Feeding

&
Food Processing
1. Structure
2. Function (behavior,
physiology)

3. Nutritional needs

4. Digestive efficiency
Food capture

 Mouth and pharyngeal cavity

 Jaws

 Teeth - jaw, mouth, pharyngeal

 Gill rakers
Fish Feeding - function

 Herbivores
 < 5% of all bony fishes, no
cartilaginous fishes
 browsers - selective - eat
only the plant
 grazers - less selective -
include sediments
 Detritivores
 5 - 10% of all species
 feed on decomposing
organic matter
Fish Feeding – function

 Carnivores
 zooplanktivores
 suction feeding
 ram feeding (engulfing
prey while swimming
forward)
 benthic invertebrate
feeders
 graspers
 pickers
 sorters
 crushers
Fish Feeding – function

 More Carnivores
 fish feeders
 active pursuit
 Stalking (hunt and kill)
 Ambushing (sit and
wait)
 Luring (attract)
Fish feeding behavior

 Fish feeding behavior integrates

morphology with perception to obtain
food:

 Search --> Detection --> Pursuit --> Capture -->

Ingestion
Feeding
behavior
 Fish show versatility in
prey choice and
ingestion

 Behavior tightly linked

to morphology
(co-evolution)
Fish feeding behavior

 Behavior tends to be optimizing when

choices are available
 Optimal = maximize benefit : cost ratio
 More for less
 Select the prey that yields the greatest
energetic or nutrient “return” on the energy
invested in search, pursuit, capture, and
ingestion
Fish digestive physiology

 After ingestion of food, gut is responsible for:

 Digestion - breaking down food into small, simple
molecules
 involves use of acids, enzymes
 Absorption - taking molecules into blood
 diffusion into mucosal cells
 phagocytosis/pinocytosis by mucosal cells
 active transport via carrier molecules
Digestive Apparati

trout
carnivore

catfish
omnivore

carp
omnivore

milkfish
planktivore
Fish Digestion

 Two major groups: w/stomach, w/out

 w/out stomach: cyprinids (carps)
 w/stomach: cold-water salmonids, warm-water
catfish, tilapia, eels, grouper
 all “pure” predators have a stomach and teeth
 relative gut length (RGL): gut:body length
 high RGL = species consuming detritus, algae (high
proportion of indigestible matter)
Relative Gut Length

Species Feeding RGL

Labeo horie Algae, detritus 15.5
Garra dembensis Algae, inverts 4.5
Barbus sharpei Plants 2.8-3.1
Chelethiops Zooplankton 0.7
elongatus
Chela bacaila Carnivorous 0.9
Fish Digestive Morphology:
Major Divisions
 Mouth
 Esophagus
 Pharynx
 Stomach
 Intestine
 Rectum
 Secretory glands (liver and pancreas)
 often difficult to distinguish
Gastrointestinal Tract
 Esophagus
 Stomach
 large in carnivores,
small in
herbivores/omnivores
 Pyloric caeca (amino acid
& carbohydrate
absorption)
 Intestine
 short in carnivores,
long in herbivores-
omnivores
 Anus - separate from
urogenital pore
GI Tract- Secretory Glands
 Liver
 produces bile (lipolysis)
 stores glycogen
 stores lipids
 Pancreas
 digestive enzymes
 proteases - protein breakdown
 amylases - starch breakdown
 chitinases - chitin breakdown
 lipases - lipid breakdown
Digestive Anatomy:
Mouth/Esophagus
 Channel catfish: large mouth/esophagus, capture prey,
slightly predaceous, mouth has no teeth, no gizzard
 Common carp: small mouth for bottom feeding,
pharyngeal teeth (teeth in pharyngeal arch) , grinds food
 Tilapia: combination of bottom feeder, predator,
efficient plankton feeder, uses gill rakers, pharyngeal
mucous
Digestive Anatomy: Stomach

 Channel catfish: have true stomach that secretes

HCl and pepsinogen (enzyme)
 Common carp: no stomach; however, “bulb” at
anterior end of digestive tract, bile and
pancreatic secretions empty into intestine
posterior to cardiac sphincter, no secretion of
gastrin (low pH)
 Tilapia: modified stomach, secretes HCl, well-
defined pocket, pH varies w/digestal flow, has
pyloric sphincter
Digestive Anatomy:
Intestine
 Channel catfish: length less than whole body, no
large/small version, slightly basic pH, digestive
secretions, nutrient absorption, many folds for
absorption
 Common carp: digestive tract is 3x whole body
length, similar in activity to that of channel
catfish
 Tilapia: tract is 6-8x that of body length,
activities similar to that of other species
Digestive Anatomy: Liver and
Pancreas
 Both organs produce digestive secretions
 Liver produces bile but is also the primary organ
for synthesis, detoxification and storage of many
nutrients
 Pancreas is primary source of digestive enzymes
in most animals
 It also produces zymogens (precursors to
enzymes)
Fish Digestive Physiology
 Digestion is accomplished in
 Stomach
 low pH - HCl, other acids (2.0 for some tilapia)
 proteolytic enzymes (mostly pepsin)
Digestive Processes:
Stomach
 Catfish as an example - its digestive processes are similar
to that of most monogastric animals

 Food enters stomach, neural and hormonal processes

stimulate digestive secretions
 As stomach distends, parietal cells in lining secrete
gastrin, assisting in digestion
 Gastrin converts the zymogen pepsinogen to pepsin (a
major proteolytic enzyme)
 Some fish have cirulein instead of gastrin
Digestive Processes:
Stomach
 Flow of digesta out of stomach is controlled by
the pyloric sphincter
 Pepsin has pH optimum and lyses protein into
small peptides for easier absorption
 Minerals are solubilized; however, no lipid or
COH is modified
 Mixture of gastric juices, digesta, mucous is
known as chyme
Fish Digestive Physiology
 Digestion is accomplished in
 Stomach
 Intestine
 alkaline pH (7.0 - 9.0)
 proteolytic enzymes - from pancreas & intestine
 amylases (carbohydrate digestion) - from
pancreas & intestine
 lipases (lipid digestion) - from pancreas & liver
(gall bladder, bile duct)
Digestive Processes:
Intestine
 Chyme entering the small intestine stimulates secretions
from the pancreas and gall bladder (bile)
 Bile contains salts, cholesterol, phospholipids, pigments,
etc.
 Pancreatic secretions include bicarbonates which buffer
acidity of the chyme
 Zymogens for proteins, COH, lipids, chitin and
nucleotides are secreted
 e.g., enterokinase (trypsinogen --> trypsin)
 Others: chymotrypsin, carboxypeptidase,
aminopeptidase, chitinase
Digestive Processes:
Intestine
 Digestion of carbohydrates is via amylase,
which hydrolyzes starch
 Others: nuclease, lipase
 Cellulase: interesting in that it is not
secreted by pancreas, but rather produced by
gut bacteria
 intestinal mucosa also secretes digestive
enzymes
Fish digestive physiology
 Absorption is accomplished in
 Intestine
 diffusion into mucosal cells
 phagocytosis/pinocytosis by mucosal cells
 active transport via carrier molecules
Digestive processes:
Absorption
 Most nutrient absorption occurs in the intestine
 Cross-section of the intestinal luma shows that it is
highly convoluted, increasing surface area
 Absorption through membrane is either by passive
diffusion (concentration gradient)
 Or by active transport (requires ATP)
 Or via pinocytosis (particle engulfed)
 Nutrients absorbed by passive diffusion include:
electrolytes, monosaccharides, some vitamins, smaller
amino acids
Digestive processes:
absorption
 Proteins are absorbed primarily as amino acids,
dipeptides or tripeptides
 triglycerides are absorbed as micelles
 COH’s absorbed as monosaccharides
 calcium and phosphorus are usually complexed
together for absorption
 all nutrients, excluding some lipids, are absorbed
from the intestine via the hepatic portal vein to the
liver
Summary of Digestive Enzymes
Site/Type Fluid/enzyme Function/notes
Stomach HCl Reduces gut pH,
pepsiongen
Gastric secretions Zymogen, pepsinogen, HCl Proteolysis
Amylase COH’s
Lipase Lipids
Esterase Esters
Chitinase Chitin
Pancreas HCO3 Neutralizes HCl
Proteases Cleave peptide linkages
Amylase COH’s
Lipase Lipids
Chitinase Chitin
Liver/bile Bile salts, cholestrol Increase pH, emulsify
lipids
Intestine Aminopeptidases Split nucleosides
Lecithinase Phospholipids to glycerol
+ fatty acids
Fish Nutritional Needs

 High protein diet:

 Carnivores - 40 - 55% protein needed

 Omnivores - 28 - 35% protein needed

 Birds & mammals - 12 - 25% protein needed

 10 essential amino acids (PVT. TIM HALL)
Fish Nutritional Needs

 High protein diet

 Proteins needed for growth of new tissue

 Proteins moderately energy-dense (ectotherms, low
gravity)
 Few side-effects - ease of NH4+ excretion
Nutritional efficiency in
fishes
 Fish more efficient than other vertebrates:

 Conversion factor = kg feed required to produce

1 kg growth in fish flesh

 Fishes: 1.7 - 5.0

 Birds & mammals: 5.0 - 15.0
Nutritional efficiency in
fishes
 Fish more efficient than other vertebrates

 Ectothermy vs. endothermy

 Energy/matter required to counterbalance
gravity
 Bias of a high-protein diet
Components of Fish Diet
PROTEIN
 30-50% in most fish diets
 IMPORTANT FOR FISH – structure/muscle,
gonads and growth
 Linear relationship between daily protein and
growth
 Utilization of protein relatively constant and
independent of feeding - (carnivore, omnivore,
herbivore)
CARBOHYDRATES

 Not very important for most fish species

 Appear as sugars and starches
 Trout have limited ability to digest
sugars/starches

 May affect fish health

 Catfish digest starch well
CARBOHYDRATES

 Fish lack the enzyme cellulase

 Unable to break down cellulose
 Fiber usually considered to have 0 nutritional
value
 Levels of 10 to 20% have resulted in growth
depression in rainbow trout
ESSENTIAL FATTY ACIDS

 Triglycerides with polyunsaturated fatty

acids
 w 3 and w 6 - Omega or linolenic series (n-
series) of fatty acids
 Digestibility of lipids is 85-95%
 Major source is fish oil (salmonids) or
soybean oil (ictalurids)
MINERALS
 Required by all animals – fish can uptake some from
water
 Formation of skeletal tissue
 Respiration
 Digestion
 Osmoregulation (SW = high minerals/salts, FW = low
 Major minerals
 Ca, Phos, Sulphur, sodium, chloride ion, K+, Magnesium
 Trace minerals
 Cobalt, Copper, Fluorine, Iodine, Iron, Manganese, etc.
VITAMINS

 COMPLEX SUBSTANCES
 FAT SOLUBLE VITAMINS
 A- Retinol - carotenoids converted in intestinal
mucosa
 E - Tocopherol - antioxidants in fish diets
 K - Two forms in green plants - blood clotting
and bacteriostatic (bacterial growth preventive)
 D - calciferols not well understood
WATER SOLUBLE VITAMINS

 B1 - coenzyme of carbohydrate metab

 digestion, reproduction, nervous system
 B2 - Riboflavin- eyes function
 B6 - Pyridoxin
 Pantothenic acid
 Fat & carb breakdown
 Inositol – improves reduced growth rates
WATER SOLUBLE VITAMINS

 Niacin – nervous system function

 Biotin – metabolism of fats and carbs
 Choline - improves growth and conversion
 Cyanocobalamic (B12)- nerve cells function
& synthesis of nucleic acids
 Folic acid – brain function
WATER SOLUBLE VITAMINS

 Ascorbic Acid (C)

 Important - Collagen skeletal systems
 Wound healing, disease resistance
 Fish and primates can not synthesize
Diet Additives: Hormones

 Hormonal control used to produce mono sex

cultures of fish
 reduces reproduction/increases growth
 ex. Androgenic steroids (ethyltestosterone) fed
to tilapia fry = 90% males
 does not work the same on all fish
 17-alpha-methyltestosterone improves growth
and survival in salmonids
 andorgenic better than estrogenic
Other dietary factors
 Attractants
 Attract fish by sight or smell (shrimp meal, fish oil,
fish meal, etc.)
 Pigments
 External

 Crayfish, red snapper

 Flesh color – pink in salmon or trout


 Must be obtained from feed (crustaceans, yeast,
plants/algae)
 Physical Properties
• Ground meals are not suitable
for feeding to aquatic animals
due to poor ingestion, feed
conversion, and reduced water
quality
• pellets need to be stable in
water until consumed by the
target animal
• good pellet stability required for
slow-feeding species such as
shrimp various sizes of fish feed
• particle size is important to particles
insure appropriate consumption
 Circulatory System

Fish
Closed system

single circuit
heart gills body heart
 Special conditions for
fish circulation
– Environment is oxygen poor (1-8 ppm on average)
– Heart is simplest of vertebrates
– Fish have less blood volume than other vertebrates
 Special conditions for
fish circulation

• Adaptations by fish
– Composition of blood (many different hemoglobins)
–polymorphism of hb-monomeric, oligomeric, different
structures and number of alpha and beta chains, hb I – hb
IV

Hb IV At low pH it displays oxygen release into the

swim bladder and the retina. At pH 7.8, hemoglobin
IV is totally oxygenated, while at pH 6.0, it is fully
deoxygenated
 Special conditions for
fish circulation
• Adaptations by fish

– Morphology of circulatory apparatus

– Behavioral responses to oxygen availability
• Temporal and spatial positions
• Aerial repiration or aquatic surface respiration
• Reducing activity
• Holding air bubble in mouth
Functions of the Circulatory System

• Delivers oxygen
• Delivers nutrients
• Removes metabolic waste
• Fights pathogens
Components of the Circulatory
System to Study
• Blood
– Erythrocytes-red cells
– Leukocytes –white cells
– Structure of Hemoglobin

• Vascular system
– Heart
– Vessels
Formation of Fish Blood Cells
• Formed from hemocytoblast
(A blood cell derived from embryonic mesenchyme)

• Blood forming site differs between fishes.

Formation of Fish Blood Cells
– Agnatha (jawless hagfish & lampreys)
• Mesodermal envelope around gut in hagfish
• Fatty tissue dorsal to nerve cord in lampreys
- “fat column”
Formation of Fish Blood Cells
– Elasmobranchs (cartilaginous fish sharks, rays
& skates)
• Leydig organ (near esophagus)
• Epigonal organ (around gonads)
• Spleen
 Formation of Fish
Blood Cells
• Teleosts (bony fish)
– Kidney, Spleen, Cranium (cranial
mesenchyme at embryonic stage)

• Fish bone has no marrow

 Components of Fish Circulatory
Systems
• Blood:
– aqueous solution
– solutes (proteins, sugars, minerals)
– blood cells
• erythrocytes (red blood cells)
• leucocytes (white blood cells)
– lymphocytes
– thrombocytes
– monocytes
– granulocytes
 Blood Cells
• Leukocytes
• are numerous, ranging from 20 to 50
thousand,
• but some species have levels of up to
• 100,000/mm3
 Erythrocytes-red cells

• Most abundant fish blood cells

(800 thousand to 3.5 million/mm3)

• Nucleated

• Size range exists (elasmobranchs usually larger,

but fewer)

• More active species have more red blood cells

 Hemoglobin of Fish Erythrocytes
• Primary means for transporting oxygen
– In some fish up to 15% may be in plasma

• A few fish have no hemoglobin (rare

situation)
e.g. Channichthyidae in Antarctic waters

– Environmental oxygen high

– Low metabolic requirements
- very low temp, high O2 solubility, no RBC
Fish Hemoglobin Characteristics

• Structure is different in different fish

– Monomeric
– Single-heme peptide molecules
– Found in Agnatha

• Tetrameric
– Four peptide chains
 Fish Hemoglobin Characteristics

• May differ in many features

– Composition of amino acids
– Affinity for oxygen
– Some salmonids have ~18 different kinds
Having Different Hemoglobin Types

• Different hemoglobins have different responses

to:
- temperature
- oxygen absorption

• Allows fish to deal with changing conditions

– Important for migratory species (salmon)

• Some fish gain or lose types as they age

 Blood Oxygen Affinity

• pH
– Decreasing pH decreases affinity
– Often associated with carbon dioxide

Root effect: increased proton or carbon dioxide

concentration (lower pH) lowers hemoglobin's affinity and
carrying capacity for oxygen
 Blood Oxygen Affinity

Bohr Effect: hemoglobin's oxygen binding

affinity is inversely related both to acidity and to
the concentration of carbon dioxide

– Increase in CO2 drives off O2

– Decrease in blood pH magnifies Bohr effect
 Blood Oxygen Affinity
• Temperature
– Increase in temperature depresses oxygen
affinity and capacity (total amount bound)
– Results in fish having narrow temperature
tolerances
 Fish Circulatory System
• Primary circulation
– Closed system
• Heart
• Arteries
• Capillaries
• Veins

• Secondary circulation
– Collects blood that is outside the primary
– Originally thought to be lymphatic

• No lymph or lymph nodes.

 Chambers of the Fish Heart
(1) Sinus venous
– Collects blood from venous ducts

(2) Atrium
– Accelerates blood flow

(3) Ventricle
– Large muscled chamber
– Provides propulsive flow for circulation

(4) Bulbus arteriosus (bony)

Conus arteriosus
– Changes blood from a pulse to continuous
flow
 Divisions of Primary Circulation

• Branchial circulation
– Blood from heart through gills

• Systemic circulation
– Blood from gills to body to heart

• Blood flow is continuous from heart, to lungs, to body,

back to heart
 Proximity of Heart & Gills

Exceptions to Normal Circulation

• Hagfish have accessory inline hearts
• Lungfish have pulmonary circulation
• There are also many small adaptations in some species.
 Structure of the Fish Heart

• Four chambered heart

• All four chambers are in line
• The heart pumps only venous blood
• Except for a few air breathing fish, all
blood is pumped to the gills
(Vascular circulation in lungfish.)
 Conus vs. Bulbus Arteriosus

• Conus Arteriosus
– Contractile
– Cardiac muscle
– More than one valve

• Bulbus Arteriosus
– Elastic
– Mostly connective tissue
– One valve dividing it from ventricle
 The Hagfish Heart
• Most primitive
• Sinus venous well developed
– Divided into two parts to receive different
veins
• Bulbus arteriosus
• Have 3 additional hearts
– Cardinal heart in head
– Caudal heart near end of tail
– Portal heart – pumps blood through liver
 Lamprey Heart

• Largest of fish hearts

• Atrium overlies ventricle
• Bulbus arteriosus
 Elasmobranch Heart
• Conus arteriosus
• Sinus venosus with almost no cardiac
muscle
• Ventricle has two muscle layers
– Compacta = compact outer layer
– Spongiosa = inner layer
 Teleost Heart

• Variation exists across the group

• Sinus venosus is thick-walled
• Most have bulbus arteriosus
• Some have conus arteriosus (usually more
primitive)
 Lungfish Heart

• Atrium is divided into two parts by an

incomplete septum
– Functional 3 chamber heart
– Like amphibians
– Right atrium larger than left
– Right = deoxygenated from sinus venosus
– Left = oxygenated from pulmonary vein
 Secondary Circulation
• analogies between the secondary system and lymphatics
• Narrow-bore capillaries (2–20 μm diameter × 200–300 μm
length)
• arterioarterial (interarterial) anastomoses arise from the
efferent filamental and efferent branchial arteries of the
gill, dorsal aorta, and segmental arteries and condense to
form progressively larger secondary arteries.
• Secondary capillaries have been identified in gills, skin,
fins, peritoneal lining, and oral mucosa of bony fish and in
gills of many nonteleostean species.
 Secondary Circulation
• Branchial secondary veins drain into primary
veins of the head, systemic secondary veins
primarily drain into the caudal heart, or into large
cutaneous veins
• from there blood enters the primary venous
system.
• Preliminary studies indicate that the secondary
circulation is a large-volume (48.4 ml·kg−1, 1.5
times greater than the primary system), low-flow
(∼0.3% of cardiac output) network.
 Respiration
• Most terrestrial vertebrates have internal lungs that must
be ventilated through bidirectional movement of air to
replenish the oxygen (O2) supply

• Most fish have external gills that are ventilated by a

unidirectional flow of water, by pumping or swimming

• Fine sieve structure of gills very efficiently extracts O2

from water.
• Efficient O2 uptake is vital to fish because of its low water
solubility.

• Solubility decreases with increased temperature & salinity

 Oxygen solubility determined by
temperature

Temp ( C) O2 con. at sat. O2 con. at sat.

(mg/l) – Fresh (mg/l) – Salt
0 10.3 8.0
10 8.0 6.3
20 6.5 5.3
30 5.6 4.6
• Gills are the main
site of gas exchange
in almost all fishes.

• The gills consist of bony or

stiffened arches (cartilage) that
anchor pairs of gill filaments.
Numerous lamellae protrude from
both sides of each filament and are
the primary sites of gas exchange.
Microscopic gill structure:
showing gill filament and
lamellae (Red blood cells
evident.)
How can fish remove 80 -
90% of O2 available from
water?
• Short diffusion distance at gill
site

• Large surface area for diffusion

at gill site

• Counter current exchange of

gases at gill site

• Large volume of water passes

over gills
1. Fill mouth cavity (open
mouth, expand volume of
mouth, expand volume of gill
chamber with operculum
closed)

2. Fill gill cavity (close mouth,

squeeze mouth cavity, expand
gill cavity, with operculum
closed)
3. Expel water from gill
cavity (squeeze mouth and
gill cavities, open operculum)

4. Reset for next cycle

Oxygen Exchange in
Fish
• Fish employ the
countercurrent
system to extract O2
from the water.

• This system moves

water flowing
across the gills, in
an opposite
direction to the
blood flow creating
the maximum
efficiency of gas
exchange.
 Countercurrant*
Close-up
• Blood flow through lamellae
is from posterior to anterior.

• Water flow over lamellae is

from anterior to posterior.

• Counter-current allows for

diffusion from high O2 in
water to low O2 in blood
across entire length of
lamella.

* When he blood and water

flows in the same direction,
the co-current system, it will
initially diffuses large
amounts of oxygen but the
efficiency reduces when the
fluids start to reach
equilibrium.
 4 gill arches on each side of body
2 rows of gill filaments on each arch (demibranchs)
100’s filaments per demibranch - closely spaced
1000’s lamellae per gill filament

gill area = 10 to 60 times that of body surface area, depending

on species

HUGE potential to extract Oxygen from water

 Auxiliary Respiratory Structures

• Skin - diffusion of oxygen from water into dense network

of capillaries in skin (eels), Thin skin (larval fish) supplies
50% of O2 needed.
• Swim bladder - vascularized physostomous swim bladders
(gars)
• Lungs - modified swim bladder (lungfishes)
• Mouth - vascularized region in roof of mouth (electric eel,
mudsuckers)
• Gut - vascularized stomach or intestinal wall (armored
catfish, loaches)
 Branchial vs. Ram Ventilation
Branchial
• Mouth
• Pharynx
• Operculum
• Branchiostegals (filaments, lamella)

Ram
• Uses same parts, but not the pumping energy required. Sharks
primarily. Once swimming speed is achieved...no need to
actively vent buccal cavity. However, this can only be used
consistently by strong swimmers (sharks, tuna).
Gas Transport in Fishes
 Hemoglobin (Hb) of Fish Erythrocytes

• Primary means for transporting oxygen

– In some fish up to 15% may be in plasma

• A few fish have no Hb (rare situation)

– Environmental oxygen high

– Low metabolic requirements
Fish Hemoglobin Characteristics
• Structure is different in different fish
– Monomeric
– Single-heme peptide molecules
– Found in Agnatha

• Tetrameric
– Four peptide chains
 Fish Hemoglobin Characteristics

• May differ in many features

– Composition of amino acids

– Affinity for oxygen

– Some salmonids have up to 18 different Hbs

 Having Different Hemoglobin Types

• Different Hbs have different responses to:

- temperature
- oxygen absorption

• Allows fish to deal with changing conditions

– Important for migratory species

• Some fish gain or lose types as they age

 Blood Oxygen Affinity

• pH
– Decreasing pH decreases Hb affinity for O2
– Often associated with carbon dioxide

• Carbon dioxide
– Increase in CO2 drives off O2 (Bohr effect)
– Decrease in blood pH magnifies Bohr effect
 Blood Oxygen Affinity

• Temperature
– Increase in temperature depresses oxygen
affinity and capacity

– Results in fish having narrow temperature

tolerances

• Organic phosphate
– ATP depresses O2 affinity
– Urea increases O2 affinity
 How to inflate a swim bladder

• Gas gland is location of action in wall of swim bladder

(rete mirabile and surrounding tissues)

• Need to pry O2 molecules from Hb molecules in gas gland

• Need to accumulate enough O2 (>pO2) in solution in blood

plasma to generate a diffusion gradient from distal end of
rete mirabile into lumen of swim bladder
 Prying O2 from Hb
• Change of pH in blood causes change in bond strength of Hb for O2

• Bohr effect--decrease in affinity of Hb for O2 due to decreasing pH or

increasing pCO2
affinity: strength of attraction of Hb for O2

• Root effect--decrease in capacity of Hb for O2 due to decreasing pH or

increasing pCO2 (extreme Bohr effect)
capacity: total quantity O2 of that Hb can carry

• more active species tend to have greater Bohr & Root effects
 Effect of pH on Hb*--2 components
100 pH 8.02
% Saturation of Hb

capacity Root
pH 7.47
affinity
50
Bohr

0
0 80 160
*data for winter flounder
pO2 mm Mercury Air saturation
 How to cause pH to drop in tissues of gas
gland?
• Regular metabolic processes result in release of H+, either
from glycolysis (lactic acid) or aerobic metabolism (CO2)

• Increase metabolic activity in tissues surrounding rete

mirabile = decrease of pH
 How to cause free O2 to accumulate in
distal end of rete?
• Another counter-current exchange system:
– long capillaries that fold back on self
– afferent (incoming) part of capillary experiences drop
in pH, Hb loses O2
– efferent (outgoing part of capillary has higher partial
pressure (concentration) of dissolved O2 than afferent,
so
– O2 diffuses into afferent arm, causing supersaturation
of blood at distal end of rete with O2
 Counter-current multiplication system

lactic acid
afferent blood

O2heme O2heme pO2

swim bladder
1 pO2
2 pO2

O2heme pO2

efferent blood
 Function of Rete Mirabile
2. Lactic Acid Secretions
heme dumps O2 to plasma
pO2 diffuses into swim bladder to equil.
 Counter-current multiplication system

lactic acid
afferent blood

O2heme O2heme pO2

swim bladder
1 pO2
2 pO2

O2heme pO2 3

efferent blood
 Function of Rete Mirabile
3. Multiplying effect: pO2 diffuses from efferent capillary to
afferent cap. Also, longer capillaries yield more efficient
exchange of oxygen, higher pressures
 What about CO2?
• the O2 inside...how do the CO2 outside

• Again, blood also is involved in CO2 transport.

• Three mechanisms to move CO2 outside cell to be excreted.

(1) Simple dissolution in plasma
(2) Binding to proteins/formation of carbamino groups.
(3) Dissociation into carbonic acid by pH change (greatest
amount of CO2 transformed this way.)
 “Chloride Shift”

dissociation is slow

CO2 + HbO2
HHbO2 + O2

HbCO2 + O2

O2 O2
Lungs
 Buoyancy strategies

1. Low density compounds

2. Lift generated by swimming
3. Reduction of heavy tissues
4. Swim bladder (air bladder)
 1. Low density compounds:

Substance Specific Gravity

Bone 2.0
Muscle 1.05
Cartilage 1.1
Freshwater 1.002 @20C
Saltwater 1.072 @20C
Lipids 0.9-0.92
Squalene 0.86
 2. Lift generated by swimming:

thrust
lift
lift
sharks
3. Reduction of heavy tissues

deepwater fishes

umbrella mouth gulper Eurypharynx pelecanoides

 4. Swim bladder

• low density
• adjustable
• most osteichthians
• lost secondarily in some species
 Two types of swim bladders:

• Physostomous
– pneumatic duct
– soft-rayed teleosts--herrings, salmonids, catfishes,
cyprinids, eels, etc.
• Physoclistous
– blood/circulatory system
– spiney-rayed teleosts--Acanthopterygii, sunfishes, perch,
most marine fishes
 Effects of depth on swim bladder volume

• pressure increases 1 ATM/10m

• swim bladder must be adjustable
• Physostomous fishes adjust volume by gulping or
spitting air.
– mostly shallow water species
– gas-spitting reflex
– gulp air at surface
 Physoclistous inflation/deflation

• circulatory system--source of gases

• rete mirabile (wonderful net) --inflation
• oval window--deflation
• Problem: fish need greater pressure in swim bladder
than is achieved by equilibrium with blood gases
 Physoclistous swim bladder
• Pressures up to 300 ATM in some deep sea fishes

• Gases mostly O2, some CO2 and N2

• Guanine crystals in SB wall reduce permeability

• Deflation occurs at oval window

– dense bed of capillaries on SB wall
– gasses diffuse into blood
– mucus layer covers window during inflation
Growth &
Metabolism
 Growth:
• Longevity
– unconfirmed reports of carp 200-400 yr.
– authenticated records for carp 50 yr.
– large fish-few > 12-20 yr.
– some marine spp > 100 yr. thornyspines, orange roughy
– many small spp-2 yr. or less (sardines, anchovies)

Note: aging with scales, bones, otoliths

 Many Generalities:
• Sexual Dimorphism: females can be larger than males

• Growth rate a function of temperature

• Longevity inversely proportional to temperature

• Stress reduces growth

• Dominance hierarchies - dominant get food

• Overcrowding can lead to stunting

• Indeterminate growth - grow throughout life

• Growth highly variable - decreased weight gain

 Bioenergetic Definition of Growth
• energy accumulation (calories) vs. length or weight
 Bioenergetics continued:

• Energy Budget:

I=M+G+E
where: I = ingested energy
M = energy expended for metabolism
G = energy stored as growth
E = energy lost to environment
Bioenergetic Energy Budget:
Bioenergetic Energy Budget:

I
Bioenergetic Energy Budget:

I M
Bioenergetic Energy Budget:

I M G
Bioenergetic Energy Budget: heat

I M G
 Bioenergetics continued:

Ex: M = energy for body repair

maintenance
activity
digestion
 Bioenergetics continued:

Ex: E = energy in feces

ammonia, or urea
mucus
epidermal cells
 Terms:

• Standard Metabolic Rate

– maintenance met.; no growth, no activity
• Routine Metabolic Rate
– typical met.; routine growth & activity
• Active Metabolic Rate
– max. aerobic metabolism
 Factors Affecting Growth: Temperature
routine
active
Metabolic Rate

standard

scope

{
activity

growth

Where would
Temperature growth be best?
 Factors Affecting Growth: Temperature

normal O2
Metabolic Rate

reduced O2

reduced scope
reduced growth

Temperature
Routine Metabolism Factors Affecting Growth: Dissolved oxygen

O2 regulator (most species)

O2 conformer
critical O2 concentration

0 4 8
Dissolved Oxygen mg/L
Osmoregulation & Excretion
 Kidney

• Mesonephric, Paired, elongated, placed above

the alimentary canal
• No sexual dimorphism
• Head kidney (lymphoid, hemopoietic,
interrenal tissue)
• Trunk kidney (nephrons)
Fig. 30.9a
Fig. 30.9b
 Nephron
• Renal corpuscle containing a vascularized glomerulus
• Ciliated neck connecting renal corpuscle with tubule
• Initial Proximal segment of tubule
• Second Proximal segment of tubule
• Intermediate segment
• Distal segment
• Collecting duct system
Figure 1. Trunk kidney, transverse section (Formalin, H&E, Bar = 325 µm).
1. kidney; 2. vertebrae; 3. spinal cord; 4. skeletal muscle; 5. abdominal cavity.
• Figure 2. Glomerulus (Formalin, H&E, Bar = 10.7 µm). 1. Bowman's space;
2. endothelial cell; 3. mesangial cell; 4. visceral epithelium of the renal capsule;
5. parietal epithelium of the renal capsule; 6. red blood cell in capillary.
Figure 3. Kidney tubules, transverse section (2) (Formalin, H&E, Bar = 16.7 µm).
1. first proximal tubule; 2. second proximal tubule; 3. intermediate tubule segment;
4. distal tubule; 5. collecting duct; 6. brush border; 7. mitotic epithelial cell;
8. melanomacrophage; 9. red blood cells.
• Figure 4. Large collecting duct, transverse section (Formalin, H&E, Bar = 18.1 µm).
1. large collecting duct ; 2. distal tubule; 3. brush border; 4. low columnar
epithelial cells;
5. intercalating cell; 6. rodlet cell; 7. pseudostratified columnar epithelial cell; 8.
mucous cell;
9. melanomacrophage; 10. artery; 11. fibrous connective tissue and smooth
muscle.
 Excretory Processes-Kidneys
• Excretory systems produce urine by refining a
filtrate derived from body fluids
• Key functions of most excretory systems
– Filtration: Filtering of body fluids
– Reabsorption: Reclaiming valuable solutes
– Secretion: Adding nonessential solutes and wastes
from the body fluids to the filtrate
– Excretion: Processed filtrate containing nitrogenous
wastes, released from the body
 1 Filtration
Capillary

Filtrate
Excretory
tubule

2 Reabsorption
Monovalent ions in FW. Urea
in elasmobranchs

3 Secretion
Water, Na+, CL+, Mg2+, So4-
Nit. wastes
Urine

4 Excretion
 Structure & number of Nephron
• Extreme diversity among spp.
• Numerous, highly vascularized glomeruli in
FW fish
• less, small, poorly vascularized glomeruli in
SW fish
• Avascular glomeruli in sea horses
 Urine
• Renal portal system
• Venous blood supply except to glomeruli
• Systemic pressure is average 20 mm HG
• FW spp. produce 0.1 – 1.4 ml of urine per hour
per 100 g of BW
• SW spp. produce 1/10 of these values
• Clearance of water via kidney is up to 30% in FW
• Nil in SW
 Urine
• Urine is more dilute as compared to plasma in
FW
• Iso-osmotic with plasma in SW
• No. of functioning glomeruli can be adjusted
• GFR and urine production will change without
changing the its conc.
• E.g. 45% of nephrons functional in FW while
5% in SW
 Kidneys-Fish nitrogenous wastes
• The type and quantity of an animal’s waste
products may greatly affect its water balance
• Among the most significant wastes are
nitrogenous breakdown products of proteins and
nucleic acids
• Fish typically produce toxic ammonia (NH3) rather
then less toxic compounds
• Abundance of water to dilute toxic materials
 A Balancing Act
• Relative concentrations of water and solutes
internally must be maintained within fairly narrow
limits
• Internal environment influenced by external
environment
 Osmoregulation & Excretion

• Osmoregulation
– Regulates solute concentrations and balances
the gain and loss of water

• Excretion
– Gets rid of nitrogenous metabolites and other
waste products
 Osmoregulation & Excretion

• Freshwater fishes in different environments

show adaptations that regulate uptake and
conservation of both water and solutes
 Osmoregulation & Excretion
• Osmoregulation is based largely on controlled
movement of solutes between internal fluids and
the external environment
 Osmosis and Osmolarity
• Cells require a balance between uptake and loss of
water
• Osmolarity, the solute concentration of a solution,
determines the movement of water across a
selectively permeable membrane
• If two solutions are isoosmotic, the movement of
water is equal in both directions
• If two solutions differ in osmolarity, the net flow of
water is from the hypoosmotic to the
hyperosmotic solution
 Osmotic Challenges
• Osmoconformers, consisting only of some marine
animals, are isoosmotic with their surroundings
and do not regulate their osmolarity
• Osmoregulators expend energy to control water
uptake and loss in a hyperosmotic or hypoosmotic
environment
 Hagfishes

• Osmoconformers
• Only vertebrate that is isotonic to seawater - much
like marine invertebrates
 Osmoregulators

• Aquatic vertebrates - gills are chief organs of

excretion/osmoregulation
• Kidneys first evolved as osmoregulatory organs in fishes to
remove water (freshwater) or conserve water (marine)
 Marine Animals

• Most marine vertebrates are osmoregulators

• Marine bony fishes are hypoosmotic to sea water
• They lose water by osmosis and gain salt by diffusion
and from food
• They balance water loss by drinking seawater and
excreting salts
• Kidney excretes creatinine, creatine and TMO
• Gills excretes ammonia & urea
(a) Osmoregulation in a marine fish
Gain of water Excretion Osmotic water
and salt ions of salt ions loss through gills
from food from gills and other parts
of body surface

Gain of water Excretion of salt ions and Key

and salt ions small amounts of water in
Water
from drinking scanty urine from kidneys
seawater Salt
 Freshwater
• A different form of osmoregulator
• Freshwater animals constantly take in water by
osmosis from their hypoosmotic environment
• lose salts by diffusion and maintain water balance
by excreting large amounts of dilute urine
• Salts lost by diffusion are replaced in foods and by
uptake across the gills
• N2 wastes are excreted more from gills than kidney
(6-10 times in carps)
• Kidney excretes creatinine & uric acid
• Gills excretes ammonia & urea
(b) Osmoregulation in a freshwater fish
Gain of water Uptake of Osmotic water
and some ions salt ions gain through
in food by gills gills and other
parts of body
surface

Key Excretion of salt ions and

large amounts of water in
Water
dilute urine from kidneys
Salt
Fig. 30.3
 Animal Inflow/Outflow Urine

Freshwater Does not drink water Large volume

fish. Lives in Salt in H2O in of urine
water less (active trans-
concentrated port by gills) Urine is less
than body concentrated
fluids; fish than body
tends to gain fluids
water, lose salt

Salt out
 Animal Inflow/Outflow Urine

Marine bony Drinks water Small volume

fish. Lives in Salt in H2O out of urine
water more
concentrated Urine is
than body slightly less
fluids; fish concentrated
tends to lose than body
water, gain salt fluids
Salt out (active
transport by gills)
Freshwater fishes
• Large volume of
urine, salts are
reabsorbed in the
distal tubules Rainbow trout
(Oncorrhynchus mykiss)

Frog (Rana)
Marine bony fishes
• Problem: gain salts from environment and tend to
lose water
• Lack distal tubule, smaller glomerulus, can adjust
amount of urine

Northern bluefin tuna (Thunnus thynnus)

• Marine cartilaginous fishes:
– Shark tissue contains a high concentration of urea
– To prevent urea from damaging other organic
molecules in the tissues, they have trimethyl amine
oxide (TMAO)
– Because of high solute concentration in tissue, water
enters the cells (sharks don’t drink)
– Produce concentrated urine
• Euryhaline organisms
like salmon:

– In sea, they drink sea

water and discharge salt
through their gills
– In freshwater, they stop
drinking and produce
large volumes of dilute
urine, gills take up salt
Reproduction in fishes
 Reproduction

sexual strategies:
females must be „careful‟ in mate selection due to cost
- energy investment in eggs
- migration, brooding

male investments in reproduction :

- advertisement, colors, tubercules, displays
- mate competition
- nest building, territorial defense, migration
- parental care, brood guarding
 Reproduction

bioenergetics: C = E + M + G + S + R

C – consumption
E – excretion
M – metabolism
G – growth
S – storage
R – reproduction
 Anatomy

hagfish, lamprey: single gonads

no ducts; release gametes into body cavity
 Anatomy

Teleosts: paired gonads

• external fertilization
• sperm released through separate opening of sperm duct
• ova maintained in continuous sac from ovary to oviduct
• exceptions: Salmonidae, Anguillidae, Galaxida
- these release eggs into body cavity when ripe
Anatomy
Testicular development

EL: empty lobules; LW: lobular wall; PSC:

primary spermatocytes; RC: residual cyst; SC:
stromal cell; SG: spermatogonia; SP: sperm; SSC:
secondary spermatocyte; ST: spermatid.
 Ovarian development

AF: atretic follicle; EV: early vitellogenic oocyte; EVE: early vesicular oocyte; GC: granulosa cells; IF: interstitial fluid;
LV: late vitellogenic oocyte; LVE: late vesicular oocyte; MC: mature cell; MV: mid vitellogenic oocyte; N: nucleus;
POC: primary oocyte; POF: post ovulatory follicle; TC: thecal cells; YG: yolk globule; YP: yolk plate; ZR: zona radiata.
 Anatomy

in general:
gametes produced only during spawning season
gonads reduced during non-reproductive season
 Timing and location of spawning

strategy:
avoid competition for spawning habitat
maximize access to food for offspring
minimize access to offspring by predators
 Timing and location of spawning

example: Lake Champlain

anadromous – salmon (SW to FW)
catadromous – eels (FW to SW)
fall spawners – lake trout, whitefish
spring spawners – smelt
littoral spawners – (near shore) sculpins, sunfishes, basses
stream spawners – suckers, darters, minnows, sturgeon
pelagic eggs – (attached to a floating object in water column)
e.g. burbot
 Reproduction

fecundity
egg size and number inversely related
egg number directly related to female size (within species)
related to food supply, competition
population-regulating mechanism
 Reproduction

fecundity
fractional spawners – produce eggs continuously,
spawn frequently
batch spawners – single reproductive season
release all eggs in a short period
 Reproduction

frequency of reproduction
semelparity - spawn and then die
- huge investment in egg production
e.g. Pacific salmon

iteroparity - repeated reproduction

allows compensation for a “bad” year
more common in more unstable environments
may not spawn every year (sturgeon)
 Reproductive strategies

fertilization
external except livebearers (elasmobranches, Poecilidae, etc)

mass spawning events (Clupeiformes, smelt, etc.)

several males to each female (Salmoniformes, lampreys)

several females to each male (Gobiidae)

single-pair matings (guppies)

 Reproductive strategies

non-guarders
- pelagic (broadcast) spawners
semi-buoyant eggs
high fecundity
egg and larval „migrations‟
 Reproductive strategies

non-guarders
- pelagic (broadcast) spawners
- benthic spawners
on coarse substrates (lake trout)
on vegetation (carp, perch)
on fine substrates (smelt)
 Reproductive strategies

non-guarders
- pelagic (broadcast) spawners
- benthic spawners
- brood hiders
build redd on coarse substrates (salmon, lamprey)

credit: Thomas B. Dunklin

 Reproductive strategies

non-guarders
- pelagic (broadcast) spawners
- benthic spawners
- brood hiders
build redd on coarse substrates (salmon, lamprey)
beach spawners (grunion)
use another species (bitterling)
 Reproductive strategies

guarders
- nest builders (largemouth bass)
 Reproductive strategies

guarders
- nest builders (largemouth bass)
rock and gravel (like a lentic redd - sunfishes)
plant material (sticklebacks)
holes, crevices, cavities (gobies, sculpin, blennies)
froth (bettas)
anemones (clown fish)
 Reproductive strategies

Bearers - carry eggs and/or fry with them

 Reproductive strategies

Bearers
- external bearers
transfer: Gasterosteidae, Sygnathidae (pipefishes, seahorses)
grade from attachment to skin, to open pouch,
to closed pouch
gill chambers, forehead
obstetrical catfish carry eggs on ventral surface
 Reproductive strategies

Bearers
- external bearers
mouth: males or females
some cichlids and bonytongues
 Alternative reproductive strategies
Hermaphroditism
synchronous (or simultaneous) hermaphrodites
Myctophiformes: (laternfishes) - several families
Atheriniformes: Aplocheilidae, Poeciliidae
Perciformes: Serranidae (sea basses, hamlets),
Labridae (wrasses), and others

"Egg-trading" in black hamlets Hypoplectrus nigricans (serranid)

 Alternative reproductive strategies
Hermaphroditism
consecutive (sequential) hermaphrodites
first male (protandrous) – less common

first female (protogynous)

Synbranchiformes (swamp eels – only freshwater example)
Perciformes: Serranidae, Maenidae, Labridae
from 100% female -> 100% male
from 100% female -> 50% male / 50% female
some do not pass thru a female stage ("primary males")
 Alternative reproductive strategies
parthenogenesis:
females produce diploid eggs, no sperm used
premeiotic endomitosis - mitotic division without cytokinesis

gynogenesis:
females produce diploid eggs, use sperm to stimulate development
male genome not used
congeneric species arne used for sperm

hybridogenesis: one genome from female in egg,

male genome discarded - then uses sperm to restore ploidy
- no crossing over
example: Poeciliopsis monacha-lucida
DEVELOPMENT
 Developmental stages
skate
(5 cm)
egg <0.5 mm - 10 cm
variable shape, attachments
variable buoyancy
water hardening
lake trout
(5 mm)

yellow perch egg mass

round goby
(0.5 mm)
Developmental stages

embryo - dependent on mother or yolk sac for food (free embryo)

Credit: Fly Anglers online

Susan Middleton & David Liittschwager

Developmental stages

embryo - dependent on mother or yolk sac for food (free embryo)

larvae - not fully functional, may look totally unlike adult
ends when axial skeleton is formed
Developmental stages

juvenile - small functional individual, immature

adult - reproductively mature
 Control of reproduction
• Advancement or retardation of spawning
• No maturation to conserve energy and
increase immunity
• Availability of fish at appropriate time/size
• Continuous production and marketing
 Control of Reproduction
• Manipulation of the reproductive cycle
• Induction of spawning
• Sex reversal
 Advancement

Continuous

&
Pineal Gland

Melatonin

GTHs
Possible
Sex Steroids Retardation
or Inhibition
Gonadal Development
 Induced reproduction

• Mammalian LH
• Mammalian HCG
• Pituitary extract
• Given in form of injections or LH pallets
implants
• Species specific dosage and timing
 Artificial fertilization
Gathering eggs and artificial fertilization

• Gathering eggs is made from

perfect sexually matured females.
At the time harvesting eggs, the
fish is hold by rolling up with a
towel around the head and tail, is
hold by head with left hand in
position with tail down so that by
gently palpation of abdomen,
eggs to fall in jar prepared
 Water Quality
Requirements/hygiene for
Aquaculture
 Parameters
• Temperature
• Dissolved Oxygen
• Alkalinity and Hardness
• pH
• Ammonia, Nitrite and Nitrate
• Miscellaneous
 Water Quality
• Water Quality (WQ) determines the ultimate
success or failure of an aquaculture operation
• The farmer must measure, record, and
manage WQ all through the growing season
• Water Quality parameters affect respiration,
feeding, metabolism, reproduction, and waste
removal from environment
 Temperature
• Affects the development and growth of fish more
than any other single factor
- Metabolic rates either increase or decrease with temp. change
- Each species has optimal growth and reproduction temp. ranges.
* Warm water species grow best above 21 C
* Cold water species grow best below 21 C
* Cool water species grow best at mid-range = 22 to 28
- Fish are ectothermic animals, therefore same as surrounding
water.
* Sudden change causes stress and even death
- Select species to culture according to the available water temp.
 Dissolved Oxygen (DO)
• Temperature, salinity and elevation affect
Dissolved Oxygen
• Salinity is the saltiness or dissolved salt content of a body of water
- As these three factors increase, DO at saturation decreases.
* Freshwater at sea level holds 9.2 ppm at 20C & 7.6 ppm at 30C
- Fish become more active and increase their metabolism as
temperature increases.
* Need more DO as temperature rises to grow muscle tissue.
* Minimum tolerable DO levels increase with a rise in temp.
With Rainbow trout, lethal level for DO is 1.6 ppm minimum at lower
temperatures and 2.5 ppm DO at higher temperatures.
 Dissolved Oxygen

• DO Ranges for cultured fish

0 to 2 ppm - small fish may survive a short exposure, but
lethal if exposure is prolonged. Lethal to larger fish.
2 to 5 ppm – most fish survive, but growth is slower if
prolonged; may be stressful; aeration devices are often used below
3ppm.
5 ppm to saturation – the desirable range for all.
 Dissolved Oxygen
• Biological Oxygen Demand (BOD)
- BOD is a measure of the oxygen used by all organisms in pond.
* Microbes (bacteria & fungi) use oxygen to decompose organics
(may use 1-3ppm DO in 24 hours).
* Phytoplankton respire at night to use oxygen (may use 5-15ppm
DO nightly).
* Fish respire day and night (may use 2-6ppm DO in 24 hours).
- DO falls at night, since all organisms are respiring; DO rises during the
day, since plants photosynthesize to use carbon dioxide and eliminate
oxygen (may gain 5-20ppm DO daily).
- Diffusion and wave/wind action add oxygen (may add 1-5ppm DO).
 Alkalinity & Hardness
• Two similar parameters, but still different
• Alkalinity
• The ability of the water to accept hydrogen ions, neutralizes pH.
• Consists of negatively charged bases – carbonates, bicarbonates, and
hydroxides.
• Expressed in equivalent concentrations of calcium carbonate.
• Carbonates and bicarbonates are sources of carbon for plants which is
used in photosynthesis to make sugars.
• Alkalinity offers a buffering system to reduce pH swings.
• An intermediate range of 20 to 80 ppm is recommended.
• E.g. striped bass prefer above 80 ppm alkalinity.
 Alkalinity & Hardness

• Hardness
- Refers to the concentration of divalent
cations (calcium, magnesium, and sodium).
- Also expressed as the calcium carbonate
equivalent concentration.
- The same carbonate rocks responsible for
most of the alkalinity are the main sources
of calcium and magnesium cations for
hardness.
- Hardness may be an index of potential
pond productivity.
* Minimum of 20 ppm
* Optimum around 100 ppm
 pH
• The negative logarithm of hydrogen-ions
concentration – a way to measure acidity
- Scale used is from 0 to 14, where lower number reflects higher acidity
and the higher number reflects higher alkalinity.
- Water with 4.5 pH or lower has no measurable alkalinity.
- Water with 8.3 pH or higher has no measurable acidity.
- Value of 7 is neutral, when donors of hydrogen ions = acceptors
- Recommended range for cultured fish is 6.5 to 9.0 pH
- Acid death point is around 4, alkaline death point is about 11 pH.
- Toxicity of ammonia to fish increases with an increase in pH.
 Ammonia, Nitrite and Nitrate
• Ammonia
- Ammonia gas from fish gills or decomposing organics dissolves in water.
Some of it reacts with the water to produce ammonia ions. The
remainder is present as un-ionized ammonia, which is acutely toxic to
aquatic life.
- The percentage of un-ionized in solution depends upon the pH and
temperature of the water. As both go higher so does the toxicity.
- Solubility decreases with rise in temp.
- Test kits normally used measure Total Ammonia Nitrogen (TAN),
therefore the fish culturist has to determine what % of total is toxic.
- Recommended that un-ionized ammonia should be < 0.02 ppm to
prevent stress and reduced growth.
- Lethal to catfish at about 0.4 ppm.
 Ammonia, Nitrite, & Nitrate
• Table lists the toxic percentage at different pH and temps.
 Ammonia, Nitrite and Nitrate

• Typical pond has bacteria, which in the presence of DO

converts (oxidizes) ammonia to the intermediate form of
nitrite and then to nitrate. Nitrite is more toxic to fish
than ammonia, however, nitrate is relatively nontoxic.
 Miscellaneous Water Quality Parameters

• Hydrogen Sulfide
- A poisonous gas produced by anaerobic decomposition of organics.
- If fish culturist smells “rotten egg” around the pond, aerate vigorously.
• Salinity and Chlorides
- Salinity is a measure of the total concentration of dissolved solids,
usually in parts per thousand (ppt). Anions (- charged) are chloride,
sulfate, bicarbonate and bromide. Cations (+ charged) are sodium,
magnesium, calcium, potassium, and strontium. Sodium and chloride are
the major solids.
- Freshwater = < 2 ppt; Brackish water = 2-16 ppt; Saltwater = 35 ppt.
 Miscellaneous

• Carbon Dioxide
- Consumed during photosynthesis
by plants and expired during
respiration by animals, plants (at
night) and bacteria in the pond.
- Levels > 20 ppm often harm the
fish, especially if DO is low.
- When added to the pond water by
respiration or diffusion, it forms a
weak carbonic acid, which lowers
pH. DO and pH follow the same
daily peaks.
 Miscellaneous
• Chlorine
- Harmful/Toxic to fish at > 0.03 ppm. City water may range from 4.0 to
8.0 ppm. Sodium thiosulfate can be used to neutralize the chlorine.
- May be used to disinfect equipment, tanks and nets at 10 ppm for 24
hours or 200 ppm for 30 to 60 minutes.
- Effectiveness is reduced by organic material such as mud, slime and plant
material.
- Sodium hypochlorite (HTH) is available at 15%, 50%, or 65% active. To
make a 200 ppm solution:
Add 2 oz. of 15% active HTH to 10.5 gal. of water,
Add 1 oz. of 50% active HTH to 18 gal. of water, or
Add 1 oz. of 65% active HTH to 23 gal. of water
 Miscellaneous
• Toxic Materials
- Some industrial and agricultural substances are toxic to fish,
such as heavy metals, herbicides and pesticides.
Examples of heavy metals are zinc, copper, cadmium, lead
and mercury.
- Minute amounts (5 parts per billion) of some toxic materials
are sufficient to be toxic to aquatic life.
 Hygiene of ponds/tanks
• De-odorase
• Removal of ammonia
• Extracted from plant Yucca shidigera
• Containing enzymes and glyco-comonents
such as sapogenin, smilogenin,chlorogenin
• Glyco-components selectively bind ammonia
• Producing non toxic nitrogenous compounds
which are used by bacteria
 Hygiene of ponds/tanks
• Chlorine dioxide
• Disinfectant in gas and liquid form
• Low toxicity
• Bactericide, viricide, slimicide, protocide
• Less corrosive
• Effective over broad range of pH (5-10)
• 150 – 250 ml per acre feet every 15 days
 Hygiene of ponds/tanks
• Aquazyn
• Blend of cultured bacteria, enzymes and
buffers
• Degradation of organic wastes from dead
phyto, zooplanktons, faeces and unconsumed
feed
• Increases average biomass and survival
• decreases biochemical oxygen demand and
pond bottom sludge
 Hygiene of ponds/tanks
• Aquazyn
• Improves FCR
• Increases nutrient availability for plankton
growth
• Non pathogenic, non toxic, biodegradable
 Hygiene of ponds/tanks
• Sokrena
• Liquid pond sanitizer
• 1 mg/L or 10 L of chemical per hectare
• Effective against broad range of bacteria, fungi
and viruses
• Most effective in alkaline environment
• Effective even in presence of organic matter
• Does not settle on pond bottom
 Hygiene of ponds/tanks
• Geolite
• Natural aggregate of inorganic salts i.e. silicon,
aluminium, calcium
• 10 kg/ hectare
• Improves growth and survival
• Water quality
• Neutralize pH of water and soil
• Decreases water turbidity
• Removes bad odor of water
Fish Diseases
Parasitic Diseases
 Numerous types of parasites
• Protozoans
• Trematodes
• Nematodes
• Cestodes
• Crustaceans
• Leeches
 Protozoa
• Ciliates
– Large protozoa up to 2mm in length that have cilia
(hair like organelles) covering their body at some
point, if not all, of their life cycle
• Flagellates
– Protozoa equipped with one or more whip-like
flagella used for propulsion
 Ciliates
• Largest group of protozoa
• Direct life cycle
• Common in pond-reared fish
• Easy to eliminate in aquarium due to
controlled conditions
• Easily transmitted through nets, hoses or
hands
 Ciliates
• Symptoms include
– Skin or gill irritation
• Rubbing and abnormal breathing
• Species
– Ichthyophthirius multifiliis
– Tetrahymena
– Apiosoma
– Epistylis
 Ichthyophthirius multifiliis
• Disease
– Ich or white spot disease
– Responsible for majority of fish fatalities
• Symptoms
– Small lesions on body or fins
– If gills only then no lesions but a thick mucus
 Ichthyophthirius multifiliis
• Identification
– Horseshoe shaped macronucleus
Ichthyophthirius multifiliis
 Ich Treatment
• Life cycle is temp dependent
– So raising the temp increases life cycle to days
instead of weeks
• Formalin
• Copper sulfate
• Treat for >3days after last spot gone
• vaccum the floor of tank to get the cysts
 Tetrahymena
– Living in organic debris
– Body surface of fish
• Treatment
– Same as Ich
 Apiosoma
• Location
– Gills, skin, or fins
• shape
– Vase like
– Oral cilia
• Treatment
– formaline, copper sulfate
Apiosoma
 Epistylis
• Stalked ciliate very similar to apiosoma
• More dangerous than apiosoma
– Greater concern than most due to proteolytic
enzymes secreted by the organism
– The enzyme breaks down proteins in the skin of
the fish making the fish susceptible to bacterial
infection
 Epistylis
• Treatment
– Salt
• .02% salt solution for extended period
• 3% salt dip
– More than one treatment required
 Flagellates
• External and internal
• species
– Spironucleus
– Ichthyobodo
– Myxozoa
– Microsporidia
 Spironucleus
• Location
– Intestinal tract
• Symptoms
– Extreme weight loss
– Distended abdomen
– Yellow mucous build up in intestines
 Spironucleus
• Diagnosis
– Intestinal prep observed at 200-400x
• Treatment
– Metronidazole bath
• 5mg/L
• Every other day for one week
 Ichthyobodo
• External Flagellate
• Large amounts of mucus
– Blue slime disease in catfish
• Location
– Gills, skin and fins
• Diagnosis
– Microscopic examination….
• Treatment
• formalin, copper sulfate
 Myxozoa
• Widespread in native or pond reared fish
– Most infections not bad
– But some may be serious in young fish
 Myxozoa
• Extremely abundant and diverse
• Speciation
– Based on spore shape and size
• Examination technique
– Preps of infected area
– Histologic sections of tissue
 Myxozoa
• Symptoms
– Vary depending on the organ affected
• Excess mucus productions
• White or yellowish nodules on target organs
• Treatment
– No remedies known
– Spores can survive >1year
• Disinfection necessary after removal of infected fish
 Microsporidia
• Intracellular parasites
– Require host tissue for reproduction
• Mode of transmission
– Ingesting spores from infected fish or food
• Symptoms
– Small tumors in various tissues
– Enlargement of hosts cells cause tumor like
masses
 Microsporidia
• Clinical signs
– Dependent on tissue infected
– from no lesions to dead
– In Serious cases, cysts enlarge to a point that
organs no longer function correctly
 Microsporidia
• Infections caused
– Pleistophora
• Infects skeletal muscle
• Cysts are observed
• Treatments
– None available
– Spores tough
• Can survive long periods
• Most environmental conditions it can handle
– Flushing of infected fish
– Disinfect the environment
Nematodes
 Capillaria

• common nematode seen in the intestines of

cichlids, cyprinids, gouramis, tetras.
• Treatment
• dewormers such as levamisole or
fenbendazole
 Eustrongylides
• can be found in muscle, “free” within the
body cavity, or encapsulated on the liver and
other organs, but they are found outside the
intestinal tract of fish
• Affected fish typically have bloated abdomens
• Treatment
• Only culling
 Camallanus
• infect the gastrointestinal tract of cichlids,
guppies, swordtails and other species of
freshwater fish.
• Usually, the first evidence of infection is a red,
worm-like animal protruding from the anus of
a fish.
• Treatment
• Dewormers
Crustacean Diseases
 Argulosis
• large ectoparasite and can move over the body
surface of the fish.
• Argulus puncture the skin and inject cytolytic toxin
through the oral sting to feed on the blood.
• The feeding site becomes a wound and
haemorrhagic, providing ready access to secondary
infection of other parasites, bacteria, virus and
fungi.
• Treatment
• 500 ppm of ammonium chloride, 410 ppm of
balsam, 10 ppm of DDT for 25 seconds dip
 Lernaeasis
The adult female is a specialized fish parasite,
worm like, which burrows into the fish flesh,
keeping its eggs cases protruding out of the fish
body.
• Treatment
• Only preventions
• 1% common salt eliminates larvae in 3 days,
250 ppm formalin for 30 to 60 minutes.
Ergasillus and salmincola
• These two parasites are responsible for huge
mortality of fishes in the culture systems.
• These two parasites are found attached to the
gill filaments and feed on blood and
epithelium.
• Later they may also be found on the fins and
body.
 Treatment
• Ergasilus can be treated successfully with a
combination of 0.5 ppm copper sulphate and
0.2 ppm ferric sulphate for 6 to 9 days.
• Salmincola can be controlled with 0.85%
calcium chloride, 0.2% copper sulphate
Fish Leeches
 Piscicola geometra
• Can affect any part
• Piercing skin and sucking
Blood
• Secondary infections
• Treatment
• Anaesthetize fish and
Manually remove
Fungal Diseases
 Class Oomycetes
• They produce a motile biflagellate spore (easy
dispersal)
• Also produce a thick-walled zoospore by the
fusion of two gametes
• usually identified as hyphae
 Saprolegniasis
• Four Orders, but most significant fish pathogens are
within Family Saprolegniaceae

• It is typically external, affecting skin and gills,

sometimes eggs
Saprolegniasis: hyphae
 Saprolegniasis
• the adult form is a mass of filaments known as
hyphae
• the mass is called a mycelium (looks like
cotton in the water)
• hyphae are unique in that they are non-
septate (no divisions)
• the asexual biflagellated zoospores are
thought to initiate most infections
 Saprolegniasis

• ubiquitous, most surface fresh waters, limited to no greater than

2.8 ppt salinity.
• Can live on dead or live matter, affect only fish which have been
compromised in some way:
1. suppression of immune system (unfavorable temps)
2. injury to skin (trauma)
3. spawning or precocious sexual maturity (thickened
epithelium = more mucus)
4. no seasonal (temperature) restrictions to infections with eggs
 Saprolegniasis
• Signs
• gray-white lesions on skin
• lesions start small and circular: spread
• can damage internal organs
• All fish susceptible
• Unfertilized eggs can be attacked by hyphae
(water hardening)
 Saprolegniasis
Treatment:
• Malachite green-topical (not approved)
• Bath: 1-2 mg malachite/litre (30 - 60 min.)

• Formalin: Bath: 0.15 to 0.25 mls/litre (60 min.)

approved but as effective
• Salt bath
• Potassium Permanganate
• Chloramine T
• Methylene blue
• Acetic acid (as a dip at 5% up to 1 minute)
Saprolegniasis
Saprolegniasis
Saprolegniasis

Fish eggs with Saprolegniasis

 Branchiomycosis
• This disease is commonly referred to as “gill rot”
• due to massive necrosis of gills
• Branchiomycosis sp. fungus invades gill blood vessels
• either B. sanguinis (only in gill blood vessels); carp,
goldfish
• B. demigrans (grows from blood vessels to tissue);
bass, pike, striped bass
Branchiomycosis
 Branchiomycosis
• Signs: sudden on-set, rapid course, high
mortality (within two days sometimes), overall
mort’s = 30-50%
• usually when temps above 20oC
• High organic loads, algae, high temps, high
density
• transmission: horizontal from other necrotic
gills (spores)
 Branchiomycosis
• Signs
• disease course so fast that fish are dead before any
signs, become sluggish
• later: necrotic patches on gills (much clubbing,
fusion of lamellae)
• hyperplasia of gill epithelium, fusion of lamellae,
massive necrosis
• Control: treatment ineffective due to rapid on-set;
strict hygiene, remove dead fish, don’t overfeed,
fertilize, crowd
 Ichthyophoniasis (Zygomycotina)
• caused by Ichthyophonus hoferi
• thick, fungus-like resting spores
• found in most cold water marine fish populations
• disease transmitted orally
• it is an obligate fish pathogen.
• life history varies from host to host
Ichythophoniasis
 Ichythophoniasis

• Life cycle complicated: produces large number of endospores

and resting spores in most internal organs

• Signs: hyphae are not visible externally

• gray-white lesions of organ, organ atrophy

• Control: disease transmitted orally

• Don’t feed infected fish to fish (feeding raw marine fish meal to
hatchery fish)
Ichythophoniasis
 Aspergillo mycosis

• Ubiquitous, involved in decay

• By products of degradation of feeds = aflatoxicosis
• mortalities of at least 20% of stock
• Signs: abdominal distension, darkening of color,
lethargy
• hyphae in liver, spleen, kidney, intestine, swim
bladder
Migration
Migration in Fish

• Migration is the movement of large number of

animals from one place to another for feeding,
reproduction or to escape weather extremes. When
large numbers of fishes come together and move
socially it is called shoaling.
• But sometimes migrating fishes exhibit high degree
of coordination in their movements and carry out
synchronized manoeuvres to produce different types
of shapes.
• This is called schooling, as seen in tunas and
sardines.
• Diadromons fishes – migrate between sea and
fresh water

– Anadromous – major part sea but fresh water during

breeding season e.g. Salmon and Hisla
– Catadromous – major part fresh water but sea during
breeding e.g. Anguilla (eel fish)
– Amphidramons – fresh water to sea and vice versa.
E.g. Gobies
• Potamodromons – migratory, confined to fresh
water eg. Carps and trouts, Mahaseer move up
stream along Himalaya rivers

• Oceanodromons – migratory, confined to sea

only eg. Tunnas, Mackerels
 Causes of migration in fishes
• Gametic migration (Spawning/breeding migration)

– Better survival and proper development of egg/larva

– Stop feeding prior to migration or reduced drastically
– Energy requirement (fat deposit) e.g. Chum salmon spp.
25,810 Ca and 28,390 cal by male and female respectively
• Alimental of Feeding Migration
– Due to shortage of food (Suitable/wintering/spawing)
– Better food facilities, better survival and fast growth
– Grow fast in size and mature and produce more eggs.

• Climate or Wintering Migration

– Due to inactive physical condition and low BMR
– Depends on fish condition and environment
– Achieved by hormonal and physiological changes
– Deposit mainly as fat deposits
– Among freshwater species (Grass carp) more to wintering
grounds.
• Osmoregulatory or Protective Migration
– Spawning, feeding and wintering migration can all be regards as
protection migration as they ensure further life of fish
– These migration are not cyclical
 Feeding or alimental migration
• Feeding or alimental migration takes place in fishes
for feeding.
• In high populations fishes exhaust food resources in
an area quickly and therefore must migrate
constantly in search of new feeding resources.
• Salmons, cods and sword fish constantly migrate for
food from one place to another in the sea.
 Spawning migration
• Spawning migration takes place in breeding
season in those fishes which have spawning
grounds far away from feeding places.
• Migratory fishes such as eels and salmons and
a large number of riverine fishes spawn in
tributaries of river in hills and migrate in large
number for laying eggs in these oxygen rich
waters.
• Juvenile migration
• involves larval stages of fishes which hatch in
spawning grounds and must migrate long
distances in order to reach the feeding
habitats of their parents.
• Recruitment migration
• takes place when large number of larvae
moves from nursery habitat to the habitat of
adults.
• which may sometimes be distinctly different.
• Adults of eels live in rivers in Europe and
America but their larval stages live and grown
in sea and migrate to reach rivers which may
take one to two years.
 Seasonal migration
• takes place in fishes that inhabit arctic areas
where in summer climate is conducive and
food abundant but as winter approaches
temperatures fall below zero and food
becomes scarce.
• fishes must migrate towards subtropical and
tropical areas to escape extremes of weather
conditions.
 TYPES OF MIGRATION IN FISHES

• Fishes live in two different types of aquatic

habitats, namely, freshwater and marine
habitats, which pose different osmotic
problems because of which it is difficult to
migrate from one type of habitat to another.
• some fishes species do migrate.
POTAMODROMOUS MIGRATION
• When fishes migrate from one freshwater habitat to
another freshwater habitat in search of food or for
spawning, it is called potamodromous migration.
• There are about 8,000 known species that migrate
within lakes and rivers, generally for food on daily
basis as the availability of food differs from place to
place and from season to season.
• Fishes also must migrate to lay their eggs in places
where oxygen concentration in water is more and
where there is abundance of food for juveniles when
they hatch from eggs.
 OCEANODROMOUS MIGRATION
• This migration is from sea water to sea water. There
are no barriers within the sea and fishes have
learned to migrate in order to take advantage of
favourable conditions wherever they occur.
• Thus there are about 12,000 marine species that
regularly migrate within sea water.
• Herrings, sardines, mackerels, cods, roaches and
tunas migrate in large numbers in search of food by
way of shoaling (migrating together socially but
without much coordination) or schooling (swimming
with high degree of coordination and synchronized
manoeuvres).
 DIADROMOUS MIGRATION
• When fishes can migrate from fresh water to
sea or from sea to fresh water, it is called
diadromous migration.
• There are about 120 species of fishes that are
capable of overcoming osmotic barriers and
migrate in these two different types of
habitats. This migration is of two types.
• Catadromous migration
• Anadromous migration
 Catadromous migration
• This type of migration involves movement of large
number of individuals from fresh water to sea water,
generally for spawning as happens in the case of
eels (Anguilla) inhabiting European and North
American rivers.
• Both European eel (Anguilla anguilla or Anguilla
vulgaris) and the American eel (Anguilla
rostrata) migrate from the continental rivers to
Sargasso Sea off Bermuda in south Atlantic for
spawning, crossing Atlantic Ocean during the journey
and covering a distance of about 5,600 km.
• The adult eels that inhabit rivers are about a metre
long, yellow in colour and spend 8-15 years feeding
and growing.
• Before migration the following changes take place in
their bodies:
• They deposit large amount of fat in their bodies
which serves as reserve food during the long journey
to Sargasso Sea.
• Colour changes from yellow to metallic silvery grey.
• Digestive tract shrinks and feeding stops.
• Eyes are enlarged and vision sharpens. Other sensory
organs also become sensitive.
• Skin becomes respiratory.
• Gonads get matured and enlarged.
• They become restless and develop strong urge to
migrate in groups.
 Anadromous migration
• Adults of anadromous fishes live and feed in ocean
waters but their spawning grounds lie in the
tributaries of rivers. Salmons, sturgeons, Hilsa and
lampreys are some of the marine fishes that
undertake anadromous migration to spawn in rivers.
• Atlantic salmon (Salmo salar) migrates to the North
American rivers for spawning while six species of
Pacific salmon(Onchorhynchus) migrate to various
rivers of Asian countries.
• Salmons living in sea are metallic silvery grey in
colour but before migration they turn reddish-brown
in colour. During fall, they enter rivers and swim
energetically against water currents
(contranatent), clearing all obstacles, including
waterfalls and reach tributaries in hilly areas where
they make a saucer-like pit in which female lays eggs
and male releases smelt over them.
• Eggs take 2-3 months to hatch in the following
spring, when the juvenile stage called Alvin emerges
out but remains within the nest, obtaining its
nourishment from the yolk sac attached to its belly.
• Alvin then transforms into Fry which feed on
planktons. Fries are denatant (they swim along with
water current) and feed and grow into fingerlings
which take the shape of adult fish. They change
into Smolt which congregate at the river mouth in
large numbers and then enter sea water in to
metamorphose into adult salmons.
 Behavioral and physiological
adaptations
• Fresh water

• In salt water…reverse of it
• The behavioral (drinking or not drinking) and
physiological changes a salmon must make when
moving from fresh water to salt water and vice
versa.
• young salmon on its seaward journey first
reaches the saline water at the mouth of its
home stream, it remains there for a period of
several days to weeks
• gradually moving into saltier water as it
acclimates.
• During acclimation
• it begins drinking the water.
• its kidneys start producing
• concentrated, low-volume urine,
• NaCl pumps in its gills now pumping NaCl out
of the blood and into the surrounding water.
• Likewise, when an adult salmon is ready to
spawn and reaches the mouth of its home
stream, it once again remains in the brackish
( = less concentrated than full-strength sea
water) water zone of the stream's mouth
• until it is able to reverse the changes it made
as a juvenile invading the ocean for the first
time.
Fish Behavior
• Fish are more intelligent than they appear.
• In many areas, such as memory, their cognitive
powers match or exceed those of ‘higher’
vertebrates including non-human primates.

• Fish hold the records for the relative brain

weights of vertebrates. Most vertebrate species
have similar brain-to-body weight ratios.
 Fixed Action Patterns
Fixed Action Patterns
• stereotypical innate behavior.
• The organism will carry it out almost no
matter what, even if it doesn’t seem
appropriate.
• These are all part of a category of behaviors
very important to survival and/or fitness.
 Fixed Action Patterns
Male three spined stickleback: attacks other
males with red bellies – attacks anything
red
 Innate behavior
• Ability to confront novel stimuli, learn about them
and adjust behavior is indicative of intelligence
and self awareness.
• Intelligence involves brain development, parental
investment or training
 Learning
• Change in behavior based on experience
Maturation is behavior change based largely on ability
due to development (eg. Use of tool)
paradise fish which avoid places where they have
experienced a single attack by a predator and continue to
do so for many months
• Habituation
Loss of responsiveness due to repetition
• Imprinting
Learning in a critical time period (tightly correlated with
innate behavior
Conditioning: Pavlov
Associating a stimulus with punishment or reward (can
also be trial and error) (visual experiments)
Associative learning/conditioning
• Associating one stimulus with another
• Pavlov: classical conditioning. Associating an
arbitrary stimulus with reward or punishment
• Operant conditioning: learning through trial and
error. BF Skinner’s experiments. This has formed
the basis for much animal training.
• Classical and operant conditioning often work
together
• Rainbow trout can be trained to press a bar to
get food, and they remember this three
months after last seeing the bar.
• Red Sea clownfish can recognize their mate 30
days after it was experimentally removed from
the home anemone.
• Channel catfish can remember the human
voice call announcing food five years after last
hearing that call
• Goldfish remember the colour of a tube
dispensing food one year after the last tube
presentation.
• Sockeye salmon still react to a light signal that
precedes food arrival up to eight months since
the last reinforcement.
• Some common rudd and European chub could
remember the person who trained them to feed
from the hand, even after a 6-month break.
• Crimson-spotted rainbowfish can learn how to
escape from a trawl by swimming through a small
hole in the center and they remember this
technique 11 months later.
 Reproductive behavior

• Sexual selection
– Courtship
– Female choice
– Male aggression
 Mating strategies

• Promiscuous
• Monogamous
• Polygamous: polygynous, polyandrous
 Fish Behaviors

• Migration
• Shoaling
• Feeding
• Aggression
• Resting
• Communication
 Migration
• Spatial orientation and mapping
– Migration: Piloting, orientation
(directional headings), navigation
(relative location)
– The role of learning in migration
– (magnetite, light, etc.)
– Fish orient themselves using landmarks and may
use mental maps based on multiple landmarks or
symbols.
– Fish behaviour in mazes reveals that they possess
spatial memory and visual discrimination.
 Fish Migration
• Fish migrations are usually round-trip
• Reasons for migration
– Food gathering
– Temperature adjustment
– Breeding
 Timing of migrations

– Annual
– Daily
– generational
 Classification of Fish Migration

• Diadromous – Travel between sea & fresh water

– Anadromous – most of life at sea, breed in fresh water

– Catadromous – most of life in fresh water, breed at sea
– Amphidromous – migrate between water types at some
stage other than breeding

• Potamodromous – Migrate within a fresh water system

• Ocenodromous – Migrate to different regions of the ocean
 Reasons for Migrations
• Take advantage of different habitats
– Feeding
– Protection
• Avoid adverse conditions
• Meet requirements for reproduction
 Orientation During Migration

• Orientation to gradients of temperature,

salinity, and chemicals
• Orientation by the sun
• Orientation to geomagnetic and geoelectric
fields
 Disadvantages of Migrations

• Expenditure of energy
– Most must store energy before migration
• Risk from predation
 Adjustments Required Due to
Migrations
• Adjusting physiologically to new water
conditions
– Temperature
– Light
– Water chemistry
• Many migratory species are now rapidly
declining due to changes caused by man
 Social behaviour
• Solitary
• Shoal - any group of fishes that remains
together for social reasons
• School - a polarized, synchronized shoal (has
coordinated, directed movements)
 Functions of Schooling Behavior

• Hydrodynamic efficiency

• Reduced predation risk

• Feeding

• Reproduction
 Functions of Schooling Behavior

• Hydrodynamic efficiency
– individuals obtain reduction in drag by following in
“slip-stream” of neighbors

– limited evidence in support of this

 Functions of Schooling Behavior
• Reduced predation risk
– creates patchy distribution of prey - large areas
with no prey
– once school is found, individual risk of being
captured is reduced by dilution
– confusion of prey by protean displays,
encirclement, other behaviors
 Functions of Schooling Behavior
• Feeding
– increases effective search space for the individual
(more eyes, separated by greater distance)
– coordinated movements to help break up schools
of prey - analogous to pack behavior in wolves - by
tunas, jacks
 Functions of Schooling Behavior
• Reproduction
– increases likelihood of finding a mate
– facilitates coordination of preparedness
(behavioral and pheromonal cues)
– facilitates arriving at right spawning site at right
time
 Fish Behavior & Communication
• Shoaling
• A social grouping of fish
• Occurs throughout life in about 25% of fish
species
• Half of all fish shoal at some time
• Benefits of Shoaling
• Gives a predator many moving targets
– Confuses predators
– Increases chances at the individual level
– Increases food finding ability
• Keeps potential mates in close proximity
 Fish Behavior & Communication
• Pods
• Tightly grouped school
• Move as a single unit (including making
quick turns)
• Makes the school appear like one large
organism
– Protection from predators
 Liabilities of Grouping Behavior
• Increased likelihood of disease & parasite
transmission
• Becoming more conspicuous to some
predators
– Harvested more easily by man
 Feeding Behavior
• Morphology is often a key to feeding behavior – many
fish have specialized habits
• Actual feeding may depend on what is available

• Optimal foraging – Take whatever is closest, as long as

it is suitable food
– Highest quality of food for the least amount of
effort
 Optimal Foraging
• All else being equal, take the largest prey
• Don’t choose prey that takes more energy
than it provides
• Be in a habitat that provides the type of
food you are looking for
 Risk Sensitive Foraging
• Foraging is sometimes restricted because of
undo risk
– It does not make sense to look for prey
where you will become the prey
– Must balance energy gain possibility with
risk of obtaining the energy
 Finding Food
• Visual detection
– Diurnal feeders
– Means being in the open in bright light
• Olfaction
– Common in bottom dwelling species
• Taste
 Agressive Behavior
• Direct charges
– Often includes biting
• Ritualistic displays
– Modified swimming
– Flaring gill covers
– Color changes
– Threatening movements
 Reasons for Aggressive Behavior

• Defense of territory
– Usually connected with reproduction
– Sometimes to keep food source
• Defense of brood
• Repelling competitors for mates
 Resting Behavior

• Inactive state
• Some fish spend a large part of the day
not doing anything
• Many species change color patterns
• Most fish rest on or near the substrate
• Many fish have a specified time of day
when resting takes place
• Some fish never rest (Sleep swimming?)
– Must keep moving (sharks)
 Communication
• Visual signals
• Auditory signals
• Chemical signals
• Electric signals
 Signals
• Visual Signals
• Most important communication signal
• Large variety of signals
– Different species use different “languages”
– Some cues are recognized between species
 How visual signals are produced
• Types of coloring
– Pigments
• Colored compounds
• Located in chromatophores
– In mostly in skin, but also in eyes & organs
• Controlled by hormones & nerves
– Structural colors
• Reflection of light
 Kinds of Pigments in Fish
• Carotenoid pigments
– Bright reds & yellow
– Green when they overly blue structural color
• Melanins
– Dark red, brown, black
• Purines (guanine)
– Colorless crystals responsible for some
structural colors
 Purpose of Color Patterns
• Thermoregulation
– Probably not very significant
• Intraspecific communication
• Evasion of predators
 Auditory Signals
• Most fish produce sounds
• Uses for sound
– Courtship singing
– Territorial defense
– Signaling shoal
 Sound Production
• Stridulation
– Rubbing hard surfaces together
– Low frequency sounds
• Vibration of swimbladder
– Can give loud croaking
• Incidental to other activities
 Chemical Signals
• Pheromones released into the water
– Reproductive cues
– Recognition
• Schreckstoff = fear scents
– Predator avoidance
– Produced in epidermal cells
 Electrical Signals
• Muscle contractions give off a weak
-Some fish have electric producing organs
– Used to locate prey or conspecifics
 Behavior in Aquarium
• Loss of Appetite
• Difficulty Swimming
• Hovering Near Tank Surface
• Fish Swimming Quickly
• Fighting Among Tank Mates
• Rearranging Tank Objects
• Fin Nipping
• Fish Are Hiding