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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
Introduction
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 ensue. 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
1.1 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 incorporates 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 is 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 848k1018i 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 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 isn't (g/kg body weight gain). This is due to the fact that fish have a lower absolute energy requirement than mammals. These 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 is 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.
1.2 Indispensable amino acid requirements
10 Indispensable amino acids Phenylalanine (Phe), Histidine (His), Isoleucine (Iso), Leucine (Leu), Lysine (Lys), Methionine (Met), Tryptophan (Trp), Valine (Val), Arginine (Arg), Threonine (Thr)
Table 1. Indispensable amino acid requirements of different species of teleost (g / 100 g protein)
Amino acids |
Salmonid |
Catfish |
Carp |
Tilapia |
Milkfish |
Sea Bream |
Sea Bass |
Arg | |||||||
His | |||||||
Ile | |||||||
Leu | |||||||
| |||||||
Thr | |||||||
Trp | |||||||
Val | |||||||
Met+Cys | |||||||
Phe+Tyr |
Table 2. Amino acid composition of common protein sources (g/ 100 g protein).
|
CP |
Met |
|
Trp |
Thr |
Ile |
His |
Val |
Leu |
Arg |
Phe |
Requirement | |||||||||||
Fish meal | |||||||||||
Soybean meal | |||||||||||
Corn gluten meal | |||||||||||
Blood meal | |||||||||||
Meat and bone meal | |||||||||||
Poultry by-product meal | |||||||||||
Feather meal |
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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 bi-layer). 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 achieve proper physical characteristic of the feed.
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 fat-soluble 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 | |
Water-soluble vitamin, mg/kg | |
Riboflavin | |
Pantothenic acid | |
Niacin | |
Vitamin B12 | |
Biotin | |
Folate | |
Thiamin | |
Vitamin B6 | |
Vitamin C | |
Vitamin-like compounds, mg/kg | |
Choline | |
myo-Inositol |
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 |
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 |
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 |
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 |
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 are the most common signs of mineral deficiencies.
Table 5. Mineral requirement of salmonid fish in freshwater.
Mineral |
Requirement (mg/kg feed)* |
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 in the absence of significant amounts of the specific mineral in the water.
This page contains material produced by the Fish Nutrition Research Lab,
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