is an
important step for helping them to grow bigger and to get
stronger.
Stage 4: Juvenile
As the fry gets larger it becomes a juvenile. The juvenile is
about the same size as an adult finger. During this stage, the
scales and the fins start to grow. However, the juvenile will
still be smaller and weaker than an adult fish
Stage 5: Adult
The last change happens when the juvenile fish turns into an
adult fish. The fish will now be fully grown and
independent.
The female fish can now lay new eggs and the life cycle
will begin all over again.
Fish maturity stages
Most of the fishes exhibit seasonal cycle in the production
of gametes. The expulsion of gametes from the body into
the surrounding water is called "spawning" resulting in
fertilization.
The entire period during which the gonads attain full
maturity and spawning takes place in the population is
called the breeding season of the species.
Maturity stages in female fish
On the basis of shape, size, colour of the ovary and other
histomorphological features, at least six maturity stages can
be recognized.
Immature
The ovaries are small, thin, thread like, translucent, pale in
colour with inconspicuous vascularisation.
The ovaries occupy only a small part of the body cavity and
ova are not visible to the naked eye.
Early maturing phase
Ovaries become slightly larger, thicker, and are light
yellowish in colour. There is an increase in the weight of the
ovary and they occupy nearly ½ of the body cavity.
Advanced maturing phase
There is a further increase in the weight and volume of the
ovaries, which have a deep yellow colour and occupy 2/3 to
¾ of the body cavity. Vascular supply increases and the
blood capillaries become conspicuous. Immature oocytes
are reduced in number.
Mature or pre spawning phase
The ovaries are further enlarged occupying almost the
entire body cavity. They are turgid, deep yellow in colour
and a large number of spherical ova are visible to the naked
eye through the thin ovarian wall.
Both translucent and opaque ova are present and the
ovaries attain their maximum weight.
Spawning phase
Ovaries are very much enlarged, occupying the entire body
cavity. They are turgid and yellow in colour with a large
number of translucent eggs. Ovarian wall is very thin,
almost transparent. Eggs are present in the oviduct also, and
the fish spawns a number of times during this period. The
ovary is now said to be in running phase. At the beginning
of this phase, ova are extruded by applying a gentle pressure
on the abdomen.
Spent phase
The ovaries are flaccid, shrunk and sac-like reduced in volume
and have a dull colour. The vascular supply is reduced. Some
unspawned large ova and a large number of small ova are
present.
Growth and Mortality of Fishes
Growth
Knowledge of fish age characteristics is necessary for stock
assessments, and to develop management or conservation
plans.
Growth is generally associated with age.
The growth of fish is quite different from that of man and
other warm-blooded animals.
They do not reach an adult size at a certain age and then
stop instead, most fish continue to grow in length and
weight until they die (indeterminate growth).
Further, the growth rate may be very fast or very slow,
depending on the amount of food and other conditions.
Growth and Mortality of Fishes - their significance in fisheries
Fish growth
1) Length weight relationship
Standard Length (SL):is distance b/n tip of the snout and tip of caudal peduncle
Fork Length (FL): Distance b/n tip of the snout and the tip of the middle ray of the caudal
fin.
Total Length (TL): is distance b/n tip of the snout and tip of he caudal fin.
Fish growth
1) Length weight relationship
Length weight relationship is curvilinear and expressed by the
following equation: TW=aTLb ;Where: TW is total weight; TL is
total length; “a” and “b” are intercept and slope, respectively.
Procedures to determine the intercept and the slope
It follows simple linear regression analysis length as independent
variable (X) and weight as dependent variable (Y).
The value of “b” ranges from 2 - 4. If b=3, the growth is called
isometric growth and there is no change in body proportion with
mass and shape hence called cube law.
In fish, isometric growth means all body dimensions (length,
width, height) increase at the same rate, maintaining a
consistent shape and weight proportional to the cube of length.
If b# 3, the growth is called Allometric growth and there is a change in body
proportions with mass and shape.
b<3 is negative allometric and describes weight is not increasing inrespect to length.
b>3 is positive allometric and describes the weight of fish is increasing more than the
length’
Length-weight relationships give information on the condition and growth patterns
of fish
Condition Factor
Condition Factor (Index) is a factor that measures the well being of fish (fish’s health)
Heavier individuals of a given length are in better condition.
Let L=20cm
Fish a=300g
Fish b=200g which one is in a better condition
Types
a) Fulton Condition Factor is calculated for each species as:
FCF= TW/TLb x100
Where: TW= Total Weight; TL= Total Length; It assumes isometric growth
Methods of fish age estimation
Fish age estimation primarily relies on analyzing growth patterns in hard body
parts like otoliths (ear stones), scales, and vertebrae, counting the number of
annuli (rings) to determine age, similar to how tree rings are used.
1) Length frequency analysis
Length frequency analysis of fish involves analyzing the distribution of fish
lengths in a sample to understand population structure, growth, and mortality,
and to assess the status of fish stocks.
Peterson introduced this method in the nineteenth century.
• The size-frequency or Peterson method of age determination depends on the
fact that fish size varies with age.
• Most fishes breed during a restricted period once a year so that size within a
given brood year is fairly uniform and distinct from size groups from other
brood years.
Data of the sample is plotted and peaks are counted which represent grouping of
2) Anatomical Method of Determining age
a) Scale method
Unlike most other animals, each fish carries in its body a record
of its age and growth that can be read accurately by experts.
The scale method of fish age determination involves counting
the number of annuli (growth rings) on a fish's scales, similar to
counting tree rings, to estimate the fish's age.
Their scales show rings that can be counted, one for each year,
like the annual growth rings in a tree.
Some of the spaces between these rings may be wide and others
narrow, depending on how much the fish grew in those years.
By measuring these rings we can trace the growth of
a fish in each year of its life.
Not applicable for fish without scales, like catfish.
Fig. scale with marked annuli (age 4+)
b) The otolith (Sagittae): The otolith or ear-stone is present in the
internal ear of the fishes and helps in balancing the body.
The otolith has annulations formed by the regular deposition of
calcium salts.
To get the otolith, fish is killed and dissected and otolith is taken out
and annulations are counted.
For this, first of all the otolith is broken and cut in transverse plane
followed by polishing with liquid of high refractive index, such as
immersion oils, creosote, etc.
The otolith thus prepared is examined under the microscope to count
the annulations for age determination and the growth rate analysis
3) Empirical method
Empirical fish age determination involves counting "growth
rings" (annuli) in hard parts like scales, otoliths, or vertebrae to
estimate a fish's age, similar to counting tree rings.
The determination of growth rate through this method requires
the knowledge of age, which can be known from time of
breeding.
The hatchlings are kept in a tank in appropriate conditions for
two or three seasons.
The growth is measured periodically. The fishes are marked with
tags and reintroduced in the water.
The fishes are recaptured at regular intervals and idea of growth
rate in relation of the time is taken.
Mortality
Fish mortality refers to the loss of fish from a stock due to
death, by natural mortality and fishing mortality.
Mortality can be defined as a decrease in number of fish
with time.
In exploited stocks
Total mortality has two components. These are:
Natural mortality (M) death due to natural causes like
predation, disease and environmental factors and Fishing
mortality (F): caused by fishing activities.
Total mortality (Z) = M + F
In unexploited stock:
F=0
Feeding
Many species are visual feeders; the position of the eye in
the head serving the particular feeding habit.
Olfaction and taste also play major roles in feeding.
Mouths structures and Feeding habit
Fishes can be divided into three feeding groups based on the
position of their mouth structure:
Surface feeders (Superior),
mid-water feeders (Terminal), and
bottom-feeders (inferior).
i. Superior (surface feeders): This kind of fish has an
upturned, scoop-like mouth which is designed to feed on prey
that swims above the fish, or perhaps on the surface of the
ocean or lake, such as insects.
ii. Terminal (pointed straight forward): Fish with a terminal
mouth position have a mouth in the middle, or center of their
head.
These fish either chase their food or feed on what is ahead
of them.
The terminal mouth position is considered the “normal”
position
iii. Inferior (mouth oriented completely downward): Bottom
feeding fish generally have underslung or inferior mouths.
Ventrally-oriented mouths or mouths located under the
fishes head that are adapted for scavenging or grazing.
Fishes are often categorized by their foods and feeding strategy into
the following:
Detritvorous:-fish that feed on detritus ( decomposed organic matter)
Scavengers:-fish species that feed on dead and decomposed animal
materials
Herbivorous:-fish that feed on plant materials.
Carnivores: fish feed on animal body. It can be:-
– Zooplanktivorus: - fish feed on zooplanktons.
– Piscivorous:-fish that feed on other fish species. eg. Nile perch.
Omnivorous:-fish feed on both plant and animal materials.
Some the morphology of fish that tells us their feeding
habits are: body shape; mouth shape and position; gill
rackers and others.
Fish Diseases
• Parasites and diseases of fish constitute one of the most
important problems in fisheries and aquaculture.
• Fishes harbor major levels of parasitic infection due to the
fact that aquatic ecosystems facilitate the reproduction,
dispersion and life history of parasitic organisms (Malta,
1984).
• In wild waters, more than 80 to 90 percent of fishes harbor
at least one species of parasite.
• Parasitism is much more common and diversified in wild
conditions than in fish farms and hatcheries.
Host -parasite relationship
• It is the outcome of three main factors, the etiological agents
(the causes of diseases), include pathogens, bacteria, viruses,
fungi, protozoa, and parasites), the host and the environment
(Natarajan, 1986).
• The study is important as it reflects the wellbeing of the fish
and helps in fish stock assessment and management
(Narasimhamurthy, 1990).
• Fish parasites may provide important information on
ecosystem conditions due to their intimate contact both the
host and aquatic environments.
• Thus, they are frequently used as bio-indicators of
environmental stress, especially with respect to pollution
(Gelnar et al., 1997; Valtonen, 1997).
Effects of Parasitism
• It causes damage to tissues and organs or it affects the
growth, reproduction or it is felt in the biochemical and
hematological characteristics of fish.
• Parasites cause damage and degrade fish tissues and make
the fish unsuitable for human consumption.
• Parasites and diseases affect both capture and culture
fisheries.
• In aquaculture systems due to the manipulation of
management practices, such as excessive fertilization,
intensive stocking, over feeding and application of
chemicals, the cultivable fish species are under severe stress
and they become subjected to infection by various kinds of
parasites and pathogens.
Stress and Disease Development in Fish
• Fish health greatly depends on aquaculture inputs such as
fertilizer, feeds, stocking density, and water quality in fish
ponds.
• Optimum stocking density, fertilization with appropriate
ration, feeding the fish with nutritionally rich feeds, and
maintaining good quality water would help in producing
healthy fish.
• Disturbance of any of these characters will stress the fish
which will ultimately affect the survival and production of
fish in culture systems.
• Increased fertilizer application, stocking density and
application of excess feeds deteriorate water quality which
ultimate cause stress to fish populations.
• Stressed fish becomes weak and subject to infection by diverse
groups of parasites and pathogens.
Stress is defined as the impact of physical, chemical or biological
factors which may cause bodily discomfort and subsequently
contribute to diseases and death of fishes.
• Stress influences the release of cortisol hormone which is
responsible for negative health impacts associated with stress.
• Besides stress being responsible for reduced growth,
reproduction and digestion,
• Stress will lower the immune function of fish, causing fish to
become susceptible to infection by microbial pathogens and
parasites.
• Aquaculture systems are always under dynamic condition
wherein the physical, chemical and biological processes are
constantly changing and their impacts are felt. These changes
often negatively impact the fish, and cause stress to fish.
Stress factors
[Link] factors
a. Temperature
Temperature influences growth rate, food consumption, feed
conversion and other body functions .
As temperature controls solubility of dissolved oxygen at high
temperature stratification, it reduces dissolved oxygen levels in
pond water which affects the fish species and the fish becomes
susceptible to infection, and in extreme temperature and
temperature shock cause fish kills.
Temperature enhances the reaction rates of chemicals, toxicity of
ammonia in freshwater causing stress to fish.
The fish stops feeding, exposed to stress and subjected to
infection and mortality occurs below optimum temperature
range (El Sayed et al., 1996).
b. Turbidity
The nutrients especially phosphates and nitrates enter the pond
through incoming water, and the application of excess inorganic
and organic fertilizers in ponds causes excessive growth of
phytoplankton in fish ponds, and the bloom thus formed besides
restricting the light penetration causes stress to fish.
c. Sound
• Fish species under culture conditions are disturbed and
stressed, when they are exposed to situations where the source
of sound is frequent and continuous.
• Areas subjected to movement of vehicles, continuous sound of
aerators, loading and unloading of fishing equipment and
unnatural and loud noises cause stress to fish which also
reduce production performance in fish ponds.
• Fish under embryonic stage are more affected under sound
waves.
d. Density
• Production to a large extent depends on the stocking density
of fish species in a unit area of water body.
• High density stocking is usually followed in intensive
rearing of fish fingerlings or in production ponds to
maximize production.
• However, such intensive stocking of fingerlings causes
stress through excessive feeding and the resultant
accumulation of metabolites in culture systems.
2. Chemical stressors
a. Oxygen
• Prolonged low oxygen levels of less than 5 mg/l produce
stress which affects feeding and metabolism of fish, and
make them susceptible to infection.
• Low dissolved oxygen syndrome (LODS) develops due to
several factors.
• It develops when biological, chemical and physical factors
interact with each other and reduce water quality thereby
exposing the fish species to poor health, and cause
production loss in fish culture ponds.
b. pH
• For healthy fish ponds, the ideal pH range is generally
considered to be between 6.5 and 8.5.
• pH levels of 5.7 and below, and levels above 9.7 cause
stress to fish in fish culture.
• pH levels of 4.0 and 11.0 are lethal to cultured fish.
• At acidic pH level of below 6.0, the fish exhibits slow
growth, and acidic pH of 4.0 or below and alkaline pH of
above 11, may lead to fish kills
c. Ammonia
• Ammonia is toxic to fish ponds and its measure in water
body or its accumulation in sediment has to be regulated.
• Ammonia is formed in ponds due the addition of protein
rich feeds, increased feeding rates, and through active
metabolism of fish.
• Excess feeds, metabolites and excreta raise the levels of
ammonia in fish ponds.
• Un-ionized ammonia levels must be kept below 0.5 mg/L.
[Link] stressors
a. Feed and phytoplankton
• Although compounded feeds with rich protein source is
very important for increased fish production, most often the
rich nutrients available in pelleted feeds negatively impact
• Application of pelleted diets is very common especially in
intensive aquaculture where the pelleted feed is
compounded with rich protein.
• It is reported that more than 80 to 85 % of the nutrients in
pelleted feeds used in aquaculture are released in to
aquaculture system as fecal matter, excretory products and
metabolized compounds such as phosphate, ammonia and
carbon dioxide which promote the production of
phytoplankton.
• Excess production of phytoplankton at high feeding rates
and the resultant waste loading and phytoplankton
proliferation produces profound impacts on cultured species
and water quality deterioration in the aquaculture ponds.
• According to Claude Boyd “As a general rule, fish
production increases linearly with feeding rate while water
quality deterioration exponentially with feeding rate”.
• The organic matter wastes that are generated through
phytoplankton photosynthesis and metabolism process of
zooplankton, bacteria and other microorganism adversely
impact water quality and fish species in ponds. (Schmittou
et al., 1998).
• Inadequate and deficient in essential amino acids in fish
feeds can be stressors causing stress to fish, and the fish
becomes susceptible to infection.
b. Parasite and pathogens
• Heterotrophic bacterial populations, fungal and viruses are
pathogenic to fish under unfavorable environmental
conditions.
• Most metazoan parasites multiply rapidly under stressful
condition and cause stress to fish when they infect the fish
in large numbers.
Based on behavioral and physiological state of fish species,
the response of fish to stress is classified as alarm stage,
adaptive stage and exhaustion stage.
At alarm stage: the fish tries to avoid and tries to escape
from the stressors.
Adaptive Stage: If fishes cannot avoid the situation and
escape from the stressors, the fish tries to adjust to the new
environment, both physiologically and behaviorally to
survive in the new environmental conditions.
At this stage its growth and metabolism may not be
maintained as in the original level because, fish at this stage
deals mainly with the stress as a priority. It is at this stage
the disease problems are more likely to occur. Extended
exposure of fish to such environmental conditions disturbs
and deteriorates its normal function and decreases its
chances of survival.
Exhaustion stage: If the environmental change is so great
that the fish cannot adapt to it, the fish reaches an
exhaustion stage and it dies.
Defense mechanism
• Stressors can evoke nonspecific responses in fish which develops
some defense mechanisms to overcome the stress mainly to
maintain its normal function.
• It is an adaptive measure to withstand the disturbance and
maintain its homeostatic state.
• As defense mechanism, the body of fish secretes excess mucus to
prevent the entry of pathogens and parasites.
• The enzymes and antibodies of the mucus kill all invading
microorganisms.
• Inflammation is another defense mechanism against parasites and
• Another defense mechanism that is developed in fish is the
production of antibodies to fight specific pathogens.
Microbial diseases
• Bacteria, fungi and virus constitute the microbes which are
abundant in water and sediment in freshwater, brackish water
and sea water.
• They are present in environment as natural flora become
pathogenic and cause diseases under adverse environmental
conditions (Austin and Austin, 1999).
• Poor water quality, bad feeding regime, discharge of excess
drugs and feeds from farming systems result in decreased
activity of the immune system, and a sudden outbreak of
diseases in fish.
• Microbial pathogens produce diseases and making the fish
unsuitable for human consumption and cause great economic