Publication

420-256
2002

Understanding Fish Nutrition,

Feeds, and Feeding
Steven Craig and Louis A. Helfrich*

Introduction

Protein

Good nutrition in animal production systems is essential to

economically produce a healthy, high quality product. In
fish farming, nutrition is critical because feed represents
40-50% of the production costs. Fish nutrition has
advanced dramatically in recent years with the development of new, balanced commercial diets that promote optimal fish growth and health. The development of new
species-specific diet formulations supports the aquaculture
(fish farming) industry as it expands to satisfy increasing
demand for affordable, safe, and high-quality fish and
seafood products.

Because protein is the most expensive part of fish feed, it is

important to accurately determine the protein requirements
for each species and size of cultured fish. Proteins are
formed by linkages of individual amino acids. Although
over 200 amino acids occur in nature, only about 20 amino
acids are common. Of these, 10 are essential (indispensable) amino acids that cannot be synthesized by fish. The
10 essential amino acids that must be supplied by the diet
are: methionine, arginine, threonine, tryptophan, histidine,
isoleucine, lysine, leucine, valine and phenylalanine. Of
these, lysine and methionine are often the first limiting
amino acids. Fish feeds prepared with plant (soybean
meal) protein typically are low in methionine; therefore,
extra methionine must be added to soybean-meal based
diets in order to promote optimal growth and health. It is
important to know and match the protein requirements and
the amino acid requirements of each fish species reared.

P r epar ed (ar tificial)

Diets
Prepared or artificial diets may be either complete or supplemental. Complete diets supply all the ingredients (protein, carbohydrates, fats, vitamins, and minerals) necessary
for the optimal growth and health of the fish. Most fish
farmers use complete diets, those containing all the
required protein (18-50%), lipid (10-25%), carbohydrate
(15-20%), ash (< 8.5%), phosphorus (< 1.5%), water (<
10%), and trace amounts of vitamins, and minerals. When
fish are reared in high density indoor systems or confined
in cages and cannot forage freely on natural feeds, they
must be provided a complete diet.

Protein levels in aquaculture feeds generally average 1820% for marine shrimp, 28-32% for catfish, 32-38% for
tilapia, 38-42% for hybrid striped bass. Protein requirements usually are lower for herbivorous fish (plant eating)
and omnivorous fish (plant-animal eaters) than they are for
carnivorous (flesh-eating) fish, and are higher for fish
reared in high density (recirculating aquaculture) than low
density (pond aquaculture) systems.

In contrast, supplemental (incomplete, partial) diets are

intended only to help support the natural food (insects,
algae, small fish) normally available to fish in ponds or
outdoor raceways. Supplemental diets do not contain a full
complement of vitamins or minerals, but are used to help
fortify the naturally available diet with extra protein, carbohydrate and/or lipid.

Protein requirements generally are higher for smaller fish.

As fish grow larger, their protein requirements usually
decrease. Protein requirements also vary with rearing environment, water temperature and water quality, as well as
the genetic composition and feeding rates of the fish.
Protein is used for fish growth if adequate levels of fats and
carbohydrates are present in the diet. If not, protein may
be used for energy and life support rather than growth.

Fish, especially when reared in high densities, require a

high-quality, nutritionally complete, balanced diet to grow
rapidly and remain healthy.

Proteins are composed of carbon (50%), nitrogen (16%),

oxygen (21.5%), and hydrogen (6.5%). Fish are capable of
using a high protein diet, but as much as 65% of the protein

*Extension Specialists, Virginia-Maryland College of Veterinary Medicine, and Department of Fisheries and Wildlife Sciences, Virginia Tech, respectively

VIRGINIA POLYTECHNIC INSTITUTE

AND STATE UNIVERSITY

Virginia Cooperative Extension programs and employment are open to all, regardless of race, color, religion, sex, age, veteran status,
national origin, disability, or political affiliation. An equal opportunity/affirmative action employer. Issued in furtherance of Cooperative
Extension work, Virginia Polytechnic Institute and State University, Virginia State University, and the U.S. Department
of Agriculture cooperating. J. David Barrett, Director, Virginia Cooperative Extension, Virginia Tech, Blacksburg;
Lorenza W. Lyons, Administrator, 1890 Extension Program, Virginia State, Petersburg.
VT/028/0402//420256

VIRGINIA STATE UNIVERSITY

may be lost to the environment. Most nitrogen is excreted

as ammonia (NH3) by the gills of fish, and only 10% is lost
as solid wastes. Accelerated eutrophication (nutrient
enrichment) of surface waters due to excess nitrogen from
fish farm effluents is a major water quality concern of fish
farmers. Effective feeding and waste management practices are essential to protect downstream water quality.

Lipids (fats)
Lipids (fats) are high-energy nutrients that can be utilized
to partially spare (substitute for) protein in aquaculture
feeds. Lipids supply about twice the energy as proteins and
carbohydrates. Lipids typically comprise about 15% of fish
diets, supply essential fatty acids (EFA) and serve as transporters for fat-soluble vitamins.
A recent trend in fish feeds is to use higher levels of lipids
in the diet. Although increasing dietary lipids can help
reduce the high costs of diets by partially sparing protein in
the feed, problems such as excessive fat deposition in the
liver can decrease the health and market quality of fish.
Simple lipids include fatty acids and triacylglycerols. Fish
typically require fatty acids of the omega 3 and 6 (n-3 and
n-6) families. Fatty acids can be: a) saturated fatty acids
(SFA, no double bonds), b) polyunsaturated fatty acids
(PUFA, >2 double bonds), or c) highly unsaturated fatty
acids (HUFA; > 4 double bonds). Marine fish oils are naturally high (>30%) in omega 3 HUFA, and are excellent
sources of lipids for the manufacture of fish diets. Lipids
from these marine oils also can have beneficial effects on
human cardiovascular health.
Marine fish typically require n-3 HUFA for optimal growth
and health, usually in quantities ranging from 0.5-2.0% of
dry diet. The two major EFA of this group are eicosapentaenoic acid (EPA: 20:5n-3) and docosahexaenoic acid
(DHA:22:6n-3). Freshwater fish do not require the long
chain HUFA, but often require an 18 carbon n-3 fatty acid,
linolenic acid (18:3-n-3), in quantities ranging from 0.5 to
1.5% of dry diet. This fatty acid cannot be produced by
freshwater fish and must be supplied in the diet. Many
freshwater fish can take this fatty acid, and through enzyme
systems elongate (add carbon atoms) to the hydrocarbon
chain, and then further desaturate (add double bonds) to
this longer hydrocarbon chain. Through these enzyme systems, freshwater fish can manufacture the longer chain n-3
HUFA, EPA and DHA, which are necessary for other metabolic functions and as cellular membrane components.
Marine fish typically do not possess these elongation and
desaturation enzyme systems, and require long chain n-3
HUFA in their diets. Other fish species, such as tilapia,
require fatty acids of the n-6 family, while still others, such
as carp or eels, require a combination of n-3 and n-6 fatty
acids

activity during feed manufacturing. Dietary starches are

useful in the extrusion manufacture of floating feeds.
Cooking starch during the extrusion process makes it more
biologically available to fish.
In fish, carbohydrates are stored as glycogen that can be
mobilized to satisfy energy demands. They are a major
energy source for mammals, but are not used efficiently by
fish. For example, mammals can extract about 4 kcal of
energy from 1 gram of carbohydrate, whereas fish can only
extract about 1.6 kcal from the same amount of carbohydrate. Up to about 20% of dietary carbohydrates can be
used by fish.

Vitamins
Vitamins are organic compounds necessary in the diet for
normal fish growth and health. They often are not synthesized by fish, and must be supplied in the diet.
The two groups of vitamins are water-soluble and fat-soluble. Water-soluble vitamins include: the B vitamins,
choline, inositol, folic acid, pantothenic acid , biotin and
ascorbic acid (vitamin C). Of these, vitamin C probably is
the most important because it is a powerful antioxidant and
helps the immune system in fish.
The fat-soluble vitamins include A vitamins, retinols
(responsible for vision); the D vitamins, cholecaciferols
(bone integrity); E vitamins, the tocopherols (antioxidants);
and K vitamins such as menadione (blood clotting, skin
integrity). Of these, vitamin E receives the most attention
for its important role as an antioxidant. Deficiency of each
vitamin has certain specific symptoms, but reduced growth
is the most common symptom of any vitamin deficiency.
Scoliosis (bent backbone symptom) and dark coloration
may result from deficiencies of ascorbic acid and folic acid
vitamins, respectively.

Minerals
Minerals are inorganic elements necessary in the diet for
normal body functions. They can be divided into two
groups (macro-minerals and micro-minerals) based on the
quantity required in the diet and the amount present in fish.
Common macro-minerals are sodium, chloride, potassium
and phosphorous. These minerals regulate osmotic balance
and aid in bone formation and integrity.
Micro-minerals (trace minerals) are required in small
amounts as components in enzyme and hormone systems.
Common trace minerals are copper, chromium, iodine, zinc
and selenium. Fish can absorb many minerals directly
from the water through their gills and skin, allowing them
to compensate to some extent for mineral deficiencies in
their diet.

Carbohydrates

Energy and Protein

Carbohydrates (starches and sugars) are the most economical and inexpensive sources of energy for fish diets.
Although not essential, carbohydrates are included in aquaculture diets to reduce feed costs and for their binding

Dietary nutrients are essential for the construction of living

tissues. They also are a source of stored energy for fish
digestion, absorption, growth, reproduction and the other
life processes. The nutritional value of a dietary ingredient

is in part dependant on its ability to supply energy.

Physiological fuel values are used to calculate and balance
available energy values in prepared diets. They typically
average 4, 4, and 9 kcal/g for protein, carbohydrate and
lipid, respectively.
To create an optimum diet, the ratio of protein to energy
must be determined separately for each fish species.
Excess energy relative to protein content in the diet may
result in high lipid deposition. Because fish feed to meet
their energy requirements, diets with excessive energy levels may result in decreased feed intake and reduced weight
gain. Similarly, a diet with inadequate energy content can
result in reduced weight gain because the fish cannot eat
enough feed to satisfy their energy requirements for
growth. Properly formulated prepared feeds have a wellbalanced energy to protein ratio.

Feed Types
Commercial fish diets are manufactured as either extruded
(floating or buoyant) or pressure-pelleted (sinking) feeds.
Both floating or sinking feed can produce satisfactory
growth, but some fish species prefer floating, others sinking. Shrimp, for example, will not accept a floating feed,
but most fish species can be trained to accept a floating
pellet.
Extruded feeds are more expensive due to the higher manufacturing costs. Usually, it is advantageous to feed a floating (extruded) feed, because the farmer can directly
observe the feeding intensity of his fish and adjust feeding
rates accordingly. Determining whether feeding rates are
too low or too high is important in maximizing fish growth
and feed use efficiency.
Feed is available in a variety of sizes ranging from fine
crumbles for small fish to large (1/2 inch or larger) pellets.
The pellet size should be approximately 20-30% of the
size of the fish species mouth gape. Feeding too small a
pellet results in inefficient feeding because more energy is
used in finding and eating more pellets. Conversely, pellets that are too large will depress feeding and, in the
extreme, cause choking. Select the largest sized feed the
fish will actively eat.

Feeding Rate,
Frequency, and
Timing
Feeding rates and frequencies are in part a function of fish
size. Small larval fish and fry need to be fed a high protein
diet frequently and usually in excess. Small fish have a
high energy demand and must eat nearly continuously and
be fed almost hourly. Feeding small fish in excess is not as
much of a problem as overfeeding larger fish because small
fish require only a small amount of feed relative to the volume of water in the culture system.
As fish grow, feeding rates and frequencies should be lowered, and protein content reduced. However, rather than

switching to a lower protein diet, feeding less allows the

grower to use the same feed (protein level) throughout the
grow-out period, thereby simplifying feed inventory and
storage.
Feeding fish is labor-intensive and expensive. Feeding
frequency is dependent on labor availability, farm size, and
the fish species and sizes grown. Large catfish farms with
many ponds usually feed only once per day because of time
and labor limitations, while smaller farms may feed twice
per day. Generally, growth and feed conversion increase
with feeding frequency. In indoor, intensive fish culture
systems, fish may be fed as many as 5 times per day in
order to maximize growth at optimum temperatures.
Many factors affect the feeding rates of fish. These include
time of day, season, water temperature, dissolved oxygen
levels, and other water quality variables. For example,
feeding fish grown in ponds early in the morning when the
lowest dissolved oxygen levels occur is not advisable. In
contrast, in recirculating aquaculture systems where oxygen
is continuously supplied, fish can be fed at nearly any time.
During the winter and at low water temperatures, feeding
rates of warmwater fishes in ponds decline and feeding
rates should decrease proportionally.
Feed acceptability, palatability and digestibility vary with
the ingredients and feed quality. Fish farmers pay careful
attention to feeding activity in order to help determine feed
acceptance, calculate feed conversion ratios and feed efficiencies, monitor feed costs, and track feed demand
throughout the year.
Published feeding rate tables are available for most commonly cultured fish species. Farmers can calculate optimum feeding rates based on the average size in length or
weight and the number of fish in the tank, raceway, or pond
(see Hinshaw 1999, and Robinson et al. 1998). Farmed
fish typically are fed 1-4% of their body weight per day.

Automatic Feeders
Fish can be fed by hand, by automatic feeders, and by
demand feeders. Many fish farmers like to hand feed their
fish each day to assure that the fish are healthy, feeding
vigorously, and exhibiting no problems. Large catfish
farms often drive feed trucks with compressed air blowers
to distribute (toss) feed uniformly throughout the pond.
There are a variety of automatic (timed) feeders ranging in
design from belt feeders that work on wind-up springs, to
electric vibrating feeders, to timed feeders that can be programmed to feed hourly and for extended periods.
Demand feeders do not require electricity or batteries.
They usually are suspended above fish tanks and raceways
and work by allowing the fish to trigger feed release by
striking a moving rod that extends into the water.
Whenever a fish strikes the trigger, a small amount of feed
is released into the tank. Automatic and demand feeders
save time, labor and money, but at the expense of the vigilance that comes with hand feeding. Some growers use
night lights and bug zappers to attract and kill flying
insects and bugs to provide a supplemental source of natural food for their fish.

Feed Conversion
and Efficiency
Calculations:
Because feed is expensive, feed conversion ratio (FCR) or
feed efficiency (FE) are important calculations for the
grower. They can be used to determine if feed is being
used as efficiently as possible.
FCR is calculated as the weight of the feed fed to the fish
divided by the weight of fish growth. For example, if fish
are fed 10 pounds of feed and then exhibit a 5 pound
weight gain, the FCR is 10/ 5 = 2.0. FCRs of 1.5-2.0 are
considered good growth for most species.
FE is simply the reciprocal of FCRs (1/FCR). In the
example above, the FE is 5/10 = 50%. Or if fish are fed 12
pounds of feed and exhibit a 4 pound weight gain, the FE =
4/12 = 30%. FEs greater than 50% are considered good
growth.
Fish are not completely efficient (FEs of 100 %, FCRs of
1.0). When fed 5 pounds of feed, fish cannot exhibit 5
pounds of growth because they must use some of the energy in feed for metabolic heat, digestive processing, respiration, nerve impulses, salt balance, swimming, and other living activities. Feed conversion ratios will vary among
species, sizes and activity levels of fish, environmental
parameters and the culture system used.

such as disease or water quality deterioration in the fish

growing system. Relatively few therapeutic drugs are
approved for fish by FDA (see Helfrich and Smith 2001),
but some medicated feeds for sick fish are available.
Although using medicated feeds is one of the easiest ways
to treat fish, they must be used early and quickly because
sick fish frequently will stop feeding.

Managing Fish
Wastes
The most important rule in fish nutrition is to avoid overfeeding. Overfeeding is a waste of expensive feed. It also
results in water pollution, low dissolved oxygen levels,
increased biological oxygen demand, and increased bacterial loads. Usually, fish should be fed only the amount of
feed that they can consume quickly (less than 25 minutes).
Many growers use floating (extruded) feeds in order to
observe feeding activity and to help judge if more or less
feed should be fed.
Even with careful management, some feed ends up as
waste. For example, out of 100 units of feed fed to fish,
typically about 10 units of feed are uneaten (wasted) and
10 units of solid and 30 units of liquid waste (50% total
wastes) are produced by fish. Of the remaining feed, about
25% is used for growth and another 25% is used for metabolism (heat energy for life processes). These numbers may
vary greatly with species, sizes, activity, water temperature,
and other environmental conditions.

Feed Care and

Storage

Useful References

Commercial fish feed is usually purchased by large farms

as bulk feed in truckloads and stored in outside bins.
Smaller farms often buy prepared feed in 50-pound bags.
Bag feed should be kept out of direct sunlight and as cool
as possible. Vitamins, proteins, and lipids are especially
heat sensitive, and can be readily denatured by high storage
temperatures. High moisture stimulates mold growth and
feed decomposition. Avoid unnecessary handling and damage to the feed bags which may break the pellets and create
fines which may not be consumed by fish.

Fish Kills: Their Causes and Prevention. 2001. Helfrich L.,

and S. Smith. Viginia Cooperative Extension Service
Publication 420- 252. Website:
http://www.ext.vt.edu/pubs/fisheries/420-252/420-252.html

Feed should not be stored longer than 90 to 100 days, and

should be inventoried regularly. Bags should not be
stacked higher than 10 at a time. Older feed should be used
first, and all feed should be regularly inspected for mold
prior to feeding. All moldy feed should be discarded
immediately. Mice, rats, roaches and other pests should be
strictly controlled in the feed storage area, because they
consume and contaminate feed and transmit diseases.

Medicated Feeds
When fish reduce or stop feeding, it is a signal to look for
problems. Off-feed behavior is the first signal of trouble

Food Intake in Fish. 2001. Houlihan, D., Bouiard, T. and

Jobling, M., eds. Iowa State University Press. Blackwell
Science Ltd. 418 pp.

Feeding Catfish in Commercial Ponds. 1998. E. Robinson,

M. Li, and M. Brunson. Southern Regional Aquaculture
Center, Fact Sheet # 181. Web Site:
http://www.msstate.edu/dept/srac/fslist.htm
Nutrient Requirements of Fish. 1993. Committee on
Animal Nutrition. National Research Council. National
Academy Press. Washington D.C. 114 pp.
Nutrition and Feeding of Fish. 1989. Tom Lovell. Van
Nostrand Reinhold, New York. 260 pp.
Principles of Warmwater Aquaculture. 1979. Robert R.
Stickney. John Wiley and Sons, New York. 375 pp.
Standard Methods for the Nutrition and Feeding of Farmed
Fish and Shrimp. 1990. Albert G.J. Tacon. Volume 1: The
Essential Nutrients. Volume 2:Nutrient Sources and

SRAC Publication No. 700

July 1999

Zooplankton Succession and Larval Fish

Culture in Freshwater Ponds
Gerald M. Ludwig*
Larval fish culture is one of the
riskiest phases of freshwater fish
culture, but it can be one of the
most profitable. Special planning
is required to overcome the risk of
high mortality during fry culture.
Producers must have a dependable larvae supply, a facility
appropriate for fry and fingerlings, the right size fry, the right
kinds and quantity of food, and
fry weaned from natural to prepared foods. They must also take
special care in handling fish and
preparing the pond. This fact
sheet concentrates on the relationship between fry size and the
types and sizes of zooplankton
found in culture ponds.
Zooplankton is required as a first
food for many cultured fish; for
others it contributes to faster
growth and higher survival.

Pond size
Ponds for fry culture and small
fingerling production should be
smaller than grow-out ponds.
Ponds from 0.1 to 3 acres are ideal
because they are easier to harvest
and will produce more natural
food per unit area. There is a
higher ratio of pond bottom area
to water volume in small ponds
than in large ponds, which
*Stuttgart National Aquaculture Research
Center

increases the availability of fertilizing nutrients and resting zooplankton eggs. Increased shoreline
to water volume increases the
number of small insects being
blown into the pond, and they
may be a significant source of
food for fingerlings. However,
ponds with lots of shoreline may
have more problems with predaceous wading birds.
Using many small ponds rather
than a few large ponds may
ensure that at least some fingerlings get to market. Smaller ponds
allow the farmer to more easily
control the size of the fish by
manipulating nutrient (either fertilizer or feed) input. Small ponds
also allow the farmer to more easily determine fish size and estimate survival rates because it is
easier to locate the fish. With
many small ponds instead of a
few large ones, farmers can grow
fingerlings of different sizes for
various markets. Spreading different sizes of fingerlings among different ponds also helps minimize
cannibalism. Farmers can rotate
the harvest among many small
ponds rather than harvesting the
same pond over and over; this
reduces stress.
Of course, the benefits of smaller
ponds must be balanced against
the increased costs and decreased
pond area per acre of land that
result when small ponds are used.

Fry size
Tiny fry eat only tiny prey, but
tiny fry are preyed upon by many
creatures bigger than they are. It
is important to know the size of
the fry you are stocking and to
make sure that the pond you are
putting them into contains plankton of the size that will be their
prey and is also void of creatures
that will prey on the fry.
The total length of cultured fish
fry (Table 1) when they hatch
varies from 2 mm for sunshine
and white bass to more than 15
mm for muskellunge. In most
cases, fry are a few millimeters
longer than the values in Table 1
when they are stocked into ponds.
Suggested stocking times in the
table are based upon the size of
the fry and the sizes and types of
zooplankton that show up at different times in ponds that have
been filled with well water and
fertilized. It is safer to stock earlier
than the time listed than to stock
later. Stocking later increases the
chance that predaceous zooplankton or insects will be present.

Plankton types and sizes

Pond plankton is composed of
tiny plants called phytoplankton
and animals called zooplankton,
as well as organisms that are not
easily classified into those two
groups (such as protozoans and

Table 1. Total lengths of fry for commonly cultured cool and warm water fish.
Common name
Sunshine bass
White bass
Black crappie
White crappie
Goldfish
Fathead minnow
Rosy-red minnow
Sauger
Golden shiner
Common carp
Yellow perch
Largemouth bass
Walleye
Grass carp
Silver carp
Bighead carp
Striped bass
Palmetto bass
Paddlefish
Spotted sucker
White sucker
Shovelnose sturgeon
Channel catfish
Muskellunge

Scientific name
Morone chrysops X M. saxatilis
M. chrysops
Pomoxis nigromaculatus
P. annularis
Carassius auratus
Pimephales promelas
P. promelas
Stizostedion canadense
Notemigonus chrysoleucas
Cyprinus carpio
Perca flavescens
Micropterus salmoides
Stizostedion vitreum
Ctenopharyngdon idella
Hypophthalmichthys molitrix
H. nobilis
M. saxatilis
M. saxatilis X M. chrysops
Polyodon spathula
Minytrema melanops
Catostomus commersoni
Scaphirhynchus platorynchus
Ictalurus punctatus
Esox masquinongy

Fry size (mm)*

2-6
3-4
3-5
3-5
3-5
4-6
4-6
4-6
4-7
5-7
5-7
6-7
6-9
6-9
6-9
7-8
7-10
7-10
8-10
8-10
8-10
>10
10-12
11-15

When to stock fry**

5
5
5
5
5
5
5
5
5
5
5
7
7
7
7
7
10
10
10
11
11
12
13
14

*25.4 mm equals1 inch

**Estimated number of days after pond starts filling at 21 to 27o C (70 to 80 o F)
Sources of fry sizes for Table 1:
Becker, G. 1983. Fishes of Wisconsin. University of Wisconsin Press. Madison, Wisconsin.
Carlander, K.D. 1977. Handbook of freshwater fishery biology. Vol. 2. The Iowa State University Press. Ames, Iowa.
Ludwig, G.M. 1996. Tank culture of sunshine bass Morone chrysops X M. saxatilis fry with freshwater rotifers Brachionus calyciflorus and
salmon starter meal as first food sources. Journal of the World Aquaculture Society 25:337-341.
Halver, J., ed., Horvath, L., G. Tamas and I. Tolg. 1984. Special methods in pond fish husbandry. Academiai Kiado, Budapest, Hungary, and
Halver Corporation, Seattle, Washington.
Huq, M.F. 1965. The effect of crowding on the growth of fry of channel catfish Ictalurus punctatus (Rafinesque). Scientific Researches
(Dacca, Pakistan). 2:112- 117.

bacteria). Planktonic organisms

are suspended in the water and
are so small that even slight currents move them about. Fish fry
eat zooplankton, phytoplankton,
and tiny plants and animals
attached to objects on the pond
bottom.
Most fish fry eat three main types
of zooplanktonrotifers, copepods and cladocerans. For the
tiniest fish fry, such as the newly
hatched fry of sunshine bass or
white bass, small rotifers may be
the only zooplankton small
enough to eat. For larger fry, the

smallest rotifers may not provide

enough nutrients to make chasing
and ingesting them worth the
effort. Copepod nauplii, which are
just-hatched copepods, are important first foods for larval fish, too.
Protozoans may also be eaten, but
little is known about their contribution to fry diets.
In general, the smallest of the
main zooplankton groups are
rotifers (Fig. 1). Body lengths of
rotifer species vary from 0.04 to
2.5 mm. This diverse group of animals obtained their name from
their wheel organ, a ring of cilia

that appears to rotate around the

mouth. They are often the earliest
visible zooplankton to appear in
ponds, hatching almost immediately after ponds are filled.
Rotifers reach maturity 2 to 8 days
after hatching and some species
can increase in number very
rapidly. Rotifers that show up
later in ponds, when larger zooplankton are present, are usually
much larger than the first that
appeared. The first rotifers hatch
from resting eggs that survived
on the pond bottom during
inclement weather or while the

Figure 1. Common zooplankton in fish culture ponds. (Drawings from

Taxonomic Keys to the Common Animals of the North Central States,
1982. Burgess Publishing Co. Minneapolis, Minnesota, used with permission from James C. Underhill.)
pond bottom was dry. Most of
them hatch into females that
reproduce asexually until pond
conditions become harsh. Then
sexual reproduction occurs and
resting eggs are again produced.
Copepod nauplii are the next
largest zooplankton to appear
after ponds are filled and fertilized (Fig. 1). They hatch from resting eggs that were dormant on the
pond bottom. Resting eggs are
produced in the fall by sexual
reproduction. At other times
reproduction may be sexual or
asexual. After hatching, the young
copepods grow by shedding their
exoskeletons up to 12 times in
stages called instars. They reach
maturity about 18 days after
ponds are filled. As they grow
they provide larger and larger
food for larval fish. The largest
freshwater copepods may reach 2
to 3 mm. Only large fry, such as
those of channel catfish, have
mouths big enough to eat adult
copepods initially.
Although copepods may be prey
for larger fish fry, sometimes the
roles are reversed. Introducing
small fish fry into a pond full of
large copepods can be disastrous.
One group of copepods, the

cyclopoids, is predaceous. They

feed on smaller zooplankton and
even on fish fry. When cyclopoid
copepods are prevalent, they may
eat all the fish fry stocked.
Cladocerans, often called water
fleas because of their shape and
hop-sink type of locomotion,
are the third major group of zooplankton found in freshwater
ponds (Fig. 1). Larger fry and
even adults of some fish species
often selectively prey on these
crustaceans. Cladocerans 2 to 3
mm long are commonly found in
culture ponds several weeks after
the ponds are filled. Often only
female cladocerans are found,
except in early spring and late fall.
Like some other zooplankton
groups, cladocerans hatch from
resting eggs when ponds are
filled. Later, eggs are held in the
females before hatching.
Cladocerans compete with rotifers
and calanoid copepods for phytoplankton.

Pond preparation
Ponds that are drained, dried, and
then filled with well water are
much safer for culturing fry than
are ponds filled with surface
water. Starting with water that

does not contain zooplankton

makes it much easier to predict
when the right size zooplankton
for the fry will appear in the
ponds. It also helps to ensure that
zooplankton, fish predators, parasites, diseases, and a variety of
fry-eating insects are not abundant in the ponds when fish are
stocked. If surface water must be
used, it should be filtered through
a filter fine enough to prevent
even small copepods from passing
through (125-micron mesh).
Ponds should be properly fertilized as they are being filled. If fry
will depend on zooplankton for
food, a combination of organic
and inorganic fertilizers is best.
Organic fertilizers are the basis of
the food chain that nourishes bacteria, protozoans, zooplankton,
and eventually the fish fry. (Some
organic fertilizers, such as rice
bran, are fine enough to be directly consumed by zooplankton). As
organic fertilizers decompose,
their nutrients are used by phytoplankton, which is consumed by
some types of fry. Or, the phytoplankton is eaten by protozoans
or zooplankton before they are
eaten by fry. Nutrients from
organic fertilizers are released
over time, so they produce less
drastic changes in plankton populations than do inorganic fertilizers.
Inorganic fertilizers add nutrients
to the pond instantly. A phytoplankton-based food chain can
develop very rapidly without the
need for bacterial action.
However, the nutrients are often
used up very rapidly by the tiny
plants, and the risk of a bloom
crash is greater than it is with
organic fertilizers.
Using a combination of organic
and inorganic fertilizers results in
a greater diversity of plankton
than if either fertilizer type is used
alone, and reduces the potential
for a dangerous bloom crash.
Fertilizer nutrients are used quickly in the pond environment. Some
nutrients are trapped in the bottom mud or otherwise lost from
water. Therefore, nutrients should
be replenished often. Frequent
applications of small amounts are
more effective than a single large

application for maintaining a supply of fry food organisms.

Additional information can be
found in SRAC Publication 469,
Fertilization of Fish Fry Ponds.

Timing of fry stocking

The proper timing of fry stocking,
in relation to filling and fertilizing
the ponds, can make the difference between having an abundant
harvest or a complete crop loss.
Proper timing is also important
for optimum growth of the fry.
Ponds must contain the appropriate type and size of food when fry
are stocked. Large fry stocked into
ponds with very tiny zooplankton
may grow slowly because the fry
must expend so much energy to
catch an adequate amount of
food. Likewise, if the zooplankton
are mostly too large for the fry to
eat they may starve, or become
prey of cyclopoids or insects.
When ponds are filled and fertilized, the plant and animal populations that invade or hatch from
within the bottom mud pass
through somewhat predictable
changes in sizes and species (Fig.
2). This process is called succession. At first there are usually a
few small species in large concentrations. Later there will be many
species in an array of sizes, but
each in moderate concentrations.
The average size of organisms
also gets larger with time. The
early community is unstable and
great changes can occur quickly;
later, the greater diversity of
organisms makes the community
more stable.
Knowing how succession happens
in fry culture ponds will help a
producer be more successful.
Figure 2 illustrates the successional process in ponds in Arkansas
during the spring when temperatures are 21 to 24 o C (70 to 75 o F)
(sunshine bass fry were also present in these ponds). When ponds
are first filled with well water,
there are few living organisms
and few nutrients. The water
rapidly gains nutrients from the
bottom, particularly when soluble
inorganic fertilizers are added. It
also gains nutrients, but more
slowly, as organic fertilizers are
decomposed by bacteria. Phyto-

plankton and other bacteria rapidly use released nutrients. Within a

few days, growing populations of
phytoplankton may provide a
green tinge or bloom to the
water. This indicates that there is
a growing food base for singlecelled protozoans and other zooplankton. In many ponds the
water first appears brownish. This
happens when the bacterial food
levels are large enough to cause
huge protozoan or rotifer blooms
without much phytoplankton
being present.
Although some protozoans may
be large enough for tiny fry to eat,
it is the next stages in succession
that are of greatest importance to
fry. Rotifers usually appear first.
Rotifers feed on bacteria and phytoplankton, and then reproduce to
form huge populations. When
water temperature is 21 to 27o C
(70 to 80 o F), rotifers can go from
nearly nonexistent to concentrations in the thousands per liter by
the second week after a pond is
filled. As rotifers eat their own
food supply the population drops
drastically. Then copepod nauplii,
adult copepods and cladocerans
make their appearance. Cyclopoid
copepods prey on small rotifers.
Calanoid copepods and herbivorous cladocerans out-compete
them for phytoplankton. Together,

copepods and cladocerans prevent

a re-bloom of the smallest rotifers.
However, modest populations of
larger rotifers may appear after
several weeks, particularly when
fish fry prey on the rotifers competitors and predatorscladocerans, copepods and insects.
Because of the ephemeral nature
of high density rotifer populations, timing is critical if the fry
being stocked are so small that
they can eat only rotifer-sized
prey. Most fry 6 mm long or less
fall into that category. Fry must be
stocked just before the rotifer population begins its rapid growth
(Fig. 2). If the fry are stocked
when rotifer populations are
rapidly rising there will be plenty
of food and the fry should grow
rapidly and be large enough to eat
copepod nauplii and larger zooplankton when those organisms
appear. The fry will also have a
much better chance of being large
enough to avoid being eaten by
cyclopoid copepods.
Larger fry (more than 6 mm)
should be stocked into ponds as
populations of copepod nauplii,
copepods and cladocerans begin
to climb (Fig. 2). That usually happens 2 to 3 weeks after ponds are
filled when water temperature is
21 to 27 o C (70 to 80 o F). The fry
will then have the right size food

Figure 2. Small fry, those less than about 6 mm long, should be stocked before
rotifers reach their initial peak density. Fry longer than 6 mm may be stocked
slightly later but before predaceous insect populations are high. This graph illus trates succession of the zooplankton community as it usually occurs in a fish
culture pond that has been filled with well water and fertilized. This data was
obtained from ponds that contained sunshine bass fry that were preying on the
zooplankton.

for rapid growth and can better

escape predation from aquatic
insects that soon begin to populate
the pond.
In general, fry must have zooplankton to survive, or at least to
be healthy and grow rapidly.
Most fry are not particular about
the types of zooplankton they eat,
but the organisms must be small
enough to fit into their mouths. To
maximize survival, stock any fry
just as populations of zooplankton
small enough for the fry to eat are
rapidly increasing and before
invading predators become
numerous.
Stocking even large fry into a
pond that has been filled for more
than 3 to 4 weeks during warm
weather can result in high mortality. By that time, a variety of fry
predators have invaded the pond
and begun to reproduce. These
include insects such as back-swimmers, diving beetles and whirlygig
beetles. Later, even larger insects
such as water scorpions, giant
water beetles and the larval stages
of dragonflies will appear.
Insects begin to colonize as soon
as ponds are filled during warm
weather. However, it usually takes
several weeks for their populations to reach levels threatening to
small fish.
Predaceous cyclopoid copepods
are often a much greater threat to
fry than insects. Many of these
tiny zooplankton will prey upon
fry unless, or until, the fry are
large enough to prey upon them.
Cyclopoid copepods often are
abundant in ponds after about 10
days when water temperature is
20 to 25 o C (68 to 77 o F). Because
there are no legal means of controlling undesirable zooplankton
or insects, it is important that fry,
particularly small fry, be stocked
into ponds as early as appropriate
after ponds are filled.
The effects of weather must also
be considered when stocking fry.
Temperature has a profound effect
on the successional process. Figure
3 shows that the colder the water
the more time is required for
rotifers to reach their initial peak
population.

Figure 3. The effect of water temperature on the time to peak rotifer populations in
a fertilized culture pond. Cold water slows the development of zooplankton popu lations.
Researchers are developing methods of predicting when some zooplankton events will occur under
different temperatures. Figure 3
shows that the time it takes to
reach an initial rotifer peak is
related to the mean morning water
temperture in the following way:
Days to rotifer peak
= 29.7 0.95 (average morning
water temperature in oC)
= 46.57 0.53 (average morning
water temperature in oF).
If you know the average water
temperature on a farm for selected
dates you can predict how long it
will take to reach a peak in the
rotifer population. If the fry being
rasied require rotifers, they should

be stocked several days before the

peak.
There is also a relationship
between the mean average daily
air temperature on days between
filling the pond and reaching the
peak and the time it takes rotifers
to reach their initial peak density
(Fig. 4). The relationship is
Days to reach rotifer peak
= 27.4 0.89 (mean average
daily air temperature in oC)
= 43.22 0.49 (mean average
daily air temperature in oF)
where average daily air temperature is defined as high daily temperature plus low daily temperature divided by 2.

Figure 4. The number of days it takes to reach a peak in the rotifer population in
culture ponds at Stuttgart, Arkansas during the spring.

Fish farmers can use this relationship to approximate the time

required for rotifer populations to
reach an initial peak starting any
spring day for any location. To do
that, normal, daily average air
temperatures for the location are
substituted into the equation and
a curve is drawn. Figure 4 illustrates this procedure for Stuttgart,
Arkansas. Normal daily air temperature data can be obtained
from local airports or from the
U.S. Department of Commerce,
National Climatic Data Center,
Federal Building, Asheville, North
Carolina 28801. Local
Climatological DataUnedited
can also be obtained at
www.ncdc.noaa.gov.

Weaning fry to artificial

feed
Some fry, such as channel catfish,
can be grown on artificial feeds
alone, but even these fry will grow
better and be healthier when zooplankton are present. However, an
objective in all fry culture is to
train the fry to eat artificial feeds,
because fry ponds are stocked at
densities so high that natural
foods alone cannot sustain good
growth of fingerlings. Unless fish
are adapted to filter feeding (such
as paddlefish) they expend more
energy catching zooplankton than
they derive from eating them.

At some point, fingerlings will be

consuming zooplankton faster
than it can be produced in the
pond. By that time they should be
well on their way to learning how
to consume artificial feeds. This
learning process should be completed before zooplankton stocks
are exhausted.
Sometimes, however, small fingerlings are brought into tanks and
trained to eat manufactured feed.
This abrupt change from natural
feeds can cause high mortality. It
is better to wean fish to manufactured feeds by a process called
feeding the pond.
Feeding the pond means spreading finely ground feed across the
pond at a rate of about 10 to 50
percent of the weight of fry in the
pond. It is hard to determine the
weight of the fry, so the typical
amount used is 1 to 2 pounds of
feed per acre for small fry and 4 to
8 pounds of feed per acre for catfish and other large species. The
fish should be fed twice a day for
3 weeks. For small fry, use the
finest size feed available. For larger fry, a fine crumble may be used.
After the initial 3-week period,
increase the amount of feed to 3 to
7 pounds per acre (12 to 24
pounds for large fry) applied
twice a day for another 3 weeks.
At the same time, increase the size
of the feed. After the second 3week period, check to see if the fry
are large enough to eat a crumbled
feed.

When the fish begin to take feed at

the surface, the amount of feed
offered can be adjusted to meet
their needs by watching the fishes
behavior. Feed just a small amount
more than they immediately consume and later check to see if the
additional feed is eaten. Adjust
the amount of feed offered accordingly.
Fry feeds have a higher protein
content than feeds for larger fish,
because fry are growing at a faster
rate and must consume large
amounts of protein and other
nutrients. Fry of fish that are piscivorous (fish eaters), such as
striped bass and their hybrids,
require a protein content of about
55 percent. Minnows and catfish
require feeds with 40 to 50 percent
protein. The general rule is that
the higher the adult fish are on the
food chain, the more protein the
fry require.
Most fingerling producers remove
their fish from the ponds and sell
them, or restock them at lower
densities, at the time when natural
food in the pond can no longer
support the standing crop of fingerlings.
A fingerling operation that is carefully planned and based on what
is happening in the pond has the
best chance of being profitable.

The work reported in this publication was supported in part by the Southern Regional Aquaculture Center through Grant No. 94-38500-0045 from
the United States Department of Agriculture, Cooperative States Research, Education, and Extension Service.

NRAC Fact Sheet No. 222-1994

Northeastern Regional
Aquaculture
Center
University of Massachusetts
Dartmouth
North Dartmouth
Massachusetts 02747

Evaluation of Artificial Diets for Cultured Fish

Michael A. Rice, Fisheries, Animal & Veterinary Science, University of Rhode Island
David A. Bengtson, Zoology, University of Rhode Island
Carole Jaworski, Rhode Island Sea Grant, University of Rhode Island

Introduction
One of the major expenses in any fish culture
operation is the cost of feeds for the fish, and the
profitability of many operations is frequently tied to
the cost of feed. Hatchery production of fish larvae
most often requires the expensive production of live
food (phytoplankton and zooplankton), because
artificial diets are either not available, or are grossly
inadequate. Artificial diets are available for growout of
fingerlings and adults of most cultured fish species, but
they may be less than optimal because they had been
formulated for another species. For example, in the
United States,commercially formulated diets are
available for catfish and salmonids, but these diets have
been used without modification to feed other species of
fish, including hybrid striped bass, tilapia, carps, and
others. Less than optimum diets for growout of fingerlings will result in lowered growth rates and excessive
waste, either by excessive fecal material, excessive
urinary nitrogen, or uneaten food. Thus, less than
optimum diets are not only wasteful in terms of money
spent on feed, but they can cause increased waste
management problems. The key challenge of producing
production feeds is the maximization of fish growth
with a minimization of waste.

problems at first feeding and disease, can often be

greater than 50 precent. The development of
high-quality artificial diets could potentially ameliorate
water quality and disease problems, as well as reduce
the high cost of live feed.

Evaluation of Production Diets

For most practical applications, evaluation of
production diets (diets for fingerling and adult production) can be adequately done in feeding trials. Since
diets are available that have a well-defined composition, growth performance of fish can be readily determined after modifications of a control diet are made.
Typically, the total feed utilization by fish, expressed as
food conversion ratio (FCR), or the protein utilization,
expressed as protein efficiency ratio (PER), are calculated. The highest quality production diets will have
relatively low FCRs and high PERs.
One of the simplest means for an aquaculture
producer to assess feed performance is to determine a
food conversion ratio (FCR). The FCR is the weight of
food supplied divided by the weight gain of the fish
during the feeding period. FCR can be expressed by the
equation:
FCR = F/(Wf-Wo)
(1)

The challenges of formulating diets for larval fish

are more formidable, as evidenced by a reliance on live
feeds. One major challenge of larviculture is the production of organisms that areas similar as possible to
those in nature. Marine fish larvae that hatch and grow
in nature typically feed on zooplankton in an environment of low fish density and good water quality;
mortality is often >90 percent, due primarily to problems at first feeding and to predation. By contrast,
those that hatch and grow in larval rearing systems
typically feed on rotifers, Artemia and prepared diets in
an environment of high fish density and (at best)
adequate water quality; mortality, due primarily to

when,

and

is the weight of food sup

plied to fish during the
study period,

wo

is the live weight of fish at

the beginning of the study
period,

is the live weight of fish at

the end of the study period.

Example: A fish pond operator starts with 1,000

fingerlings at an average weight of 200g each. The
aggregate W0 is 200 kg. The fish are fed 7g food/fish/
day for 6 months when they are harvested at a final
average weight of 900g each, but there has been 2
percent mortality. The aggregate Wf would be:
1,000- (1,000 X 0.02)= 980 fish x 900g = 882kg

Now, if a feed were chosen with the reduced crude

protein content of 35 percent and the fish growth is the
same, the PER would be:
PER = (882kg - 200kg)/(1,274kg x 0.35) = 1.53
The PER values are reduced when protein levels in
the feed are either insuffient or are in excess. Optimum protein content in fish feeds is species specific
and occurs when PER is maximized.

The amount of food supplied would be:

7g/day/fish x 182 days x 1,000 fish= l,274kg
Then, the FCR would be:

Evaluation of Larval Diets

Diets for larval fish are notoriously difficult to
evaluate because there are no completely defined artificial diets that are adequate for fish growth. Larval fish
producers are currently reliant upon live feeds, so active
work with artificial diets is largely confined to the research community. A comprehensive evaluation of an
artificial diet only begins with a well-controlled experiment to compare survival, growth, and perhaps other
indicators (e.g., stress/activity tests) of the larvae to
those obtained with live food (either rotifers, Artemia or
natural zooplankton). If equivalency is not obtained,
one then needs to investigate the causes of the deficiency, realizing that those causes may not even be in
the formulation of the diet. Two basic categories of research are required: 1) research on the physical and
chemical state of the diet in the water column; 2) re search on the physiological and biological requirements
of the larvae. Two special caveats should be noted here:
1) all research should be conducted, and the results
expressed, relative to live food; and 2) if the artificial
diet happens to be microncapsulated, it is necessary to
investigate deficiencies in the diet and the
microcapsule separately. One way to study the
microcapsule separately is to microencapsulate live
food, as Leibovitz (1991) has done with Artemia nauplii.

FCR = 1274kg/(882kg - 200kg) = 1.87

A very important factor to remember when FCRs
are compared is that they are based on the wet weight
of the feed. Different feeds may have very different
moisture levels. For example, a dry catfish production
diet may have a moisture content of around 10 percent,
whereas a semi- moist diet for sea bass may have a
moisture content of over 60 percent. Moisture does not
contribute to the growth of fish, but does add a bias to
the FCR values. Thus, if comparisons are made between
two or more diets, it is often useful to calculate the FCR
on a dry weight basis. To make this easier, it is important to know the percent of moisture and dry weight
in both your feeds and fish.
High protein ingredients are frequently the most
expensive components of artificial diets. Consequently,
feeding a diet too high in crude protein will not only
be wasteful in terms of cost, but excess excretory
nitrogen resulting from the breakdown of protein for
energy metabolism may be a stressor to the fish. One
means for determining the optimum level of protein in
a selected feed is to compare the protein efficiency
ratios (PER) of different feeds fed to fish. PER is the
weight gain of fish divided by the dry weight of protein
in the feed. An equation describing PER would be:

Diets in the Water Column

After they are introduced into the water column,
diets should remain both available and palatable to the
larvae without leaching significant amounts of nutrients. Ideally, the diet should be neutrally buoyant; but
in practice, this is very difficult to achieve. Many
sinking diets can be kept in the water column with
sufficient aeration, but the aeration levels required may
be detrimental to the larvae due to the excessive agitation. Estimation of the availability of the diet to the
larvae is possible even without the larvae in the system
simply by measurement of the residence time of the
diet: 1) at the surface; 2) in the water column; and 3) at
the bottom. For example, Leibovitz (1991) quantified
the percentages of diet particles (microencapsulated
Artemia nauplii) at each of the three locations over an
8-hour period to demonstrate that they spent 2-4 hours

PER = (Wf-W0)/F x p

(2)
when,

F is the weight of feed

supplied over the test period,

and

p is the fraction of crude

protein weight in the feed,

For example, if the percentage of crude protein in

the feed from the above example were 40 percent, the
PER over the 6 month growth period would be:
PER = (882kg - 200kg)/(l,274kg x 0.4) = 1.34

at the surface, 1-2 hours in the water column, and the

remainder at the bottom.

enous enzymes in the digestion of live vs. artificial food

in the larval fish digestive tract has been studied and
debated for years (Dabrowski and Glogowski 1977a,
1977b); however, the addition of digestive enzymes to
artificial diets has had varying degrees of success/failure
(Dabrowski and Glogowski 1977c; Dabrowska et al.
1979; Lauff and Hofer 1984; Tandler and Kolkovsky
1991).

Leaching of essential nutrients from larval diets

has long been considered a serious problem, with water
soluble vitamins being the most susceptible (Meyers
1979). Microencapsulated and microbound diets can
help to overcome leaching, but the diets should still be
analyzed to determine the extent of the problem in a
comprehensive examination scheme. The degree of
leaching can be determined by chemical analyses for
various substances conducted either on the diet particles recovered from the water at various time intervals, or on the water itself, or on both. For example,
Leibovitz (1991) showed that microencapsulated
Arternia exhibited no significant change in proximate
composition after 2 hours in seawater. The simplest
determination of leaching includes simply the measurement of dry weight of particles recovered from the
water at various time intervals.

Morphological studies of development of larvae

include histological and histochemical methods with
light, scanning electron and transmission electron
microscopy. In the context of determination of larval
capabilities for utilization of artificial diets, the most
useful studies include examination of sensory apparatus
(e.g., taste buds) (Appelbaum et al. 1983), the alimentary canal (especially the mucosal epitheliums) (e.g.,
Kjorsvik et al. 1991; Verreth et al. 1992), the liver and
pancreas (Alami-Durante 1990). Larval fish are characterized by significant uptake of nutrients by the hindgut epithelial cells and intracellular digestion in the
supranuclear vacuoles of those cells (Iwai and Tanaka
1968; Watanabe 1984). Any morphological examination of fish larvae by researchers should emphasize
the development of hindgut epitheliums.

Palatability can be determined by the rate of the

rate of ingestion of feed particles by larvae. Simple
visual observations can be useful, provided that the
larvae and particles are large enough to be seen with
the naked eye. Alternatively, larvae can be videotaped
to record the number of strikes at prey (or particles),
number of successful ingestions, and number of rejections. The recent use of image analysis to determine
number of prey remaining in a bowl with predator(s) at
frequent time intervals (Letcher 1990) could be adapted
for palatability determinations, but has so far been
attempted only with live Artemia as prey.

Once the diet is ingested, the digestion, absorp

tion, and assimilation of the food can be studied using
fluorescence, radiolabeling, and/or histological methods. Walford et al. (1991) and Walsh et al. (1987) have
used fluorescence methods to follow particles passing
through the larval fish digestive tract, particularly
noting time of passage and bottlenecks to passage.
Determination of assimilation efficiency with radiolabeled carbon has long been practiced with larval fish
fed live food (e.g., Boehlert and Yoklavich 1984), but
has recently been used also to compare uptake of
artificial and live diets (Tandler and Kolkovsky 1991).
Assimilation effiency data, when combined with data
on rates of ingestion of live vs. artificial diets, can
provide valuable insight into artificial diet deficiencies
(e.g., to what extent reduced growth is due to reduced
ingestion vs. reduced digestibility). Whereas fluorescence and radiolabeling studies are most useful in
estimating process rates for the whole organism,
histological studies are most useful in identifying
specific digestion and absorption sites within the
digestive tract. Bengtson (1993) and Bengtson et al.
(1993) have studied uptake of live vs. artificial food by
examination of mucosal epitheliums in larval fish. By
sampling larvae at time intervals after a single feeding
and examining histological sections, one can follow the
passage of particles through the digestive tract and
answer the question: Are there differences in the
digestion and absorption of live vs. artificial diets?
Larval striped bass appear to absorb all of the nutrients

Determining Digestive Capabilities and

Nutritional Requirements of Larvae
The digestive capabilities and dietary requirements
of the larvae can best be determined through a combination of biochemical, physiological, and morphological studies. Perhaps the greatest challenge to larval fish
nutritionists is the integration of information from
those studies in the formulation of adequate artificial
diets. Some approaches to estimation of the nutritional
requirements of a given species have included biochemical analyses of: 1) yolk material in eggs of that
species; 2) zooplankton on which the species feeds, or
3) Artemia (Leibovitz et al. 1987). Estimation of the
physiological capabilities of larvae have included
studies of the development of digestive enzyme production (e.g., Baragi and Level 1986) and determinations of
the pH of the digestive tract in which the enzymes
must function (e.g., Buddington 1985). Of particular
value are those studies that locate the portions of the
digestive tract responsible for the addition of specific
enzymes through histochemical means (e.g., Segner et
al. 1989). The relative roles of exogenous and endog-

from live Artemia nauplii through the hindgut epithelial cells, but do not absorb nutrients from artificial
diets through those cells.

on the structure and function of the digestive system of Menidia beryllina (Pices,
Atherinidae).pp. 11%208. In: B. T.Walther and H. J. Fyhn, editors Physiology and
biochemistry of fish larval development. University of Bergen, Norway.
Boehlert, G. W. and M. M. Yoklavich. 1984. Carbon assimilation as a function of
ingestion rate in larval Pacific herring, Clupea harengus pallasi Valenciennes.
Journal 1 of Experimental Marine Biology and Ecology 79:251-262.

Conclusion
The evaluation of artificial diets for adult and
juvenile fish has been largely based upon feeding trials
with great success, because defined basal diets are
available. Aquaculture producers can use simple feeding
trial techniques to evaluate the efficiency of feed
utilization by their fish and the cost-effectiveness of
feeds from different sources. Evaluating feeds for larval
fish is not as simple because of the lack of an adequate
artificial basal diet. Research has evolved from the
one-dimensional approach of formulating a variety of
diets and simply obtaining survival and growth results
from feeding trials with larval fish. The multidisciplinary nature of a comprehensive evaluation of an
artificial diet now requires that many groups communicate and cooperate with each other. These groups
include, but are not limited to, nutritional biochemists,
food chemists and engineers, physiologists and
morphologists, and the Aquaculturists themselves.
Continuation (and undoubtedly expansion) of such a
multidisciplinary approach provides the best chance of
defining an inert or artificial diet that can compete
with live rotifers and Artemia as a nutritional source for
marine fish larvae. The ideal artificial diet, however,
will produce marine fish larvae that biochemically,
physiologically, and behaviorally resemble wild larvae
that feed on natural zooplankton.

Buddington, R. K. 1985. Digestive secretions of lake sturgeon, Acipenser

fulvescens, during early development. Journal of Fish Biology 26:715-723.
Dabrowska, H.C. Grudniewski and K. Dabrowski 1979. Artificial diets for
common carp effect of addition of enzyme extracts. Progressive Fish-Culturist
41:196-200
Dabrowski, K. and J. Glogowski. 1977a. Studies on the proteolytic enzymes of
invertebrates constituting fish food Hydrobiologia 52:171-174.
Dabrowski, K. and J. Glogowski. 1977b. The role of exogenic proteolytic enzymes
in digestion processes in fish Hydrobiologia 54:129-134.
Dabrowski, K and J. Glogowski. 1977c. A study of the application of proteolytic
enzymes to fish food. Aquaculture 12:349-360.
Iwai, T. and M. Tanaka. 1%8. The comparative study of the digestive tract of
teleost larvae-III. Epithelial cells in the posterior gut of halfbeak larvae. Bulletin of the
Japanese Society of Scientific Fisheries 34:44-48.
Kjersvik, E., T. van der Meeren, H. Kryvi, J. Amfinnson and P. G. Kvenseth
1991. Early development of the digestive tract of cod larvae, Gadus morhau L.,
during start- feeding and starvation. Journal of Fish Biology 38:1-1S.
Lauff, M. and R. Hofer. 1984. Proteolytic enzymes in fish development and the
importance of dietary enzymes, Aquaculture 37:335-346.
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Island, Kingston, Rhode Island, USA.
Leibovitz, H. E., D.A. Bengtson, P.D. Maugle and K. L. Simpson.1987. Effects
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Acknowledgments
The authers gratefully acknowledge the constructive criticisms of four anonymous NRAC reviewers who
helped to improve the manuscript. Preparation of this
fact sheet was supported by grants from the Northeastern Regional Aquaculture Center. This is Contribution
Number 2960 from Rhode Island Cooperative Extension and RIU-G-94 -001 and P1363 from Rhode Island
Sea Grant. Portions of this NRAC fact sheet have been
reprinted from the Journal of the World Aquaculture
Society (vol. 24, pp. 285-293), with permission.

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Rombout, R. Booms and H. Segner. 1992. The development of a functional
digestive system in the Afican catfish Clarias gariepinus (Burchell). Journal of the
World Aquaculture Society 23:286-298.

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SRAC Publication No. 186

Southern
Regional
Aquaculture
Center

May 1993

Channel Catfish
Dietary Effects on Body Composition and Storage Quality
Catfish diets must provide enough
energy, protein, vitamins and minerals in the proper proportions for
fast, efficient growth and health
maintenance. Choosing the right
feed plays an important role in determining the productivity and
profitability of aquaculture operations. But, producers arent the
only people who are interested in
diet quality. Certain characteristics
of the diet influence the quality of
catfish products during processing
and storage. As a result, catfish
processors, wholesale marketers
and retailers also depend on
proper feed quality to yield desirable results.

Effects of dietary protein

Because excess fat decreases the
dress-out yield and potential shelflife of processed catfish, questions
regarding the impact of diet composition on product quality have
arisen. Research projects evaluated
the effects of dietary protein content on body composition of various-sized catfish in different
production systems.

The Southern Regional Aquaculture

Center has supported research to determine how diet affects the composition
and quality of processed channel catfish
products. The results of-these research
projects should help producers and processors modify management practices to
improve the value and marketability of
their products. This publication was
compiled by James T. Davis, D.M. Gat
lin, III and Max R. Alleger, based on research conducted at Auburn University,
University of Georgia, Kentucky State
University and Texas A&M University,
Details are available from the applicable
publications listed.

the fish weighed an average of 1

pound. Results of this experiment
indicated that neither feeding frequency nor protein content of the

diet within this range affected

growth or important body composition characteristics such as fat
content or fillet yield.
In an Auburn University study,
fingerlings were fed commercialtype diets containing 26,32 or 36
percent protein for 125 days on
either a restricted basis or to satiation. Dressing percentage increased as dietary protein was
increased from 26 to 32 percent,
but then decreased as protein concentration was increased to 36 percent. Whether fed on a restricted
basis or to satiation, body fat decreased as the diets protein level
increased. Body fat content of fish
fed to satiation was higher than
those fed on a restricted basis,

Catfish grow-out in ponds

At Kentucky State University, fingerlings stocked in intensively

managed ponds were fed commercial-type diets containing 34 or 38
percent protein to satiation once or
twice daily. At the end of 170 days

Choosing the right feed plays an important part in profitability of aquaculture

operations.

suggesting that feeding rate does

influence important body composition variables.
Another study conducted at
Auburn University measured the
effects of feeding commercial-type
diets containing 24,28,32,36 or 40
percent protein to fingerlings in
ponds. Fingerlings were fed to an
average of 1 pound in 151 days.
They were fed to satiation once
daily during the growing season.
Dressing percentage increased as
dietary protein was increased
from 24 to 36 percent, but decreased when increased to 40 percent. Fat content in fillets
decreased, while protein and moisture increased when dietary protein was increased.
Conclusions
These studies suggest that fillet
yield may improve as dietary protein is increased up to 36 percent,
and that feeding to satiation may
increase body fat concentrations.
However, the same studies suggest that producers can save
money without sacrificing weight
gain by feeding diets that contain
much less than 36 percent protein.
These trade-offs between
economic savings and potential
changes in product quality deserve further attention, especially
if fat content and other body composition characteristics are proven
to reduce the quality and consumer acceptance of catfish products.
Catfish grow-out in cages
Cage culture offers an opportunity
to produce fish in ponds that may
be poorly suited for conventional
pond culture because of their size,
depth or the presence of other fish.
However, successful cage culture
also provides unique management
challenges to the producer.
A study conducted at Kentucky
State University focused on the nutritional needs of fingerling channel catfish stocked in cages. Fish in
cages were fed to satiation once or
twice daily for 105 days with complete, commercial-type feeds con-

taining either 34 or 38 percent protein.

Body composition of fingerlings in
this study was not affected by feeding frequency or dietary protein
level. Fish grew faster on the
higher protein diet, and fish fed
twice daily had a higher dressing
percentage than those fed only
once per day.
Grow-out of third-year fish in
ponds
Little information is available regarding how diet affects the
growth or body composition of
third-year fish. Studies conducted
at Auburn University and Kentucky State University measured
the effects of feeding commercialtype diets containing various concentrations of protein to third-year
catfish in ponds. Results of these
studies indicate that, although
body fat decreases when dietary
protein is increased, fish growth
and dressing percentage were unaffected. It remains unclear
whether diets containing more
than 32 percent protein improve
the quality of fillets from thirdyear fish enough to justify the accompanying higher feed costs.

Supplementing amino
acids in catfish diets
Amino acids are the building
blocks of protein; they are essential for good fish growth and
weight gain. Several research projects have focused on the effects of
specific amino acid supplements
in catfish diets. These studies were
conducted with catfish fingerlings
in aquaria maintained under optimum conditions.
Lysine is one of ten amino acids
that must be provided by the diet;
it is also the least abundant amino
acid in many feedstuffs. As a result, extra care must be taken to
provide enough lysine when formulating catfish diets containing a
large percentage of protein from
plant sources. Also, lysine supplementation above requirement levels has been shown to reduce body

fat of some terrestrial animals. Researchers at Texas A&M University compared diets that contained
either 25 or 30 percent protein
from soy isolate or casein and gelatin, and either 0 or 0.5 percent supplemental lysine. Fingerlings fed
diets with protein from casein and
gelatin gained more weight than
those fed diets containing soybased protein. Also, fish fed soybased diets contained more lipid
and less protein than those fed the
casein-based diet.
Supplemental lysine improved
protein conversion efficiency and
feed efficiency of catfish fed soybased diets, but not of those fed casein-based diets. Fish fed a 30
percent protein soy-based diet
without added lysine performed
better than those fed a 25 percent
protein soy-based diet with extra
lysine. However, supplementallysine did not influence body composition characteristics at any protein
level.
Results suggest that both the
source and concentration of dietary protein impact catfish performance, and that supplemental
lysine does not influence body
composition.
Carnitine is a naturally-occurring
compound that animals typically
produce from lysine. Some research suggests that providing
supplemental carnitine in the diet
increases the quality of processed
animal products by reducing fat
content. A study conducted at the
University of Georgia compared
the benefits of feeding diets that
contained 0.1 percent carnitine
and 1.1, 1.4 or 1.7 percent lysine.
Feeding diets that included both
supplemental carnitine and lysine
proved most beneficial. When carnitine was added to diets containing lysine close to or above the
required dietary level (1.4 and 1.7
percent, respectively), fat content
in the viscera and dark muscle tissue decreased and whole-fish protein levels increased.
Results indicate that feeding highquality diets supplemented with

carnitine may reduce body fat content.

Effects of vitamin
fortification
Improved storage quality depends
on management practices during
grow-out, as well as procedures
carried out during and after
processing. Storage quality of poultry and some kinds of fish has
been improved by feeding diets
that increased concentrations of vitamin E in muscle tissues prior to
processing. Vitamin E, and similar
synthetic products, are called antioxidant because they help reduce
lipid oxidation and maintain the
freshness of products during storage.
Researchers at Texas A&M University evaluated the benefits of adding synthetic and natural
antioxidants to channel catfish
diets. Fingerling catfish were fed
experimental-type diets that satisfied all known requirements and
contained one of two concentrations of vitamin E (60 or 240
mg/kg), either alone, or in combination with one of four synthetic
antioxidants. None of the synthetic
antioxidants affected weight gain,
feed efficiency, survival or tissue
composition. Fillet samples from
fish receiving each diet were frozen at -10F for six months. The
TBA number, a measure of rancidity caused by oxidation, was determined for fillets to assess how
stability during frozen storage was
affected by diet composition. Fish
fed the higher level of vitamin E
had reduced TBA numbers, but
synthetic antioxidants did not affect this measure of storage quality.
Fortification of catfish diets with
high levels of vitamin E probably
offers an effective means of maintaining fillet stability during frozen storage. Results from another
study indicate that maximum
benefits from vitamin E supplementation are achieved within 2
weeks of feeding a diet fortified
with 1,000 mg vitamin E/kg. Feeding diets fortified with this high
level of vitamin E throughout the

Dietary protein concentration does not appear to affect storage quality of catfish
fillets.

grow-out period may be unnecessary.

Vitamin C is essential for normal
fish growth and has some properties that allow food products to resist oxidation. Rutin, a compound
classified as a bioflavonoid, may
produce beneficial responses similar to vitamin C when available in
the diet. A Texas A&M University
study compared diets containing
various concentrations of vitamin
C and rutin. By the end of eight
weeks, no differences in weight
gain, feed efficiency or survival
due to diet were observed. But,
within 10 weeks, fish fed diets
without supplemental vitamin C
or rutin had developed deformed
spinal columns, external hemorrhages and eroded fins. By week
12, fish receiving supplemental
rutin but no vitamin C showed the
same symptoms. By the end of 16
weeks all fish that had not received supplemental vitamin C
had reduced weight gain and feed
efficiency and decreased survival.
Rutin had no significant effect on
weight gain, feed efficiency or
other variables, either alone or
when fed with added vitamin C.
According to this study, high concentrations of supplemental vitamin C (1,500 and 3,000 mg/kg)
improved the oxidative stability of
channel catfish fillets, but supplemental rutin was not beneficial.

Dietary impacts on
storage quality
Studies at the University of Georgia focused on how dietary protein concentration and packaging
method may affect the quality of
frozen catfish fillets. Year-2 and
year-3 catfish stocked in research
ponds were fed diets containing
24,28,32,36 or 40 percent protein
to an average harvest weight of 3.3
pounds. Upon processing, fillets
were packaged using PVDC film
overwrapping, vacuum packaging
with Eva bags or vacuum skin
packaging and stored at -10F. Fillets were removed from frozen
storage after O, 30 and 90 days for
chemical analysis and sensory
evaluation. Chemical analyses included pH, TBA number, ammonia and free fatty acid content. A
consumer panel evaluated broiled
samples for off-flavor, greasiness
and texture. Although lower dietary protein increased fillet fat content, it did not directly affect TBA
number, pH or sensory attributes.
Sensory panelists reported that all
fillets became tougher, but greasiness decreased as storage time increased. Packaging treatment did
not impact the free fatty acid characteristics of fillets.
Results indicate that lower protein
diets may increase the fat content
of catfish fillets, but not to a degree that reduces consumer satisfaction. Also, current processing
and packaging methods for catfish
provide adequate quality protection for up to 3 months of frozen
storage.

Another University of Georgia

study evaluated the impact of dietary protein on channel catfish
stored on ice. Fish were fed to an
average size of 3.3 pounds on commercial-type diets containing
either 24, 28, 32, 36 or 40 percent
protein in production ponds. Fish
were processed upon harvest, and
fillets were placed on polystyrene
trays overwrapped with plastic
wrap and stored in drained ice
chests.
Sensory and chemical evaluations
of iced fillets were conducted after
1,7,14 and 21 days. Catfish fed
the lowest-protein diet had more
body fat than those fed higher-protein diets, but fillet fat content had
no direct effects on free fatty acid
content during storage. Also, dietary protein did not affect ammonia concentration, pH, TBA
number or bacterial counts of fillets stored on ice for up to 2
weeks. Sensory panelists reported
that the texture of fish fed the 32
percent protein diet was superior
to those fed 36 percent protein
after 1 day of storage. After 1
week, fish fed 28 percent protein
were more greasy than those fed
the other diets, but other differences were not detected.
Results suggest that dietary protein does not influence important
quality attributes of channel catfish fillets, and that fillets can be
stored on ice for up to 2 weeks
without compromising quality.

References
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Bai, S. C. and D. M. Gatlin III.
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The work reported in this publication was supported in part by the Southern Regional Aquaculture Center through Grant No. 89-38500-4516 from the United
States Department of Agriculture.