Bureau and Cho 1999 Introduction To Nutrition and Feeding of Fish

1
An Introduction to
Nutrition and Feeding of Fish
Dominique P. Bureau and C.Young Cho
Fish Nutrition Research Laboratory
Dept. of Animal and Poultry Science
University of Guelph, Guelph, Ontario, N1G 2W1, Canada
email: dbureau@aps.uoguelph.ca
In culturing fish in captivity, nothing is more important than sound nutrition and adequate
feeding. If the feed is not consumed by the fish or if the fish are unable to utilize the feed
because of some nutrient deficiency, then there will be no growth. An undernourished animal
cannot maintain its health and be productive, regardless of the quality of its environment.
The production of nutritionally balanced diets for fish requires efforts in research, quality
control, and biological evaluation. Faulty nutrition obviously impairs fish productivity and
results in a deterioration of health until recognisable diseases ensues. The borderlines between
reduced growth and diminished health, on the one hand, and overt disease, on the other, are very
difficult to define. There is no doubt that as our knowledge advances, the nature of the
departures from normality will be more easily explained and corrected. However, the problem of
recognizing a deterioration of performance in its initial stages and taking corrective action will
remain an essential part of the skill of the fish culturist.
1. Protein and Amino Acid Requirements of Fish

Protein
Protein is required in the diet to provide indispensable amino acids and nitrogen for
synthesis of non-indispensable amino acids. Protein in body tissues incorporate about 23 amino
acids and among these, 10 amino acids must be supplied in the diet since fish cannot synthesise
them. Amino acids are need for maintenance, growth, reproduction and repletion of tissues. A
large proportion of the amino acid consumed by a fish are catabolized for energy and fish are
well-adapted to using an excess protein this way. Catabolism of protein leads to the release of
ammonia.
Protein is the most important component of the diet of fish because protein intake
generally determines growth (protein growth has, in general, priority), has a high cost per unit
and high levels are required per unit of feeds.
First observations on fish protein and amino acid requirements came from studies on
natural diet of different fish. Natural diet (plankton, invertebrates, fish) is generally rich in
protein and has a good amino acid balance. All dietary proteins are not identical in their
Bureau, D.P. and C.Y. Cho. 1999. Nutrition and feeding of fish. OMNR Fish Culture Course, Unversity of Guelph,
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Guelph, Ontario, 21-25 June 1999.
nutritive value. The nutritional value of a protein source is a function of its digestibility and
amino acid makeup. A deficiency of indispensable amino acid creates poor utilization of dietary
protein and hence growth retardation, poor live weight gain, and feed efficiency. In sever cases,
deficiency reduces the ability to resist diseases and lowers the effectiveness of the immune
response mechanism. For example, experiments have shown that tryptophan-deficient fish
become scoliotic, showing curvature of the spine, and methionine deficiency produces lens
cataracts. Salmonid diets generally contain 35-45% digestible protein (DP), or 40-50% crude
protein. However, amino acids or protein must be supplied in relation to digestible energy (DE).
The recommended ratio of protein to energy in the salmonid diet is 20-26 g DP/MJ DE (92-102 g
protein per Mcal). Increasing these proportions increases ammonia excretion; the requirement for
dissolved oxygen is also increased because the efficiency with which the energy is used is
decreased.
Why do fish have such high requirements for protein? The main factors explain this
phenomenon:
1)
The protein requirement in terms of dietary concentration (% of diet) is high but

the absolute requirement isnt (g/kg body weight gain). This is due to the fact that
fish have a lower absolute energy requirement than mammals. This results in
similar g body weight gain/g protein ingested as mammal but better feed
efficiency (gain:feed).
2)
Protein (amino acids) is used as a major energy source. Some economy can be
made here if other dietary fuel are present in adequate amounts, e.g. increasing
the lipid (fat) content of diet can help reduce dietary protein (amino acid)
catabolism and requirement. This is referred to as protein-sparing effect of lipids.
Protein to useful energy ratio is the factor that should be considered, not %
protein of the diet per se.
Indispensable amino acid requirements
10 Indispensable amino acids

Phenylalanine (Phe) Histidine (His)
Lysine (Lys)
Methionine (Met)
Arginine (Arg)
Threonine (Thr)
Isoleucine (Iso)
Tryptophan (Trp)
Leucine (Leu)
Valine (Val)
2
Table 1. Indispensable amino acid requirements of different species of teleost (g / 100 g protein)
Amino
acids
Salmonid
Catfish
Carp
Tilapia
Milkfish
Arg
4.2
4.3
4.4
4.1
5.6
His
1.6
1.5
2.4
1.7
2.0
Ile
2.0
2.6
3.0
3.1
4.0
Leu
3.6
3.5
4.7
3.4
5.1
Lys
4.8
5.0
6.0
4.6
4.0
Thr
2.0
2.1
4.2
3.8
4.9
Trp
0.6
0.5
0.8
1.0
0.6
Val
2.2
3.0
4.1
2.8
3.0
Met+Cys
2.4
2.3
3.5
3.2
4.8
Phe+Tyr
5.3
4.8
8.2
5.6
5.2
Sea
Bream
Sea
Bass
5.0
4.8
0.6
4.0
4.4
Table 2. Amino acid composition of common protein sources (g/ 100 g protein).
CP
Met
Lys
Trp
Thr
Ile
His
Val
Leu
Arg
(+Cys)
Requirement
Fish meal
Soybean meal
Corn gluten meal
Blood meal
Meat and bone
meal
Poultry by-product
meal
Feather meal
Phe
(+Tyr)
4.8
0.6
2.0
2.0
1.6
2.2
3.6
4.2
68
48
60
85
50
1.7
(2.4)
3.1
1.6
3.2
1.2
1.2
7.9
6.7
1.7
6.3
4.9
1.1
1.3
0.5
1.2
0.4
4.0
4.2
3.3
4.5
4.0
4.2
5.5
3.8
0.9
3.8
8.8
2.7
2.0
3.6
3.3
7.9
5.7
4.5
6.1
5.3
7.1
8.0
15.7
12.2
5.7
8.3
8.0
3.2
2.8
6.0
2.7
(5.3)
3.6
5.7
6.3
6.0
4.0
65
1.7
5.9
0.9
4.0
2.9
2.2
4.8
5.7
7.5
2.5
85
0.7
1.2
0.5
3.3
3.1
0.3
5.4
9.2
4.6
3.1
3
2. Lipids (Fats)
Lipids (fats) encompass a large variety of compounds. Lipids have many roles: energy
supply, structure, precursors to many reactive substances, etc. In the diet or carcass of fish,
lipids are most commonly found as triglycerides, phospholipids and, sometimes, wax esters.
Triglycerides are composed of a glycerol molecule to which three fatty acids are attached.
Phospholipids are also composed of a glycerol molecule but with only two fatty acids. Instead of
a third fatty acid a phosphoric acid and another type of molecule (choline, inositol, etc.) are
attached. Wax esters are made of a fatty acid and a long chain alcohol and are a common form
of lipid storage in certain species zooplankton . The main role of triglycerides is in the storage of
lipids (fatty acids). Phospholipids are responsible for the structure of cell membranes (lipid bilayer). Fatty acids are the main active components of dietary lipids. Fish are unable to
synthesize fatty acids with unsaturation in the n-3 or n-6 positions yet these types of fatty acids
are essential for many functions. These two types of fatty acids are, therefore, essential for the
animal and must be supplied in the diet.
Deficiency in essential fatty acid result in general, in reduction of growth and a number
of deficiency signs, including depigmentation, fin erosion, cardiac myopathy, fatty infiltration of
liver, and shock syndrome (loss of consciousness for a few seconds following an acute stress).
Salmonids require about 0.5 to 1% long chain polyunsaturated n-3 fatty acids (EPA (20:5 n-3)
and DHA (22:6 n-3)) in their diet. This amount is easily covered by ingredients of marine
origins, such as fish meal and fish oil, which are always present in significant amounts in
salmonid feeds.
3. Carbohydrates
Carbohydrates represent a very large variety of molecules. The carbohydrate most

commonly found in fish feed is starch, a polymer of glucose. Salmonid and many other fish
have a poor ability to utilize carbohydrates. Raw starch in grain and other plant products is
generally poorly digested by fish. Cooking of the starch during pelleting or extrusion, however,
greatly improves its digestibility for fish. However, even if the starch is digestible, fish only
appear to be able to utilize a small amount effectively. Carbohydrates only represent a minor
source of energy for fish. A certain amount of starch or other carbohydrates (e.g. lactose,
hemicellulose) is, nevertheless, required to achieved proper physical characteristic of the feed.
4
4. Vitamins
The vitamins are generally defined as dietary essential organic compounds, required only
in minute amounts, and which play a catalytic role and but no major structural role. So far, 4 fatsoluble and 11 water-soluble vitamins or vitamin-like compounds have been shown to be
essential to fish. Requirement is generally measured in young fast growing fish. However,
requirements may depend on the intake of other nutrients, size of the fish, and environmental
stress. The recommended levels and the deficiency signs are summarized in Tables 3 and 4.
Many symptoms of vitamin deficiency are non-specific. It is also tedious and expensive to
analyze diets for vitamins. Therefore, diagnostic of vitamin deficiencies is often difficult.
Nutritional disorders caused by vitamin deficiencies can impair utilization of other nutrients,
impair the health of fish, and finally lead to disease or deformities. Nutritional deficiencies signs
usually develop gradually, not spontaneously. However, the culturist may obtain clues of
deficiency indirectly through low feed intake and poor live weight and feed efficiency.
Table 3. Vitamin requirement of salmonids.
Vitamin
Requirement
Fat-soluble vitamins
Vitamin A, IU/kg
Vitamin D, IU/kg
Vitamin E, IU/kg
Vitamin K, mg/kg
2,500
2,400
50
1
Water-soluble vitamin, mg/kg

Riboflavin
Pantothenic acid
Niacin
Vitamin B12
Biotin
Folate
Thiamin
Vitamin B6
Vitamin C
4
20
10
0.01
0.15
1.0
1
3
50
Vitamin-like compounds, mg/kg

Choline
myo-Inositol
1,000
300
5
Table 4. Deficiency signs associated with various nutrients.
Deficiency Sign
Nutrient
Anemia
Folic Acid, Inositol, Niacin, Pyrodoxine, Rancid Fat

Riboflavin, Vitamin B12, Vitamin C, Vitamin E
Vitamin K
Anorexia
Biotin, Folic Acid, Inositol, Niacin, Pantothenic Acid

Pyrodoxine, Riboflavin, Thiamin, Vitamin A
Vitamin B12, Vitamin C
Acites
Vitamin A, Vitamin C, Vitamin E
Ataxia
Pyrodoxine, Pantothenic acid, Riboflavin
Atrophy of Gills
Pantothenic Acid
Atrophy of Muscle
Biotin, Thiamin
Caclinosis : renal
Magnesium
Cartilage abnormality
Vitamin C, Tryptophan
Cataracts
Methionine, Riboflavin, Thiamin, Zinc
Ceroid liver
Rancid Fat, Vitamin E
Cloudy lens
Methionine, Riboflavin, Zinc
Clubbed gills
Pantothenic Acid
Clotting blood: slow
Vitamin K
Colouration: dark skin
Biotin, Folic Acid, Pyrodoxine Riboflavin
Convulsions
Biotin, Pyrodoxine, Thiamin
Discolouration of skin
Fatty Acids, Thiamin
Deformations: bone
Phosphorous
Deformations: lens
Vitamin A
Degeneration of gills
Biotin
Dermatitis
Pantothenic Acid
Diathesis, exudative
Selenium
Distended stomach
Inositol
Distended swimbladder
Pantothenic Acid
Dystrophy, muscular
Selenium, Vitamin E
6
Table 4. Continued
Deficiency Sign
Nutrient
Edema
Niacin, Pyrodoxine, Thiamin, Vitamin A, Vitamin E
Epicarditis
Vitamin E
Equilibrium loss
Pyrodoxine, Thiamin
Erosion of fin
Fatty Acids, Riboflavin, Vitamin A, Zinc
Exophthalmos
Pyrodoxine, Vitamin A, Vitamin C, Vitamin E
Exudated gills
Pantothenic Acid
Fatty liver
Biotin, Choline, Fatty Acids, Inositol, Vitamin E
Feed efficiency: poor
Biotin, Calcium, Choline, Energy, Fat, Folic Acid,

Inositol, Niacin, Protein, Riboflavin
Fragility: erythrocytes
Biotin, Vitamin B12, Vitamin E
Fragility: fin
Folic Acid
Fragmentation of erythrocytes
Biotin, Vitamin B12, Vitamin E
Gasping, rapid
Pyrodoxine
Goitre
Iodine
Growth, poor
Biotin, Calcium, Choline, Energy, Fat, Folic Acid

Inositol, Niacin, Pantothenic Acid, Protein, Pyrodoxine
Riboflavin, Thiamin, Vitamin A, Vitamin B12
Vitamin C, Vitamin E
Hematocrit, reduced
Iron, Vitamin C, Vitamin E
Hemoglobin, low
Iron, Vitamin B12, Vitamin C
Hemorrhage: eye
Riboflavin, Vitamin A
Hemorrhage: gill
Vitamin C
Hemorrhage: kidney
Choline, Vitamin A, Vitamin C
Hemorrhage: liver
Vitamin C
Hemorrhage: skin
Niacin, Pantothenic Acid, Riboflavin, Vitamin A, Vitamin C
Irritability
Fatty Acids, Pyrodoxin, Thiamin
7
Table 4. Continued
Deficiency Sign
Nutrient
Lesion: colon
Biotin, Niacin
Lesion: eye
Methionine, Riboflavin, Vitamin A, Vitamin C, Zinc
Lesion: skin
Biotin, Inositol, Niacin, Pantothenic Acid
Lethargy
Folic Acid, Niacin, Pantothenic acid, Thiamin
Lipoid liver
Fatty Acids, Rancid fat
Lordosis
Vitamin C
Myopathy, cardiac
Essential Fatty Acids
Necrosis : liver
Pantothenic Acid
Nerve disorder
Pyrodoxine, Thiamin
Pale liver (glycogen accumulation)
High Digestible Carbohydrate, Biotin
Photophobia
Niacin, Riboflavin
Pinhead
Starvation
Pigmentation, iris
Riboflavin
Prostration
Pantothenic Acid, Vitamin C
Rigor mortis, rapid
Pyrodoxine
Scoliosis
Phosphorus, Tryptophan, Vitamin C, Vitamin D
Shock syndrome
Essential Fatty Acids
Slime, blue
Biotin, Pyrodoxine
Spasm, muscle
Niacin
Swimming, erratic
Pyrodoxine
Swimming, upside down
Pantothenic Acid
Tetany, white muscle
Niacin, Vitamin D
Vascularization, cornea
Riboflavin
8
5. Minerals
Inorganic elements (minerals) are required by fish for various functions in metabolism
and osmoregulation. Fish obtain minerals from their diet but also from their environment. Many
minerals are required in trace amounts and are present in sufficient quantity in the surrounding
water for the fish to absorb through their gills. In freshwater, there is generally sufficient
concentration of calcium, sodium, potassium and chloride for the fish to absorb and cover its
requirements. The totality of the requirement for other minerals must, in general, be covered by
the diet. Dietary minerals play many roles. There generally have a structural (e.g. bone
formation) or catalytic (e.g. metalloenzyme) role. Minerals required by fish included calcium,
phosphorus, sodium, potassium, magnesium, iron, copper, zinc, cobalt, selenium, iodine, and
fluorine. The recommended levels of minerals in the diet are shown in Table 5. There are
numerous deficiency signs and some are highlighted in Table 4. Reduced growth, feed
efficiency and skeletal deformities is the most common signs of mineral deficiencies.
Table 5. Mineral requirement of salmonid fish in freshwater.
Mineral
Calcium (Ca)
Chlorine (Cl)
Potassium (K)
Sodium (Na)
Phosphorus (P)
Magnesium (Mg)
Iron (Fe)
Zinc (ZN)
Manganese (MN)
Copper (Cu)
Iodine (I)
Selenium (Se)
Requirement (mg/kg feed)*
10,000
9,000
7,000
6,000
6,000
500
60
30
13
3
1.1
0.3
* Requirement in the absence of significant amounts of the specific mineral in the water.
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6. Digestion
Digestive tract anatomy and physiology

Digestive system of fish is, in general, relatively simple compared to digestive system of
birds and mammals but there are numerous similarities.
Structure
Characteristics
Barbel
taste buds
Mouth
teeth, no chewing, taste buds
Pharynx
pharyngeal teeth (calcified structures)
Oesophagus
short, thick, taste buds, gizzard
Stomach
present or absent, acid, enzymes, rate of digestion correlates with

mass of food remaining in stomach, emptying is affected by
temperature.
Anterior intestine
secretive and absorptive epithelial cells, no villi but numerous

folds present, microvilli present, enterocytes with brush border
membrane
Pyloric caeca
present in some case, variable # between species and individuals

(rainbow trout 50-200 p.c.). Increase absorptive surface, number
apparently shows weak correlation with digestibility and growth.
Ratio intestine/fork length = 0.7, ratio intestine + p.c./fork length =
3.9
Pancreas
Generally a diffuse tissue, except for eel, pike, flat fish
Hindgut
Not really morphologically distinct from anterior intestine but cell

type changes. Squamous epithelial cells, mucus production, highly
vacuolated cells, absorption of macromolecules by pinocytosis
(tissue reabsorb proteins (enzymes) for recycling).
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Figure 1
Various digestive configurations
Reference :
Smith, L.S. 1989. pp.331-421. In: Halver, J.E. (Ed.). Fish Nutrition. 2nd Edition.
Academic Press, San Diego. 798p.
The Figure 1 show that anatomy of the gastrointestinal tract differs quite significantly
between species, especially between species with difference feeding habits. Difference in total
enzymes activity between species are not very pronounced for proteases and lipases but difference
are rather significant for carbohydrases.
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Estimates of apparent digestibility for salmonids

Table 6 presents the apparent digestibility coefficients for commonly used ingredients in
salmonid feeds as measured by Cho et al. (1982). Fish have different digestive capabilities
compared to terrestrial animals, and many feedstuffs, particularly cereal grains and their by-products
which contain high levels of starch and fiber, are very poorly digested by carnivorous fish. The
apparent digestibility of good quality protein by fish is very high. However, several factors can
affect the digestibility of protein. The type of drying technique used during processing is a very
important factors. A good demonstration of this is seen in blood meal. The protein digestibility of
flame-dried blood meal is very low whereas the digestibility of spray-dried blood meal is very high.
The same phenomenon can occur with fish meal.
Table 6. Apparent digestibility coefficients of ingredients measured with rainbow trout.

Apparent digestibility coefficients (%)
Ingredients
Dry
Matter
Crude
Protein
Lipid
Energy
Alfalfa meal
Blood meal
39
87
71
43
ring-dried
spray-dried
flame-dried
Brewers dried yeast
Corn yellow
Corn gluten feed
Corn gluten meal
Corn distiller dried soluble
Feather meal
Fish meal, herring
Meat and bone meal
Poultry by-products meal
Rapeseed meal
Soybean, full-fat, cook.
Soybean meal, dehulled
Wheat middlings
Whey, dehydrated
Fish protein concentrate
Soy protein concentrate
87
91
55
76
23
23
80
46
77
85
70
76
35
78
74
35
97
90
77
85
96
16
91
95
92
96
85
77
92
85
89
77
96
96
92
96
95
97
86
92
50
77
39
29
83
51
77
91
80
82
45
85
75
46
94
94
84
71
97
94
-
12
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7. Feed Formulation and Manufacturing

Diet formulation
Diet formulation and preparation are the process of combining feed ingredients to form a
mixture that will meet the specific goals of production. It is often a compromise between the
ideal formula and practical considerations. The primary objectives are to produce a mixture that
(is) :
Nutritionally balanced (to support maintenance, growth, reproduction, health)
Economical
Palatable
Water stable
Minimizes waste output & effect on water quality
Produces desirable final product (attractive & safe)
Practical considerations :
Ingredients price and availability
Pelletability of mixture
Anti-nutritional factors
Storage and handling requirements
Table 7. Composition of the grower formulae used by the OMNR Fish Culture Stations over the
past 10 years.
Ingredients
MNR89G
Formulae
MNR91H MNR95HG
MNR98HG
%
Fish meal, herring, 68% CP
Blood meal, spray-dried, 80%
CP
Corn gluten meal, 60% CP
Soybean meal, 48% CP
Poultry meal, 68% CP
Brewers dried yeast, 45% CP
Wheat middlings, 17% CP
Whey, 12% CP
Vitamin premix
Mineral premix
L-Lysine
Fish oil
20
9
35
9
18
-
18
-
17
12
20
8
0.5
0.5
13
15
14
10
0.5
0.5
16
49
6
11
1
1
14
37.6
13
9
0.5
0.5
1.4
20
37
17
22
44
20
22
44
20
22
42
21
20
Digestible Composition
Digestible protein, %
Digestible energy, MJ/kg
DP/DE, g/MJ
13
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Ingredient Quality
The first consideration for formulation and production of successful diets is the quality of
the feed ingredients. Diets produced with poor quality raw materials and under adverse
processing conditions have inferior nutritive value and adverse effects on fish health. Quality
criteria for the ingredients must be respected to insure that the final product is of consistent
quality and that deleterious effects are avoided. The chemical composition (nutrient, energy,
antinutrients, contaminants) of the ingredient obviously plays a determinant role the quality.
However, biological aspects, such as digestibility and utilization of nutrients are most important
and often overlooked.
The loss of indigestible matter from the diet as feces is the primary reason for variation in the
nutritional value of feed ingredients. Measurement of digestibility provides, in general, a good
indication of the availability of energy and nutrients, thus providing a rational basis upon which diets
can be formulated to meet specific standards of available nutrient levels. Several factors can affect
the digestibility of protein or specific amino acids. The type of drying techniques used during
processing, the composition of the protein fraction are the factors which have a determinant
effect on the digestibility of protein of a feed ingredients.
Fishery by-products:
There are various qualities of fish meals on the market, relating to the original raw fish
quality, level of ash in the meals, and the type of processing techniques used. The most important
factor is the freshness of the product. Fish must be processed as soon as possible after capture.
Ageing and spoilage decrease the nutritive value and also lead to the contamination with
potential toxic compounds, such as histamine, cadaverine, and agmatine. The second most
important factor is the type of raw material used (whole fish or by-products). By-products, such
as those generated by the filleting industry (sometime referred to as white fish meal) have higher
level of ash and lower level of protein than whole fish meals. High level of ash generally affects
digestibility of dry matter and results in high waste outputs, and can also produce mineral
imbalances (e.g. Zn deficiency).
The type of fish used is not necessarily a determinant factor in the quality of the products.
At equal freshness and if the same processing technique is used, whole capelin, anchovy,
herring, menhaden meals will support similar growth. During processing, the drying treatment is
a key factor. Flame-dried products are less digestible and produce lower performances.
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15
Table 8. Quality Standards of Fish Meal Required for Salmonid Diets.

Compound
Levels
Crude protein (%N x 6.25)
> 68%
Lipid
< 10%
Ash, total
< 13%
Salt (NaCl)
< 3%
Moisture
< 10%
Ammonia-N
< 0.2%
Antioxidant (sprayed liquid form)
< 200 PPM
ADC dry matter
> 85%
ADC crude protein
> 90%
Particle size
< 0.25 mm
Steam processed
Animal by-products
Animal protein by-products can very useful complementary protein sources in fish diets.
It is important to use highly digestible products with limited ash content. High ash content
ingredients are generally more polluting and the ash dilute useful nutrient. It is especially
important when buying these products to deal with suppliers who consistently provide high
quality products. Apparent digestibility of animal by-product is relatively high (Table 6) and
they have been used at significant levels in practical diet with success. For blood meal, the type
of drying is of primary importance. Spray-drying produce the best results.
Plant protein by-products :
There are several plant proteins and grain by-products that are used on a regular basis in
fish diet formula. Certain plant protein products have a good nutritional value (high in digestible
protein, good amino acid profile) and are economical at the same time. Other products improve
the physical characteristics of the pellets. The incorporation of certain products must be limited
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for various reasons, such as their content in starch and fibre, the presence of antinutritional or
undesirable factors and their acceptability (palatability).
Many plant products contain antinutritional factors. Most plant protein ingredients are
heat treated during processing, which greatly reduce the level of several antinutritional factors,
such as soybean trypsin inhibitors. Excess heat, however, generally decreases the nutritional
quality of plant protein products by destroying amino acids.
Fish diets formulated with high levels of certain plant protein ingredients appear to be
nutritionally adequate but not very acceptable to certain fish species. For example, diets
containing high levels of soybean meal are poorly accepted by chinook salmon and other
salmonids. Recent experimental evidences suggest that soyasaponins may be a factor affecting
performance of salmonids fed soybean meal.
Corn gluten meal is a plant protein ingredients known to be highly palatable for
salmonids. Studies with rainbow trout and Atlantic salmon show that it complements soybean
meal very well nutritionally. Recent results from our laboratory showed that corn gluten meal or
combination of corn gluten meal and soybean meal can replace most of the fish meal without any
effect on performance of the fish. Nonetheless, the incorporation of corn gluten meal must be
limited in food fish production feeds due to its high concentration in xanthophylls which can
produce undesirable pigmentation of the skin and flesh and may compete with expensive
synthetic pigment added in the feed. However, recent evidences from our laboratory do not
support this hypothesis.
Fats and Oils
Fish oil is the main source of lipid in salmonid diet. Marine fish oils are, in general,
excellent sources of long chain n-3 PUFA (EPA & DHA), fatty acids required by salmonid.
Other types of oils and fats can be used in salmonid diets. Vegetable (canola, soya, safflower,
etc.) oils and animal fats (tallow, lard, poultry fat) can also be used at certain levels in feeds
without effect on growth performance and health of the fish.
Rancidity problems: Marine oils are rich in polyunsaturated fatty acids and are susceptible to
rancidity. In all circumstances rancid oil must be avoided in the preparation of fish feeds.
Rancid fat has deleterious effect on some of the nutrients present in fish feed and health of the
fish. fatty liver disease is usually seen in fish fed rancid fat. Histologically, the main feature is
the extreme infiltration of hepatocytes by lipids. Peroxide (PV), thiobarbituric acid (TBA) and
anisidine (AV) values are in general parameters used to determine the degree of rancidity of lipid
sources. Acceptable quality parameters for fish oil as suggested by Cho et al. (1983) are
presented in Table 9. There is no unequivocal technique to measure rancidity and there is still
doubts about the reliability of PV, AV and TBA value. High PV, AV or TBA suggest problems of
lipid deterioration but are not always indicative of harmful rancidity. The easiest way to determine if
16
17
a feed is rancid may be its smell. Feed with a rancid smell must not be fed. It is preferable to
discard such feeds instead of jeopardising the health of the fish by feeding them.
Table 9. Quality Standards of Oils and Lipid in Final Product Required for Salmonid Diets
Parameters
Levels
Oils
Iodine value
Report value
Peroxide value
< 5 meq/kg
Anisidine value
< 10
Pesticides, total
< 0.4 PPM
PCB's
< 0.6 PPM
Nitrogen
< 1%
Moisture
< 1%
Antioxidant (liquid form)*
< 500 PPM
No vitamin fortification
Clean odour
Lipid in final product

Iodine value
> 135
n-3 polyunsaturated fatty acids
> 15%
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18
Diet preparation and manufacturing
The are several forms of fish feed, including wet, moist, and steam-pelleted and extruded
dry pellets. However, two basic types of formulated feed are generally used in intensive fish
culture: dry and semi-moist diets. The diets are similar, the basic difference being that semimoist pellets contain a larger proportion of raw fish and by-products which contribute a higher
moisture level to the final product. Moist feeds have some merit in coastal regions where fresh
raw fish and by-products are regularly available and economical. It is also possible that the
physical characteristics of moist pellets are more palatable to some fish species. However, there
is no evidence that such feeds are nutritionally superior to dry feeds. Moist feed may contain
pathogens since the feed ingredients are only submitted to moderate heat treatment
(pasteurization). In contrast to moist diets, dry feed are heat-treated and generally free from
pathogens. They are also easier to transport and store. The bulk purchase and storage of quality
dry ingredients is possible and ensures a continuous supply of quality feed. The dry ingredients
on the commodity market are more quality defined than raw fisheries products and can be
supplied regularly. Hence it is possible to formulate dry feeds more precisely with the available
knowledge of fish nutrition. Most nutrient in dry feeds are stable are room temperature and
therefore dry feeds can be stored safely without freezing for periods which depend on storage
conditions (approx. 3 months in a cool, shady, and well-ventilated location).
Widely used dry feeds today may divide into three types: (1) steam-pelleted feed; (2)
partially extruded, slow-sinking pellets, and (3) expanded and floating pellets. Feeding dry
pellets either by hand or with automatic feeders is much simpler than that of moist feeds. The
problem of acceptability of dry feeds by some fish species can usually be solved by better
feeding techniques and fish culture management. Otherwise, fry which have difficulty in
accepting dry feeds can be started with semi-moist feed and gradually shifted over to dry feed
within 3-5 weeks.
A formulated dry fish feed must be pelleted and/or crumbled so as to be durable and
water stable. Formulated feeds must also have desirable physical and textural characteristics, and
be of the correct sizes to be readily acceptable by different sizes of fish. Disintegrated and
uneaten feed pollutes the water and creates stresses from low oxygen and high nitrogen and
organic wastes, with serious effects on growth and health. Some of the important factors in
manufacturing a durable, dry fish feed without fines are (1) physical properties of the
ingredients, (2) particle size of ingredients, (3) conditioning time and temperature in the pellet
mill, (4) quality of steam supply, (5) compression pressure through the die, and (6) efficiency of
sifting/grading and fat-spraying equipment. Many of the dietary problems experienced in fish
culture in the past have been related to the physical quality of the pellets and granules, which
was in turn related to poor quality ingredients, inadequate manufacturing processes, and
negligent practices. Unfortunately for fish feed, the manufacturing process is of crucial
importance. Having to transfer dietary nutrients into the fish through the water medium presents
problems which are unknown in other animal-feeding practices. Therefore, all newly opened
bags should be checked for the presence of excess fines, undersized granules, durability, foreign
particles, too little or too much oil, mildew, and other evidence of poor quality. Any bag or batch
of feed judged to be questionable and any with a detectable rancid smell should not be fed.
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19
All questionable feed should be immediately reported to a qualified nutritionist and returned to
the manufacturer for replacement.
Table 10. Recommended particle size for salmonid diets

Feed
Broodstock
7 Pt.
Grower
Feed size
Feeding
per day
Fish size
(g)
0.5 - 2
> 200
Pellets
6.4 mm x 7 mm long
Pellets
6 Pt.
6.4 mm x 6 mm long
1-2
> 200
5 Pt.
4.8 mm x 5 mm long
< 200
4 Pt.
3.4 mm x 4 mm long
100
3 Pt.
2.4 mm x 3 mm long
50
Grower
Granules
3 Gr.
3 mm
< 50
2 Gr.
2 mm
20
Starter
Granules
1.5 Gr.
1.5 mm
< 10
1 mm
0.5 mm
6-8
1 Gr.
0.5 Gr.
19
20
8. Feeding Systems
Feeding systems may be defined as all feeding standards and practices employed to deliver
nutritionally balanced and adequate amount of diets to animals, so maintaining normal health and
reproduction together with efficient growth and/or work performance. Until now the feeding of fish
has been based mostly on folkloric practices while the main preoccupation has been to develop
magic diet formulae. Many hypes such as mega-fish meal and mega-vitamin C diets have come
and gone, and we are now in the age of the Norwegian Fish Doughnut (>36% fat diet)! Whichever
diet one decides to feed, the amount fed to achieve optimum or maximum gain is the ultimate
measure of ones productivity in terms of biological gain, economical benefit and/or environmental
sustainability.
Scientific approaches have been used in the feeding of land animals for over a century. The
first feeding standard for farm animals was proposed by Grouven in 1859, and included the total
quantities of protein, carbohydrate and ether extract (fat) found in feeds, as determined by chemical
analysis. In 1864, E. Wolf published the first feeding standard based on the digestible nutrients in
feeds.
Empirical feeding charts for salmonids at different water temperatures were published by Deuel and
his colleagues and were likely intended for use with meat-meal mixture diets widely in use at that
time. Since then several methods of estimating daily feed allowance have been reported.
Unfortunately all methods have been based on the body length increase or live weight gain, and dry
weight of feed and feed conversion, rather than on biologically available energy and nutrient
contents in feed in relation with protein and energy retention in the body. These methods are no
longer suitable for todays energy- and nutrient-dense diets, especially in the light of the large
amount of information available on the energy metabolism of salmonids.
Many problems are encountered when feeding fish, much more so than with feeding domestic
animals. First, delivery of feed to fish in a water medium requires particular physical properties of
feed together with special feeding techniques. It is not possible in the literal sense to feed fish on an
"ad libitum" basis, like it is done with most farm animals. The nearest alternative is to feed to "nearsatiety" with very careful observation over a pre-determined number of feedings per day; however,
this can be very difficult and subjective. Feeding fish continues to be an "art" and the fish culturist,
not the fish, determines "satiety" as well as when and how often fish are fed. The amount of feed not
consumed by the fish can not be recovered and, therefore, feed given to them must be assumed eaten
for inventory and feed efficiency calculations. This can cause appreciable errors in feed evaluation
as well as in productivity and waste output calculations. Meal-feeding the fish pre-allocated amounts
by hand or mechanical device based on theoretical energy requirement may be the only logical
choice. Uneaten feed represents an economical loss and becomes 100% solid and suspended wastes!
Meal-feeding a pre-allocated amount of feed calculated based on the theoretical energy requirement
of the animal may not represent a restricted feeding regime as suggested by some since the amount
of feed calculated is based on the amount of energy required by the animal to express its full growth
potential.
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21
There are few scientific studies, based on nutrition and husbandry, on feeding standards and
practices; however, there are many duplications and "desktop" modifications of old feeding charts
with little or no experimental basis. Since the mid-1980's, development of high fat diets has led to
most rations being very energy-dense, but feeding charts have changed little to reflect these changes
in diet composition. Most feeding charts available today tend to over-estimate feed requirements
and this overfeeding has led to poor feed efficiencies under most husbandry conditions, and this
represent a significant, yet avoidable, waste of resources for aquaculture operations. In addition, it
may results in self-pollution which in turn may affect the sustainability of aquaculture operations.
Recent governmental regulations imposing feed quota, feed efficiency guidelines and/or stringent
waste output limit may somewhat ease the problem. Sophisticated feed management systems, such
as underwater video camera or feed trapping devices, have been developed to determine fish
satiation or the extent of feed wastage and are promoted by many as a solution to overfeeding.
However, regardless of the feeding system or method used, accurate growth and feed requirement
models are needed in order to forecast growth and objectively determine biologically achievable
feed efficiency (based on feed composition, fish growth, composition of the growth). These
estimates can be used as yardsticks to adjust feeding practices or equipment and to compare results
obtained.
The development of scientific feeding systems is one of the most important and urgent
subjects of fish nutrition and husbandry because, without this development, nutrient dense and
expensive feeds are partially wasted. Sufficient data on nutritional energetics are now available to
allow reasonably accurate feeding standards to be computed for different aquaculture conditions.
Presented here is a summarized review of the basis of a nutritional energetic approach to estimating
feed requirement and waste output of fish culture operation as well as the development of the FishPrFEQ computer program. Results obtained from a field station are presented and provide a
framework to examine the type of information that can be derived from bioenergetic models and
generate a feed requirement scenario for the next production year.
PRODUCTION RECORDS
Evaluating and/or predicting growth performance of a fish culture operation or a stock of
fish firstly requires production records of past performance. These records may become databases
for calculating growth coefficients, temperature profiles during growth period and feed intake and
efficiency for various seasons etc. One such production records for a lot of rainbow trout from a
field station is shown in Table 11. A lot of 100 000 fish was reared over a 14-month (410 days)
production cycle between May, 1995 and June, 1996. Cumulated live weight gain (fish production)
was 72 tonnes with feed consumption of 60 tonnes which gave an overall feed efficiency (gain/feed)
of 1.19 (ranged between 1.11 1.22). Water temperature ranged from 0.5C in winter to 21C in
summer which is typical of most lakes in Ontario. In spite of the wide fluctuation in water
temperature, the thermal-unit growth coefficients (TGC) was fairly stable ranging between 0.177
0.204. Total mortality was around 9% over 410 days.
From the production record (Table 11) one can extrapolates an overall growth coefficient of
0.191 and this coefficient can be used for the growth prediction of next production cycle with
assumption of similar husbandry conditions and fish stock are used. Total feed requirement and
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22
weekly or monthly feeding standards can be computed on the basis of this growth predictions plus
the quality of feed purchased.
Table 11. - Fish production records from a field station

Month- Days
End
No.
Fish
Weight TGC
(g/fish)
Total
Biomass
(kg)
Total Gain/
Feed Feed
(kg)
Temp
Flow Rate
(C)
(L/min)
1995
Initial
May
Jun
Jul
Aug
Sep
Oct
Nov
Dec
Jan
Feb
Mar
Apr
May
Jun
TOTAL
15
30
31
31
30
31
30
31
31
28
31
30
31
30
410
days
100000
10.00
98900
95000
95000
94500
94000
93500
93200
93000
92000
91500
91200
91000
91000
90800
12.05
36.45
89.84
177.43
296.26
396.06
451.03
455.85
460.77
465.23
470.39
475.54
534.65
783.37
0.184
0.189
0.197
0.175
0.184
0.199
0.197
0.176
0.178
0.177
0.184
0.188
0.200
0.204
1191.75
167
3462.75 2000
8534.80 4300
16767.14 7200
27848.44 9500
37031.61 7800
42036.00 4300
42394.05
400
42390.84
400
42568.55
370
42899.57
420
43274.14
420
48653.15 4500
71130.00 18500
1.22
1.18
1.18
1.15
1.18
1.20
1.19
1.12
1.14
1.11
1.12
1.12
1.20
1.22
0.191
60277
kg feed
1.19
5.00
18.00
19.00
21.00
19.00
11.00
5.50
0.50
0.50
0.50
0.50
0.50
5.00
18.00
2500
6000
10000
16000
20000
25000
25000
25000
25000
25000
25000
25000
30000
50000
13.5 mill. m3
water used
Procedures for the Estimation of Feed Requirement and Waste Output
Using production records as a starting point, feed requirements and waste output can
scientifically be estimated based on the following three concepts:
1)
2)
3)
1)
Prediction of growth and nutrient and energy gains

Estimation of excretory and feed waste outputs
Quantitation of energy and nutrient needs
Prediction Of Growth And Nutrient And Energy Gains:
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23
Accurate prediction of growth potential of a fish stock under given husbandry condition is an
inevitable prerequisite to the estimation of energy or feed requirement (e.g. weekly ration). The
formula most commonly used for fish growth rate expression is instantaneous growth rate known as
"specific growth rate (SGR)" which is based on the natural logarithm of body weight:
SGR = (ln FBW - ln IBW) / D.
(1)
where
FBW is final body weight (g)
IBW is initial body weight (g)
D = number of days
SGR has been widely used by most biologists to describe growth rate of fish. However, the
exponent of natural logarithm underestimates the weight gain between the IBW and the FBW used
in the calculation and it also grossly overestimates predicted body weight at weights greater than
FBW used. Furthermore the SGR is dependent on the IBW, making meaningless comparisons of
growth rates among different groups unless IBW are similar.
A more accurate and useful coefficient for fish growth prediction in relation to water
temperature is based on the exponent 1/3 power of body weight. Such a cubic coefficient has been
applied both to mammals and to fish. The following modified formulae were applied to many
nutritional experiments:
Thermal-unit Growth Coefficient (TGC)
= [FBW1/3 - IBW1/3] / [T x D] x 100
(2)
Predicted Final Body Weight

= [IBW1/3 + (TGC/100 x T x D)]3
(3)
where:
T is water temperature (C)
(NOTE: 1/3 exponent must contain at least 4 decimals (e.g. 0.3333) to maintain good accuracy)
This model equation has been shown by experiments in our laboratory and several field
stations to represent very faithfully the actual growth curves of rainbow trout, lake trout, brown
trout, chinook salmon and Atlantic salmon over a wide range of temperatures. Extensive test data
were also presented by Iwama and Tautz (1981). An example of the relationship among growth,
water temperature and TGC is shown in Figure 11. Growth of some salmonid stocks used for our
experiments in freshwater gave the following TGC:
Rainbow trout-A
0.174
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24
Rainbow trout-B
Rainbow trout-C
Lake trout
Brown trout
Chinook salmon
Atlantic salmon-A
Atlantic salmon-B
0.153
0.203
0.139
0.099
0.098
0.060
0.100
Since these TGC values and growth rate are dependent on species, stock (genetics), nutrition,
environment, husbandry and others factors, it is essential to calculate the TGC for a given
aquaculture condition using past growth records or records obtained from similar stocks and
husbandry conditions.
Once the expected TGC and water temperature profile during the production period are
established, expected live weight gain (LWG) and recovered energy (RE), nitrogen (RN) and
phosphorus (RP) on basis of dry matter (DM, 20-35% of live body weight) in carcass can be
computed in the following manners:
LWG = FBW - IBW
RE (or RN, RP) = LWG x DM x GE (or N, P)
(4)
(5)
where
LWG is live weight gain (g)
FBW is final body weight (g)
IBW is initial body weight (g)
RE, RN, RP are the recovered energy (kJ), nitrogen (g) and phosphorus (g)
DM is dry matter content (%) of the fish
GE, N, P is gross energy (kJ), nitrogen (%) and phosphorus (%) content of dry matter
Because of a large proportion of the nutrients (e.g. protein, lipid) and, consequently of the
dietary energy, consumed by fish is retained as carcass body constituents, carcass energy gain is a
major factor driving dietary energy requirement of the fish. Carcass moisture, protein and fat
contents in various life stages dictate energy level of fish. These factors are influenced by species,
genetics, size, age and nutritional status. The dry matter and fat contents of the fish produced are, in
general, the most variable factors and have a determinant effect on energy content of the fish. For
example, relatively fatty Atlantic salmon and rainbow trout may require more dietary energy per unit
of live body weight than leaner salmonids such as brown trout, lake trout and charr. Fish containing
less moisture (more dry matter) and more fat require more energy allocation in feeding standards.
The simplistic assumption of the constant body composition within a growth stanza in
certain published models is not necessarily valid for different species and sizes. Dry matter and
energy content of fish can increase dramatically within a growth stanza, especially in the case of
small fish. Underestimation or overestimation of the feed requirement is likely to occur if constant
carcass energy content is assumed in calculations. Reliable measurements of carcass composition of
fish at various size are essential. Nutrient and energy gains should be calculated at relatively short
size intervals as possible, at least for small fish (<100 g). Additionally, composition of the diet,
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25
notably the digestible protein to digestible energy ratio and the lipid content of the diet, can have a
very significant influence on the composition and energy content of the carcass. Estimation of
carcass composition and energy content should rely on data obtained with fish fed diets similar to
those one intends to use.
2) Estimation of Excretory And Feed Waste Outputs:

Waste output loading from aquaculture operations can be estimated using simple principles
of nutrition and bioenergetics. Ingested feedstuffs must be digested prior to utilization by the fish
and the digested protein, lipid and carbohydrate are the potentially available energy and nutrients for
maintenance, growth and reproduction of the animal. The remainder of the feed (undigested) is
excreted in the feces as solid waste (SW), and the by-products of metabolism (ammonia, urea,
phosphate, carbon dioxide, etc.) are excreted as dissolved waste (DW) mostly by the gills and
kidneys.
The total aquaculture wastes (TW) associated with feeding and production is made up of
SW and DW, together with apparent feed waste (AFW):
TW = SW + DW + AFW
(6)
SW, DW and AFW outputs are biologically estimated by:

SW = [Feed consumed x (1-ADC)]
(7)
DW = (Feed consumed x ADC) - Fish produced (nutrients retained) (8)
AFW = Actual feed input Theoretical feed requirement
(9)
in which ADC is the apparent digestibility coefficients of diets. Measurements of ADC and feed
intake provide the amount of SW (settled and suspended, AFW-free) and these values are most
critical for accurate quantification of aquaculture waste. ADC for dry matter, nitrogen and
phosphorus should be determined using reliable methods by research laboratories where special
facility, equipment and expertise are available. More information on the equipment and procedures
may be obtained from the website www.uoguelph.ca/fishnutrition.
DW (N or P) can be calculated by difference between digestible N or P intake and retained N
(RN) or P (RP) in the carcass if this information is available, or by using a digested nutrient retention
efficiency (NRE = Retained/Intake). Reliable NRE are necessary and should be determined or
estimated for each type of diet used by research laboratories where expertise is available. However,
controlled feeding and growth trial(s) with particular diets at production sites are essential to validate
and fine-tune the coefficients from the laboratory. Dissolved nitrogen output depends very much on
dietary protein and energy ratio and amino acid balances and rate of protein deposition by the fish,
therefore all coefficients must be determined on a regular basis, particularly when feed formulae are
changed. Assuming constancy of many coefficients is a dangerous exercise.
Accurate estimation of total solid waste (TSW) requires a reliable estimate of AFW.
Feeding the fish to appetite or near satiety is very subjective and unfortunately TW contains a
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26
considerable amount of AFW under most fish farming operations. The use of biomass gain x
feed conversion as an estimate of real feed intake of the fish to calculate waste output used in
certain waste output prediction models can grossly overestimate the feed intake in many operation
where overfeeding is common and result in an underestimation of the TSW output.
It is very difficult scientifically to determine the actual feed intake by fish in spite of many
attempts (mechanical, radiological and biological) that have been made by biologists. Since
estimation of AFW is difficult and almost impossible, the best estimates can be made based on
energy requirements and expected gain in which the energy efficiency (energy gain/intake)
indicates the degree of AFW for a given operation. The theoretical feed requirement (TFR) can be
calculated based on nutritional energetic balance as follows:
TFR = Retained + Excreted
(10)
and the amount of feed input above the TFR should be assumed as AFW and all nutrient contents of
the AFW must be included in solid waste quantification. This approach may yield relatively
conservative estimates.
Biological procedures based on the ADC for SW and comparative carcass analyses for DW
were shown to provide very reliable estimates. Biological methods are flexible and capable of
adaptation to a variety of conditions and rearing environments. It also allows estimation of the
theoretical feed requirement and waste output under circumstances where it would be very difficult
or impossible to do so with a chemical/limnological method (e.g. cage culture). Properly conducted
biological and nutritional approaches to estimate aquaculture waste outputs are not only more
accurate but also more economical than chemical/limnological method.
The waste outputs from the field station (see Table 11) are tabulated in Table 12. SW was
estimated at 10 610 kg (fish production 72 t; 60 t feed input over 14 months). SW represented 90%
of TSW, since AFW (actual feed input theoretical feed requirement) was estimated at 1 201 kg or
2 % of feed input (60 277 kg in Table 11). The TSW outputs were equivalent to 164 kg per tonne
fish produced. Phosphorus waste was 5.11 kg/t fish produced and nitrogen 30.64 kg. Total water
consumption during 14 months was 13 469 m3, therefore the average effluent quality can be
estimated at: solid 0.877 mg/L, nitrogen 0.163 and phosphorus 0.027 (Table 12). The diet (MNR91HG) and the procedures to estimate waste production as well as comparative data of chemical and
biological estimations from field experiments at the Ontario Ministry of Natural Resources (OMNR)
Fish Culture Stations are described elsewhere (Cho et al., 1991, 1994).
26
27
Table 12 - Waste outputs and effluent quality from fish production operation in Table 1
WASTE OUTPUT
Solid
Nitrogen
Phosphorus
(kg)
(kg)
(kg)
Feed Wastage (2.2 %) *
1201
80.69
12.008
Solid
10610
356.49
212.194
1764.60
143.231
TOTAL
11811
2201.79
367.433
- per tonne fish produced

- % of dry matter fed
164.3
21.8 %
30.64
60.4 %
5.113
67.7 %
Average CONCENTRATION (mg/L)

in EFFLUENT (13469 mill. L)
during 410 days
0.877
0.163
0.027
(Total Load Estimate)
Dissolved
* Actual amount of feed fed Theoretical amount of feed required
3)
Quantitation Of Energy And Nutrients Needs

3.1) Dietary energy and protein requirements
A relatively large portion of dietary energy is expended for maintenance or basal

metabolism, which is the minimum energy and nutrients required necessary to maintain basic life
processes. Maintenance energy requirement is approximately equal to the heat production of a
fasting animal. This amount of dietary energy represent as an absolute minimum of "energyyielding" nutrients must be covered before any nutrients can be used for growth and reproduction of
the animal. Otherwise body tissues will be catabolized because of a negative energy balance
between intake of dietary fuels and energy expenditure.
A review of available data suggest that a HEf of about 36-40 kJ/kg0.824 per day appear
accurate for rainbow trout at 15C, at least for fish between 20 and 150 g live weight with which
most of studies have been conducted. Water temperature has a major influence on basal metabolism
of fish. The following equation to estimate HEf of salmonids as a function of water temperature
(10):
HEf = (- 0.0104 + 3.26T - 0.05T2) (BW 0.824 ) D -1
(11)
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28
where
HEf is fasting heat production in kJ
T is water temperature (C)
BW is body weight (kg)
D is number of days
Ingestion of food by an animal which has been fasting results in an increase in the
animal's heat production, this heat production is known as heat increment of feeding (HiE). The
physiological basis of this increased heat production includes the post-absorptive processes
related to ingested food, particularly protein-rich food and the metabolic work required for the
formation of excretory nitrogen products, as well as the synthesis of proteins and fats in the
tissues from the newly absorbed, food-derived substrates such as amino acids and fatty acids.
The HiE of rainbow trout fed a balanced diet was observed to be approximately 30 kJ/g
digestible N or the equivalent of 60% HEf (Cho and Kaushik, 1990), but these relationships do
not always hold true. Studies with farm animals suggest that HiE associated with growth may be
more appropriately quantified as a factorial function of protein and lipid deposition rates. Protein
and lipid oxidation rates also appear to contribute to HiE (Cho et al., 1982). Experimental
observations suggest that HiE is approximately equivalent of 17% of net energy intake, i.e.
0.17(RE+HEf) for rainbow trout and other salmonids. This value is used in the bioenergetic
model presented here. Studies are underway to quantify HiE as a function of protein and lipid
deposition and oxidation rates.
Biological oxygen requirement of feeding fish is equal to the total heat production (HEf +
HiE / Qox) in which the oxycalorific coefficient (Qox) used in the model is 13.64 kJ energy per
g oxygen. This represent the absolute minimum quantity of oxygen that must be supplied to the
fish by the aquatic system. Oxygen requirement per unit of BW per hour will vary significantly
for different fish sizes, water temperatures and growth rates.
3.2)
Total energy requirement and calculation of feeding standard
The calculation of total energy requirement and consequently feed allocation of the animal
can be accomplished as follows:
1.
Calculation of expected live weight gain (LWG = FBW - IBW) and
recovered energy (RE) based on carcass dry matter content (DM = 20-35% of live
body weight and gross energy (GE) contents = 25-30 kJ/g DM):
RE = LWG x DM x GE
(12)
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29
2.
Allocation of approximate maintenance or fasting energy requirement at a
given water temperature (T):
HEf = (- 0.0104 + 3.26T - 0.05T2) (BW 0.824) d-1
(11)
3.
Allocation of approximate heat increment of feeding for maintenance and
growth ration:
HiE = (RE + HEf) x 0.17
(13)
4.
5.
6.
Allocation of approximate non-fecal energy loss:

ZE + UE = (RE + HEf + HiE) x 0.09
or
(14)
Theoretical/minimum energy requirement:

TER = 1) + 2) + 3) + 4) = [(RE + HEf) x 1.2753]
(15)
Feed allowance or feeding standard:

FA = TER / DE x Qfi
(16)
Where
TER is theoretical/minimum energy requirement (MJ)
FA is feed allowance (kg)
DE is digestible energy content of the feed (MJ/kg)
Qfi is an adjustment factor
Qfi is an adjustment factor determined by the fish culturist to provide flexibility for
estimating realistic FA under a given husbandry condition (if one observes that more or less feed
may be required than predicted by the model). The minimum digestible energy requirement that
should be fed to the fish is the sum of retained energy (RE) and energy lost as HEf + HiE + ZE +
UE. The amount of feed can be estimated on a weekly or monthly basis, and recalculated if any
parameter (growth rate, water temperature, etc.) is changed. The computed quantity of feed should
be regarded as a minimum requirement under most conditions and fish culturists should fine-tune the
feeding level to own local conditions using the adjustment factor (Qfi).
The overall energy cost of producing one kg of rainbow trout is around 15-16 MJ DE, but
this ranges from 10 MJ for fry to more than 20 MJ for fish of near 3 kg. Even though maintenance
energy requirement per kg BW is much higher in small than in large fish, overall energy cost of
production is much higher in large fish because of high "growth-fattening cost". This may become
much more significant when feeding overly high energy (fat) diets, hence more than 50 kJ DE per g
DP (or less than 20 g DP/MJ DE) is not recommended. Water temperature greatly affects heat
production and oxygen consumption of poikilotherms and a growing fish of 100 g is expected to
consumed 110, 210 and 300 mg oxygen/kg BW/hr at 5, 10, 15C, respectively.
29
30
Table 13 summarizes the monthly fish sizes and feed rations predicted by the bioenergetic
models program for the field station based on the production records (see Table 11). The feed
requirements were calculated using a single TGC (0.191) for the whole production cycle (14
months) and actual temperature profile. The nutrient and energy gains used in the calculations were
based on carcass composition values for rainbow trout of various sizes obtained in different
laboratory trials at the University of Guelph. Nutrient and energy retention efficiencies (NRE and
ERE) used were derived from previous studies at another fish culture station (Harwood Fish Culture
Station, Harwood, Ontario) using comparable diets (Cho et al., 1994). The main discrepancy is
between the actual and predicted feed amount for the first four months with actual feed input being
greater than predicted allocation. This may indicate that overfeeding occurred in 1995, however,
real feed intake by the fish could be somewhere between the predicted amount and the actual
amount. Using this information, the fish culturist can fine-tune the program in the next production
cycle. In the remaining 10 month, the feed allocation estimated by the model was very close to the
actual feed fed, the largest discrepancies (in terms of predicted/actual) occurring at very low water
temperature (0.5C).
This simulation may not be considered a perfect example of independent or objective
validation of the model but is, nevertheless, an adequate demonstration of the realism of the
predictions from bioenergetic models. Most of the parameters used in the calculations are fairly
independent from the actual data. For example, the carcass composition data were from a
number of laboratory trials which had nothing to do with actual data. The TGC and the
temperature profile used in the calculation are not independent from the actual data because it is
essential to use actual values or values from previous production cycle if these are available and
repeatable. TGC and water temperature are main inputs required from the fish culturist by the
models. The predicted values from Table 11 were calculated a posteriori and their main use is
as production scenario for following year based on 1995 production performance. The predicted
values can also be used as yardsticks to compare the results obtained with what was predicted to
be biologically achievable and adjust feeding practices or equipment in the following production
cycle.
30
31
Table 13. - Prediction of fish body weight and feed requirement based on 1995 production records in
Table 11.
Month
-End
No.
Fish
TGC
(%)
Body
Weight
(g/fish)
Total
Feed
(kg)
Gain/
Feed
Ratio
Body
Weight
(g/fish)**
Total
Feed
(kg)**
Gain/
Feed
Ratio
Actual
Actual
Predicted
Predicted
167
2000
4300
7200
9500
7800
4300
400
400
370
420
420
4500
18500
1.22
1.18
1.18
1.15
1.18
1.20
1.19
1.12
1.14
1.11
1.12
1.12
1.20
1.22
Predicted
10.00
12.15
37.39
87.94
181.93
310.23
406.58
461.46
466.68
471.94
477.24
482.58
487.96
543.95
780.78
120
1498
3446
6732
9495
7775
4602
451
454
452
453
456
4627
18228
1.81
1.68
1.47
1.40
1.35
1.24
1.19
1.16
1.16
1.17
1.18
1.18
1.21
1.30
Temp
(C)
1995
Actual
Initial
100000
May
Jun
Jul
Aug
Sep
Oct
Nov
Dec
Jan
Feb
Mar
Apr
May
Jun
98900
95000
95000
94500
94000
93500
93200
93000
92000
91500
91200
91000
91000
90800
10.00
0.184
0.189
0.197
0.175
0.184
0.199
0.197
0.176
0.178
0.177
0.184
0.188
0.200
0.204
12.05
36.45
89.84
177.43
296.26
396.06
451.03
455.85
460.77
465.23
470.39
475.54
534.65
783.37
5.0
18.0
19.0
21.0
19.0
11.0
5.5
0.5
0.5
0.5
0.5
0.5
5.0
18.0
** Overall TGC = 0.191 from Table 1 was used to predict body weight and total feed
requirement
31
32
Development of Fish-PrFEQ Computer Program
A stand-alone multimedia program for the Windows 95 platform was developed in visual
basic language with database functionality by the Ontario Ministry of Natural Resources. The
program has 4 modules for fish production/growth prediction, waste output quantification, feed
allowance estimation and oxygen requirement table and is based on the bioenergetic models
presented above. Feed composition, body weight, water temperature, flow rate and mortality are
entered by the user but waste, retention and other coefficients are parameters that are locked and
may only be revised with an authorized program update diskette. These coefficients should be
determined by qualified nutritionists from feed manufacturers or research institutions since specific
coefficients are required for each type of diets. The use of unrelated coefficients result in under or
overestimation of feed requirements and waste outputs. Live weight gain, feed efficiency, growth
coefficients, solid, nitrogen, phosphorus in the effluent, total waste load, feed ration and oxygen
requirements are some of the output parameters generated by the models.
Factors Affecting Feed Utilization
Feed costs represent a very significant proportion of the production cost in salmonid fish
culture. Many fish culture operations have poor feed efficiencies (gain/feed) and this contributes
to the high cost of production and often results in significant water pollution. It is necessary to
optimize feeding regimes to improve the economical and environmental sustainability of
aquaculture.
It is not always clear if low feed efficiencies observed under certain conditions are due to
feed wastage or due to a real decrease in feed utilization efficiency of the fish. The effect of
feeding level on the efficiency of feed utilization in rainbow trout and other salmonids is the
subject of controversy. It has been suggested that optimum feed efficiency is achieved at feeding
levels below that required for maximum growth in salmonids. Other studies suggest that feed
efficiency improves to its maximum at moderate feed restriction (e.g. 50% of maximum ration)
and this optimum is maintained up to the ration required for maximum growth of the fish. It has
also been suggested that maximum feed efficiency of fish is attained at maximum feed intake
and maximum growth. Most of these observations are derived from studies using fixed ration
(% live body weight) which may not represent the fishs actual feed requirement or studies
conducted under poorly controlled experimental conditions (mechanical distribution of feed,
variable temperature, etc.).
While important from a production point of view, feed efficiency can be a misleading
expression of nutrient and energy utilization. Physical quantity of feed used is not a measure of
biologically available nutrients and energy supplied to the animal. In addition, weight gain does
not always accurately reflect protein, lipid and energy gains since the composition of weight gain
is often variable. Protein deposition is associated with substantial water deposition whereas lipid
depots contain little water. The ratio between protein and lipid deposition will have an impact
32
33
on live weight gain and, consequently, feed efficiency.

Being poikilothermic animals, the metabolic rate, growth, energy expenditure, and feed
intake of fish are highly influenced by water temperature. It is, therefore, important to study
how water temperature affects these parameters, as well as to determine the effect of temperature
on the efficiency of nutrient and energy utilization. Studies have suggested that temperature can
affect the efficiency of energy utilization in salmonids.
The effect of feeding level and water temperature on feed utilization was recently reexamined under highly controlled conditions (careful hand-feeding to avoid feed waste,
controlled temperature, etc.). The results from the study indicated that fish consuming more feed
as a result of an increase in water temperature or an increase feed allocation grew faster but
appeared to utilise digestible nutrients with similar efficiencies (Table 14, Figure 2). Feeding
level or water temperature had very little effect on feed efficiency. The main factors affecting
feed efficiency of fish fed a balanced practical diet under practical is, therefore, feed wastage.
Feeding frequency and timing is another factor that has been suggested as affecting feed
intake and utilization by fish. On a weekly basis, studies have suggested that feeding the
equivalent of six days a week resulted in growth performance similar to feeding 7 days a week.
Feeding five days a week resulted, however, in significantly less growth (Table 15). There is no
good evidence that daily feeding frequency and timing affect feed utilization. The most
important factor is to insure frequent and spaced enough meals to insure that the animal can
consumed enough feed to meet its growth potential. This generally means more frequent feeding
for fish of smaller size. Table 10 provides informal guidelines for daily feeding frequency (# of
meal/day) as a function of fish size. There is, in general, slight between and within day
variations in the appetite of fish, especially if they are free to choose when to feed (i.e. with a
demand feeder). Fish will, however, easily adapt to a feeding schedule. Being attentive to
changes in appetite of fish is, nevertheless, a very important skill fish culturist must acquire.
33
34
Table 14. Growth performance and feed efficiency of rainbow trout (IBW=13.3 g/fish) fed the
experimental diet for 12 weeks at 3 feeding levels and 4 water temperatures. N = 3 for each
feeding level within temperature and for each temperature.
Water
temp.
Feeding
level
6oC
NS
R1
R2
SEM
HSD
NS
R1
R2
SEM
HSD
NS
R1
R2
SEM
HSD
NS
R1
R2
SEM
HSD
9oC
12oC
15oC
Weight
gain
(g/fish)
24.8 a z
20.4 b
17.5 c
0.32
1.60
47.4 a y
37.7 b
32.5 c
0.58
2.91
71.3 a x
58.5 b
49.9 c
1.23
6.21
96.8 a w
74.4 b
63.9 c
2.06
10.38
Feed
intake1
(g/fish)
23.0 a z
18.2 b
15.3 c
0.16
0.82
39.8 a y
30.0 b
25.7 c
0.24
1.21
62.0 a x
47.4 b
40.1 c
0.55
2.75
86.3 a w
63.6 b
53.7 c
1.52
7.67
FE2
(gain : feed)
1.15 a z
1.20 a
1.22 a
0.024
0.12
1.27 a y
1.34 a
1.34 a
0.022
0.11
1.22 b y z
1.31ab
1.32 a
0.019
0.10
1.19 a y z
1.25 a
1.26 a
0.027
0.12
Thermal - unit Growth

Coefficient (TGC) (%)
0.188 a z
0.163 b
0.143 c
0.0016
0.0081
0.204 a z
0.175 b
0.159 c
0.0013
0.0065
0.192 a z
0.172 b
0.159 c
0.002
0.010
0.191 a z
0.164 b
0.149 c
0.0026
0.0129
NS = near satiation, R1, R2 = restricted diets. SEM = standard error of mean. HSD = Tukeys
honestly significant difference (P < 0.05). Means in the same column (within each temperature)
with different superscripts (a, b, c) are statistically different (P < 0.05). The superscripts w, x, y,
z are used for comparing results between temperatures at the NS feeding level (P < 0.05).
1
weight as fed basis, 2FE = wet weight gain / dry feed.
34
35
RE (kJ/kg MBW)
Figure. 2 Efficiency of metabolizable energy utilization above maintenance by rainbow trout at

various water temperatures and feeding levels.
250
6C
200
9C
150
12C
100
15C
50
0
0
50
100
150
200
ME intake - HeE (kJ/kg MBW)
Table 15. Effect of weekly feeding frequency on growth and feed efficiency of rainbow trout.
No. of days fed / week
Gain, g/fish
Feed efficiency, gain:feed
6*
5**
SEM
HSD
84.9a
0.70
86.2a
0.70
70.0b
0.70
2.2
13.1
Initial weight = 6.1 g/fish

n= 2 tanks per treatment, Duration = 168 days, water temperature = 15C
* No feeding on Sunday
** No feeding on Saturday and Sunday
SEM = pooled standard error of a mean
HSD = Tukeys Honestly Significant Difference
Mean in the same row not sharing the same subscript are significantly different (P<0.05).
35
36
Reference and Suggested Readings
Azevedo P.A., Cho C.Y., Bureau D.P., Effects of feeding level and water temperature on growth,
nutrient and energy utilization and waste outputs of rainbow trout (Oncorhynchus mykiss). Aquat.
Living Resour. 11 (1998) 227-238 .
Cho, C.Y., Bureau, D.P., Development of bioenergetic models and the Fish-PrFEQ software to
estimate production, feeding ration and waste output in aquaculture. Aquat. Living Resour. 11
(1998) 199-210.
Cho C.Y., Bureau D.P., Reduction of waste output from salmonid aquaculture through feeds and
feeding. Progress. Fish. Cult. 59 (1997) 155-160.
Cho C.Y., Cowey C.B., Watanabe T., Finfish nutrition in Asia, Methodological approaches to
research and development. International Development Research Centre, Ottawa. Publication No.
IDRC-233e (1985) 154 p.
Cho C.Y., Feeding systems for rainbow trout and other salmonids with reference to current estimates
of energy and protein requirements. Aquaculture 100 (1992) 107-123.
Cho C.Y., Fish nutrition, feeds, and feeding with special emphasis on salmonid aquaculture. Food
Rev. Int. 6 (1990) 333-357.
Cho C.Y., Hynes J.D., Wood K.R., Yoshida H.K., Development of high nutrient-dense, low
pollution diets and prediction of aquaculture wastes using biological approaches, Aquaculture 124
(1994) 293-305.
Cho C.Y., Hynes J.D., Wood K.R., Yoshida H.K., Quantitation of fish culture wastes by biological
(nutritional) and chemical (limnological) methods; the development of high nutrient dense (HND)
diets, In: Cowey C.B., Cho C.Y. (eds) Nutritional Strategies and Aquaculture Waste, Proceedings
of the 1st International Symposium on Nutritional Strategies in Management of Aquaculture Waste,
University of Guelph, Ontario, Canada (1991) pp.37-50.
Cho C.Y., Kaushik S.J., Nutritional energetics in fish: energy and protein utilization in rainbow trout
(Salmo gairdneri), World Rev. Nutr. Diet. 61 (1990) 132-172.
Cho C.Y., Slinger S.J., Bayley H.S., Bioenergetics of salmonid fishes: Energy intake, expenditure
and productivity, Comp. Biochem. Physiol. 73B (1982) 25-41.
Iwama G.K., Tautz A.F., A simple growth model for salmonids in hatcheries, Can. J. Fish. Aquat.
Sci.,38 (1981) 649-656.
Shearer K.D., Factors affecting the proximate composition of cultured fishes with emphasis on
salmonids, Aquaculture 11 (1994) 63-88.
36

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1. Protein and Amino Acid Requirements of Fish

The protein requirement in terms of dietary concentration (% of diet) is high but

Indispensable amino acid requirements

10 Indispensable amino acids

Carbohydrates represent a very large variety of molecules. The carbohydrate most

Water-soluble vitamin, mg/kg

Vitamin-like compounds, mg/kg

Table 4. Deficiency signs associated with various nutrients.

Folic Acid, Inositol, Niacin, Pyrodoxine, Rancid Fat

Biotin, Folic Acid, Inositol, Niacin, Pantothenic Acid

Vitamin A, Vitamin C, Vitamin E

Pyrodoxine, Pantothenic acid, Riboflavin

Methionine, Riboflavin, Thiamin, Zinc

Rancid Fat, Vitamin E

Methionine, Riboflavin, Zinc

Clotting blood: slow

Colouration: dark skin

Biotin, Folic Acid, Pyrodoxine Riboflavin

Biotin, Pyrodoxine, Thiamin

Fatty Acids, Thiamin

Niacin, Pyrodoxine, Thiamin, Vitamin A, Vitamin E

Fatty Acids, Riboflavin, Vitamin A, Zinc

Pyrodoxine, Vitamin A, Vitamin C, Vitamin E

Biotin, Choline, Fatty Acids, Inositol, Vitamin E

Feed efficiency: poor

Biotin, Calcium, Choline, Energy, Fat, Folic Acid,

Biotin, Vitamin B12, Vitamin E

Biotin, Vitamin B12, Vitamin E

Biotin, Calcium, Choline, Energy, Fat, Folic Acid

Iron, Vitamin C, Vitamin E

Iron, Vitamin B12, Vitamin C

Choline, Vitamin A, Vitamin C

Niacin, Pantothenic Acid, Riboflavin, Vitamin A, Vitamin C

Fatty Acids, Pyrodoxin, Thiamin

Methionine, Riboflavin, Vitamin A, Vitamin C, Zinc

Biotin, Inositol, Niacin, Pantothenic Acid

Folic Acid, Niacin, Pantothenic acid, Thiamin

Fatty Acids, Rancid fat

Essential Fatty Acids

Pale liver (glycogen accumulation)

High Digestible Carbohydrate, Biotin

Pantothenic Acid, Vitamin C

Rigor mortis, rapid

Phosphorus, Tryptophan, Vitamin C, Vitamin D

Essential Fatty Acids

Swimming, upside down

Tetany, white muscle

Table 5. Mineral requirement of salmonid fish in freshwater.

Requirement (mg/kg feed)*

Digestive tract anatomy and physiology

teeth, no chewing, taste buds

pharyngeal teeth (calcified structures)

short, thick, taste buds, gizzard

present or absent, acid, enzymes, rate of digestion correlates with

secretive and absorptive epithelial cells, no villi but numerous

present in some case, variable # between species and individuals

Generally a diffuse tissue, except for eel, pike, flat fish

Not really morphologically distinct from anterior intestine but cell

Various digestive configurations

Estimates of apparent digestibility for salmonids

Table 6. Apparent digestibility coefficients of ingredients measured with rainbow trout.

7. Feed Formulation and Manufacturing