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November | December 2010 Feature title: Niacin: one of the key B vitamins for sustaining healthy fish growth and production

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Niacin: one of the key B vitamins for sustaining healthy fish growth and production by Simon J Davies and Mark Rawling, Aquaculture Nutrition & Health Group, Plymouth University, United Kingdom

n 1951 Dr John E Halver of the School of Fisheries Science, University of Washington, USA presented the ‘model semi-purified fish diet’ to the aquatic nutrition research community. This innovation allowed for the proliferation of deficiency studies with mainly salmonid fish such as rainbow trout and Pacific salmon to evaluate the significance of vitamins in complete diets for cultured fish.

to have a loss of appetite and poor food conversion (food intake to body weight ratio) that progressed into a darker skin colour, anorexia, posterior gut lesions, oedema of the stomach and intestine, erratic motion and at-rest muscle spasms. In the late 1950s and 1960s, a predilection to sunburn in fish was discovered and, in carp, subcutaneous haemorrhages developed under chronic and acute niacin deficiency. In the 1970s, eels were found to develop skin lesions and display erratic swimming, while lesions, deformed jaws, and anaemia were discovered in catfish, Ictalarus punctatus. The period from 1980 to date encompassed a series of investigations that augments earlier knowledge, but there have been relatively few studies in the early 21st century except for the work of Shaik Mohammed et al. (2001) where pathological effects of niacin deficiency similar to this described above were reported from studies with Indian catfish (Heteropneustes fossilis).

fish nutrition. In mammals, there is a recognised and documented conversion pathway from tryptophan to niacin, thus allowing tryptophan, and proteins rich in tryptophan, to be an important reservoir for niacin biosynthesis. Although the essential amino acid tryptophan is a precursor of niacin, this endogenous synthesis, comprising 13 steps in a metabolic sequence is not deemed efficient. Studies in man have shown that approximately 60 mg of tryptophan are required to produce 1 mg of niacin and this ratio varies considerWith such an ‘ideal’ diet, vitamins could ably within different vertebrate groups. easily be assayed by using this vitamin test Fish, however, may even lack this converdiet, consisting of ‘vitamin free’ carbohydrate sion capacity or have very a poor efficacy for and protein sources i.e. casein, purified gelatin, this metabolic pathway. By supplementing potato starch, hydrogenated cotton seed oil, both a niacin deficient and niacin complete alpha-cellulose flour, minerals, cod liver oil, diet with varying amounts of tryptophan, it combined with crystalline vitamins. Each vitawas previously determined that tryptophan min could then be systematically assessed by levels have no effect on niacin accumulation. selective exclusion from this advanced basal Serrano and Nagayama (1991) found that the diet formulation. The water soluble vitamins 3-hydroxyanthranilic acid (3-HAA) to picolinic such as the B-complex and especially vitamin acid carbolase (PC) activity ratio in rainbow C (ascorbate) were all found to be essential trout suggested an ineffective conversion from in fish as in other terrestrial animals of comtryptophan to niacin. This finding will help mercial importance and indeed having the Metabolic considerations same basic functions as in humans. Exogenous proteins within the diet supply explain higher niacin requirements for some The role of niacin (vitamin B3) is no the metabolic pool with essential and non- fish, as others do carry the capacity in some less important within aquatic species; as fish essential amino acids. Among these is tryp- degree but this cannot be an insurance against farming became more prevalent, the health tophan which has considerable importance in providing a separate dietary supply. Niacin and niacinamide status of stocks fluctuare required by ated due to the wide Figure 1: Niacin in its two biologically active forms as presented to fish for assimilation all living cells spectrum of feed forand their chemimulations at that time. cal structure A number of negais depicted in tive symptoms were Figure 1. attributed to niacin They are deficiency and steps essential comwere taken to protect ponents of two against them based on coenzymes, early evidence. niacinamide In the 1940s and Nicotinic Acid Nicotinamide adenine dinucle1950s fish were found 26 | International AquaFeed | March-April 2013

FEATURE drates, lipids and proteins at the cross roads of metabolism and vital for energy production from these nutrients and protein turnover. With respect to genomic stability, the need for niacin seems most imminent when the organism is under genotoxic or oxidative stress, with particular reference to UV exposure of the animal (Hageman & Stierum, 2001). A deficiency of niacin will result in an increase or disrepair of DNA nicks within chromosomes, and conseFigure 2: Niacin requirements for selected aquatic quent increase in chromoanimal species (from compiled literature sources) somal breakage, and a heightened sensitivity to mutagens otide (NAD), and niacinamide adenine dinu- (Fenech, 2002). In general, fish with niacin cleotide phosphate (NADP) that are involved deficiencies displayed an increased risk of sunin numerous enzymatic pathways especially burn when under even natural UV radiation. In the expanding aquaculture industhose involving energy mediation and protein synthesis and degradation. More than 40 try, feed conversion ratios must be optibiochemical reactions have been identified mised in order for production costs to as being dependent on these coenzymes as be minimised. Greater efficiency present co-factors. Their major function is the removal throughout culturing conditions will lead of hydrogen from specific substrates and the to shorter growing time and a greater transfer of hydrogen to another coenzyme. demand for micronutrients such as vitaReactions in which NAD and NADP are mins. Surplus nutrients, such as vitamins involved include the metabolism of carbohy- supplied above levels useful to the species,

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can be removed from the diet if exact requirements are met. In the past, many vitamins have been included in excess of recommended levels to be certain that the requirements were fully complied (NRC 2011). However, studies have reported excess niacin can inhibit growth (Poston & Lorenzo, 1973; Poston & Combs, 1980); conversely, sub-optimal absorption of nutrients can be avoided if requirements are correctly defined and adequately presented in feed. For maximal efficiency of production, target provisions of all essential nutrients, as specified through research, must be provided through additional mineral and vitamin supplementation. If levels are unknown, further research is needed to clarify the degree of vitamin fortification necessary to maintain health and production for all phases of rearing and conditions. In relation to the other water-soluble vitamins, niacin requirements in fish procure a ranking amongst the highest needs, with the exception of choline (NRC, 2011). While many other vitamins are synthesised from precursor compounds obtained through feed ingredients, in aquatic animals, niacin is usually obtained solely through niacin presented in the diet.

Niacin requirements Caution must be expressed due to the

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FEATURE

variety of methodological approaches used in ascertaining vitamin requirement levels. In many cases, age and genetic strain of the species varies together with the pre-nutritional history of the aquatic animal under investigation. In particular, the nature of the carbohydrate component employed in experimental diets is not fully reported in the scientific literature. For example, it is well known that the carbohydrate level and complexity may influence the requirement of niacin in terms of processing of dietary energy (Shiau & Suen, 1992). This may be evident when raw materials are subjected to extrusion processing in which carbohydrates such as starch in cereals may undergo gelatinisation yielding dextrin and thereby increasing the digestible energy value of the carbohydrate fraction. It was found that for hybrid tilapia that the niacin requirements for fish fed glucose or dextrin as the carbohydrate energy source was 26 and 125 mg/Kg diet respectively. Previous formulations of fish diets often failed to address the true bioavailability of micronutrients present in fish feed ingredients pursuant to a limited common database describing this knowledge. The general niacin requirements for different species are shown in Figure 2 and these vary considerably depending on many factors. Dietary requirements have been reported to range from just 1-5 mg per kg of feed for rainbow trout to

150-200 mg for pacific salmon and 14 mg per kg for channel catfish. Clearly much will depend on the carnivorous, omnivorous or herbivorous nature of the fish species in question and rearing conditions. Investigations on Gilthead sea bream (Sparus aurata) by Morris and Davies (1995) and by Morris et al. (1995), where the qualitative and quantitative requirements for this important marine fish were first established using semi-purified diet ingredients similar to the Halver concept. The minimum nicotinic acid requirement for sea bream was determined to be 52 mg/ Kg to achieve optimum growth performance and 25 mg/kg for normal haematological balance and liver to body weight ratio. In 1997, Shiau reported parallelism between the niacin requirement of warm water fish and a varying source of dietary carbohydrate. In general, certain warm water fish, namely carnivorous species, utilise dietary carbohydrate poorly and it is recognised that carbohydrate obtained from different sources may not be equally available to all fish of the same species. There is merit for consideration of the changes in protein level, quality, and protein to energy ratio for optimum vitamin levels to be recommended. Modern fish diets are much higher in energy, presented as oil for carnivorous fish, whilst carbohydrate in the 28 | International AquaFeed | March-April 2013

form of starch is quite acceptable for omnivores such as tilapia and carp. Niacin is given special importance in this area due to its relevancy in the metabolism of protein and the release of energy from these nutrients as stated previously. However implications towards dietary requirement and variability, warrants a need to establish additional scientific information regarding the digestibility of niacin and subsequent availability coefficients within varying diets formulations based on practical ingredients. From the data of Ng et al. (1998), it was suggested that niacin supplementation can be reduced to a more efficient level due to the relatively high amount of biologically available niacin found in typical feed ingredients used in modern fish feed formulations. However, the provisions may not be adequate to meet current safety margins to guarantee production and health criteria for all species. Also, the inability to utilise particular fish feeds due to varying dietary constraints would justify continued supplementation and refinement. In addition, it was found that the bioavailability of niacin increased by some 57 percent when corn meal was extrusion cooked rather than administered in the diet in its native form. This suggests that processing technology is an important area for further investigation for determining the

FEATURE optimum inclusion levels of niacin for a range of aquatic species.

changing feed formulations where plant by products are increasingly being incorporated.

Stability and processing losses

Selected References

Niacin is regarded as a highly stable vitamin in animal nutrition and is usually added to feed as nicotinic acid or nicotinamide within the vitamin premix formulations within a dry mixture with a carrier material along with other vitamins and possibly mineral supplements as well. The advent of high energy and nutrient dense feeds in many countries engaged in intensive fish farming operations has also placed a higher burden on maintaining the health of fish, whilst promoting faster growth rates and efficient feed utilisation. The use of expanded and extruded feeds offer more scope in feeding management but may greatly influence the levels of vitamins available to fish under various conditions. Extrusion of diets has the tendency to reduce the activity of vitamins especially those within the water soluble class and the processing of raw materials may lead to serious losses. Generally this is in the order of 10-20 percent for most vitamins reported (Tacon, 1985, Gabaudan and Hardy, 2000). Further reductions are caused by storage of pelleted feed and this may result in impairment to fish health and production efficiency over extended time.

Future perspective Indeed, the movement towards new fish species in aquaculture such as flounders; turbot, sole and halibut as well as sea bass and sea bream in Europe, cobia in the USA and Brazil have generated considerable interest in producing specific diets that can meet their individual requirements for growth, development and health. Much is known about the gross nutritional requirements of these emerging species but little on vitamins, especially niacin. Intensive rearing conditions (i.e. UV light exposure to outdoor pens) and husbandry related factors may adversely affect the physiological status of fish and induce metabolic stress causing tissue damage and impaired performance. The potential of niacin supplementation in reducing such effects could prove a valuable area for future investigation. It is evident that the vitamin requirements of fish are subject to numerous factors. Recent advances in our understanding of aquatic animal biochemistry and physiology together with aquafeed technology increase the advantageous value of a thorough reexamination of the vitamin requirements of fish. This is particularly pertinent for niacin given its role in aquatic animal nutrition. There is a paucity of information in the literature for niacin in fish compared to other vitamins, and this matter needs to be addressed in the light of new candidate species for aquaculture and

Fenech, M. (2002). “Genomic Stability: a new paradigm for recommended dietary allowances (RDA’s).” Food and Chemical Toxicology. vol. 40. pp 1113-1117. Gaubadan, J and Hardy R. W. (2000). Vitamin sources for fish feeds pp, 961-965 In Encyclopaedia for Aquaculture, R. R. Stickney, Editor, New York, John Wiley and Sons, Inc. Hageman, G.J. and Stierum, R.H. (2001). “Niacin, poly (ADP-ribose) polymerase-1 and genomic stability.” Mutation Research. – Fundamental and Molecular Mechanisms of Mutation. vol 475. nos 1-2. pp 45-56. Halver, J.E. (1957). “Nutrition of salmonid fishes: 3. Water-soluble vitamin requirements of Chinook salmon.” Journal of Nutrition. vol. 62. pp. 225-43. Halver, J.E., (Halver, J.E. and Hardy, R.W. (Editors). Fish Nutrition. 3rd Edition. Oxford: Academic Press, 2002. Morris, P.C. and Davies, S.J. (1995) The requirement of the gilthead sea bream (Sparus aurata L). for nicotinic acid. Animal Science, 61: 437-443 Morris, P.C. Davies, S.J. and Lowe, D.M. (1995) Qualitative requirements for B vitamin in diets for the gilthead sea bream (Sparus aurata) Animal Science, 61; 419-426. Ng, W.K, Serrini, G., Zhang, Z and Wilson, R.P. (1997) “Niacin requirement and inability of tryptophan to act as a precursor of NAD+ in channel catfish, Ictalurus punctatus” Aquaculture. vol. 154. nos. 1-4. pp 273-285. NRC (2011) “Nutrient Requirements of Fish,” NAS/NRC, Academic Press, Washington D.C. Poston, H.A. (1969) “The effect of excess levels of niacin on the lipid metabolism of fingerling brook trout.” In: Fisheries Research Bulletin, Albany, N.Y.: State of New York Conservation Department. no. 32. pp. 9-12.

Poston, H.A. and DiLorenzo, R.N. (1973) “Tryptophan conversion to niacin in the brook trout (Salvelinus pontinalis).” Proceedings. Society for Experimental Biology and Medicine. vol. 140. pp. 110-12. Poston, H.A. and Combs, G.F. (1980) “Nutritional implications of tryptophan catabolising enzymes in several species of trout of salmon,” Proceedings. Society for Experimental Biology and Medicine. vol. 163. pp. 452-454. Poston, H.A., and Wolfe, M.J. (1985). “Niacin requirement for optimum growth, feed conversion and protection of rainbow trout, Salmo gairdneri from ultraviolet-B irradiation” Journal of Fish Diseases. vol 8. no. 5. pp. 451-460. Serrano, A.E. and Nagayama, F. (1991). “Liver 3-hyroxyanthranilic acid oxygenase activity in rainbow-trout (Oncorhynchus-mykiss).” Comparative Biochemistry and Physiology B-Biochemistry & Molecular Biology. vol. 99. no. 2. pp. 275-280. Shaik Mohammed, and Ibrahim, A. (2001) Quantifying the niacin requirement of the Indian catfish (Heteropneustes fossilis) (Bloch), fingerlings, Aquaculture Research, 32: 157-162. Shiau, S.Y., and Suen, G.S. (1992) “Estimation of the niacin requirements for tilapia fed diets containing glucose or dextrin.” Journal of Nutrition. vol .122. no. 10. pp. 2030-6. Tacon, A.G.J. (1985) Nutritional fish pathology: morphological signs of nutrient deficiency and toxicity in farmed fish.” Aquaculture Development and Coordination Programme. ADCP/REP/85/22.

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