Other Trace Elements

Information on the dietary requirements of other trace elements is limited. Increased dietary fluoride enhanced fluoride
accumulation in the vertebrae of rainbow trout (Tiews et al., 1982; Bowser et al., 1988). Evidence also indicated that elevated
concentrations of fluoride may reduce the prevalence of bacterial kidney disease (Lall et al., 1985; Bowser et al., 1988). The
importance of chromium and other trace elements essential for other animals and humans is also recognized in fish nutrition,
although the effects of their deficiencies have not been reported. Tacon and Beveridge (1982) found that rainbow trout fed a
low-chromium chemically defined diet did not show deficiency signs or change in tissue chromium distribution.
VITAMINS
Vitamins are organic compounds, distinct from amino acids, carbohydrates, and lipids, that are required in trace amounts
from an exogenous source (usually the diet) for normal growth, reproduction, and health. Vitamins are classified as water-
soluble and fat soluble. Eight of the water-soluble vitamins are required in relatively small amounts, have primarily
coenzyme functions, and are known as the vitamin B complex. Three of the water-soluble vitamins, choline, inositol, and
vitamin C, are required in larger quantities and have functions other than coenzymes. Vitamins A, D, E, and K are the fat
soluble vitamins that function independent of enzymes or, in some cases such as vitamin K, may have coenzyme roles. In
mammals the absence of vitamins leads to characteristic deficiency diseases, but in fish such diseases are less specifically
identified.
Some vitamins may be synthesized from other essential nutrients to spare a portion of the dietary requirement. For
example, channel catfish appear to synthesize choline if adequate methyl donors such as methionine are present in the diet;
however, if the concentration of dietary methionine is limiting, a choline requirement can be demonstrated (Wilson and Poe,
1988). An exogenous source of some water-soluble vitamins for certain warm-water fish has been shown to be derived from
microorganisms in the gastrointestinal tract (Limsuwan and Lovell, 1981; Lovell and Limsuwan, 1982; Burtle and Lovell,
1989). In cold-water carnivorous fish microorganisms are not a significant source of vitamins (Hepher, 1988).
Both qualitative and quantitative vitamin requirements of fish have been determined by feeding chemically defined diets
deficient in a specific vitamin. The quantitative requirements for most of the vitamins have been established for chinook
salmon, rainbow trout, common carp, channel catfish, and yellowtail, while only some of the requirements are known for red
sea bream and tilapia. Qualitative requirements
DIETARY REQUIREMENTS 21
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Nutrient Requirements of Fish
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have been identified in several other species. The requirements are affected by size, age, and growth rates as well as by
various environmental factors and nutrient interrelationships. Thus, different researchers have reported fairly wide ranges in
requirement values for growth in the same species (see Table 1-15). Recent studies with spring chinook salmon indicate that
the dietary requirements for certain vitamins may be lower than previously reported for this species (Leith et al., 1990). In
addition, the requirement values listed in Table 1-15, as determined by maximum liver storage or based on certain enzyme
data, are often much higher than the requirement values based on weight gain and absence of deficiency signs; therefore,
professional judgment must be used in selecting which requirement value best fits the user's needs. Thus, more studies are
needed to refine the requirements for various species for normal growth, health, and enhancement of defense mechanisms, as
suggested by Ikeda (1985). A summary of vitamin deficiency signs reported in several cultured fishes are presented in
Appendix Table A-3. Further information on vitamin nutrition research in fishes is discussed by Halver (1989).
Fat-Soluble Vitamins
The fat-soluble vitamins, A, D, E, and K, are absorbed in the intestine along with dietary fats; therefore, conditions
favorable for fat absorption also enhance the absorption of fat-soluble vitamins. Fat-soluble vitamins are stored by animals if
dietary intake exceeds metabolic needs. Thus, animals can accumulate enough fat-soluble vitamins in their tissues to produce
a toxic condition (hypervitaminosis). This has been demonstrated in the laboratory with trout for vitamins A, D, and E, but it
is unlikely to occur under practical conditions (Poston et al., 1966; Poston, 1969a; Poston and Livingston, 1969).
Since fat-soluble vitamins can be stored in the body, the nutritional history of experimental fish prior to their use in
requirement studies becomes critical. The time required to deplete fish of their stored fat-soluble vitamins is highly variable.
Differences in vitamin intake prior to an experiment may be responsible for some of the conflicting findings on the induction
and severity of deficiency signs.
VITAMIN A
Vitamin A is required in vertebrates for the regeneration of the light-sensitive compound rhodopsin in the retina of the
eye. Vitamin A has also been shown to be essential for proper growth, reproduction, resistance to infection, and the
maintenance of differentiated epithelia and mucus secretions. Blomhoff et al. (1992) have presented a recent review of
metabolic functions of vitamin A in vertebrates.
Vitamin A occurs in three forms: as an alcohol (retinol), an aldehyde (retinal), and an acid (retinoic acid). Vitamin A 1
(retinol) is found in mammals and marine fishes, whereas both vitamin A 1 and vitamin A2 (3-dehydroretinol) are found in
freshwater fishes (Braekkan et al., 1969; Lee, 1987). In freshwater fish, the oxidative conversion of retinol to 3-
dehydroretinol occurs (Goswami, 1984) as well as the reversible oxidation and reduction reactions of retinol to retinal and of
3-dehydroretinol to 3-dehydroretinal (Wald, 1945-1946). For example, tilapia has been shown to convert dietary retinol into
3-dehydroretinol and retinal into 3-dehydroretinal (Katsuyama and Matsuno, 1988). Channel catfish were found to convert β-
carotene to vitamin A1 and A2 in about a 1:1 ratio (Lee, 1987).
Cold-water fish can use β-carotene as a vitamin A precursor (Poston et al., 1977). Dupree (1970) found that channel
catfish could use β-carotene as a vitamin A source only if the dietary concentration exceeded 2,000 international units per
kilogram (IU/kg). It has recently been shown that β-carotene and canthaxanthin can be biotransformed in the liver of tilapia
into vitamin A1 and that dihydroxycarotenoids such as astaxanthin, zeaxanthin, lutein, and tunaxanthin were directly
bioconverted into vitamin A2 (Katsuyama and Matsuno, 1988). In mammals, carotenoids have been found to fulfill various
biological functions independent of vitamin A (Olson, 1989). Thus, more studies are needed on the metabolic role of
carotenoids in fish and on the possibility that carotenoids serve as a provitamin A.
Vitamin A deficiency in rainbow trout causes anemia, twisted gill opercula, and hemorrhages in the eyes and base of
fins (Kitamura et al., 1967a). Brook trout exhibited poor growth, high mortality, and eye lesions, such as edematous eyes,
displaced lens, and degeneration of the retina, when fed a vitamin A-deficient, purified diet from first feeding (Poston et al.,
1977). Channel catfish fed 0.4 mg of β-carotene/kg of diet for 3 years developed exophthalmia, edema, and hemorrhagic
kidney (Dupree, 1966). Anorexia, pale body color, hemorrhagic skin and fins, exophthalmia, and twisted gill opercula
occurred in common carp fed a vitamin A-deficient diet after 8 to 11 weeks (Aoe et al., 1968). Rapidly growing yellowtail
fingerlings fed a vitamin A-deficient diet developed deficiency signs in 20 days including arrested growth of gill opercula,
dark pigmentation, anemia, and hemorrhage in the eyes and liver, accompanied by high mortality (Hosokawa, 1989).
High dietary intake (2.2 million IU/kg diet) of retinyl palmitate caused slow growth, anemia, and severe necrosis of the
caudal fin of brook trout at 8.3°C (Poston et al., 1966). Feeding up to 2.5 million IU retinyl palmitate to trout at 12.4°C also
reduced body fat and liver size (Poston, 1971a). A high intake of dietary protein (Poston and Livingston, 1971) or methionine
(Eckhert and Kemmerer, 1974) by young trout reduced the toxicity of excess dietary vitamin A observed in fish fed a low-
protein diet.