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Nutrient requirements and feeding of Haddock

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Early Rearing of Haddock : State of the Art : Proceedings of the Workshop, 16-17 October 2002, St. Andrews, New Brunswick, Canada, Aquaculture Division, Fisheries & Oceans Canada, pp. 79-86, 2003

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Nutrient requirements and feeding of Haddock

Lall, Santosh P.; Nanton, Dominic A.; Tibbetts, Sean M.; Roy, Prabir K.;

Milley, Joyce E.

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Nutrient Requirements

and Feeding of Haddock

Santosh P Lall,

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Dominic A Nanton, Sean M Tibbetts,

Prabir K Roy and Joyce E Milley

National Research Council, Institute for Marine Biosciences. Halifax Nova Scotia, B3H 3Z1 Canada

Abstract

The development of feeds for potential new marine fish species for aquac-ulture must be based on sound information regarding the nutrient require-ments, digestion, absorption and retention of major nutrients and energy utilization from various feed ingredients. Until recently, the field of had-dock nutrition remained unexplored. Our preliminary research has shown that diet containing high amounts of protein (50-55 %), low carbohydrate (<14%), low lipid (<15 %) with a sufficient amount of n-3 long chain polyunsaturated fatty acids (1.5-2.0 % eicosapentaenoic and docosahexaenoic fatty acids) and well fortified with vitamins and trace elements is suitable for initial feed formulations of haddock growout diets. Higher amounts of dietary lipid (>12 %) cause fatty liver and an increase in hepatosomatic index. The role of certain critical nutrients (protein and amino acids, essential fatty acids, minerals, vitamins), energy utilization, feeds and feeding for haddock is briefly reviewed.

H

addock (Melanogrammus aeglefinus), an important commercial groundfish of the Gadidae family, has been one of the most popular food fish for many decades in Canada, United Kingdom and USA. It also has good potential for marine aquaculture in Atlantic Canada (Frantsi 2002). A paucity of information exists on quantitative nutrient requirements of this carnivorous fish. Our laboratory has directed a significant amount of research into developing feed formulations for haddock based on the initial studies of the quantitative protein requirement, the published information on body composition of gadoids and chemical composition of their natural diet, and the estimate of nutrient requirement values derived from salmonids and other marine fish. Recent data on the natural diet of haddock ranging in size from 200 g to 6 kg caught from the Scotian Shelf of Atlantic Canada show their stomach content to consist of echinoderms (38%), crustaceans (10%), molluscs (9%), fish (8%) and polychaetes (6%). Other species of marine

A shotgun

approach has been

used to formulate

feeds for haddock

and cod . . .

(1) Direct correspondence to: Santosh P. Lall, National Research Council, Institute for Marine Biosciences, 1411 Oxford St., Halifax Nova Scotia, B3H 3Z1Canada. Tele-phone: (1) 902 426 6272; Fax: (1) 902 4269413. E-Mail: santosh.lall@nrc-cnrc.gc.ca

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organisms, fish eggs and organic material constituted the remaining portion of the stomach content.(2) A shift in the prey composition with the size of fish has

also been observed. In the stomach of small haddock (<25 cm), the major food organism was krill (73-94% of total weight). It appears that the natural diet of haddock is rich in protein. Krill, other crustaceans and molluscs consumed at early stages of development provide a low level of lipid in their diet. This paper briefly reviews some of our recent findings on nutrient requirements and lipid utilization by haddock.

Nutrient

requirements

Feeds are formulated to provide essential nutrients (protein and amino acids, fatty acids, minerals and vitamins) and energy for growth, reproduc-tion, and health. The development of feeds for farmed fish requires sound knowledge of the quantitative nutrient requirements for economic feed formulation. Limited information is available on the nutrient requirements of most marine fish including haddock (NRC 1993). The protein requirement of most marine fish species ranges between 50-60% and the requirement for juvenile haddock (Kim 2001) has been estimated to be 50%. Most cold water fish species including cod and haddock poorly utilize carbohydrate (Hemre 1989). Dietary lipids supply essential fatty acid (EFA) and energy in the diet of fish and the EFA requirement of most marine fish can only be met by supplying the long-chain highly unsaturated fatty acids (HUFA),

eicosapentaenoic acid (EPA), 20:5n-3, and/or docosahexaenoic acid (DHA), 3. Like other marine fish, haddock tissue contains 20:5n-3 and 22:6n-3 HUFA and a preponderance of these fatty acids in cell membrane phospho-lipids has been reflected in high dietary requirement for these fatty acids (Sargent 2002). The quantitative EFA requirement and deficiency signs in either cod or haddock have not been reported. Based on total body and tissue fatty acid composition data, the estimated EFA requirement of haddock is between 1.5 to 2.0 % of the diet for 20:5n-3 and 22:6n-3 HUFA combined; a value higher than EFA requirement reported for salmonids but close to other marine fish species (NRC 1993). Another EFA for marine fish could be 20:4n-6 (arachidonic acid), a major precursor for production of the eicosanoid compounds that are involved in a variety of stress-related functions (Sargent 1995). Our findings indicate that haddock diets should contain approximately 12% lipid to minimize the prevalence of fatty liver condition in juvenile haddock (50-250 g; see Figure 1). If the dietary lipid source is marine fish oil, the HUFA content will be sufficient to meet the EFA requirement of had-dock.

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The distribution of most water and fat-soluble vitamins and minerals in tissues of haddock has been reported (Lie 1994), however their quantitative requirements have not been established. High amounts of vitamin A and D are known to occur in gadoid livers (Lall 1993). Pathological signs of vitamin A toxicity and vitamin C deficiency have been characterized in marine fish larvae, however no abnormalities due to excess level of vitamin A have been reported for gadoids. The recommended levels of vitamins for salmonids have been successfully used in the formulation of experimental feeds for haddock in our laboratory. At least part of the requirement for certain minerals such as calcium, cobalt, iron, magnesium, potassium, sodium and zinc can be obtained directly from the seawater by haddock and other marine fish. Juvenile haddock require 9.6 mg P/g of diet for optimum growth, feed utilization and bone mineralization and their requirement is much higher than salmonids (Roy 2003a). Deficiency signs of P in haddock have been recently characterized (Roy 2003b)(see Figure 2).

Figure 1. Fatty liver condition in haddock. HSI 18.4%

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Growth and development of male and female gonads require essential nutrients from diets for the synthesis of various biological compounds in the reproductive process. Broodstock diets must therefore be tailored to meet the specific demands for reproduction. Nutrient reserves of the egg and early developmental stages of larvae originate from the female. Few studies have been conducted to define the role of key nutrients in reproduction of gadoids. Broodstock nutrition, as well as the timing of nutrient release, influences the nutrient reserve of eggs and larvae (Bromage 1995). Diets containing inappropriate balances and/or deficiencies of nutrients have been shown to negatively impact spawning frequency as well as egg and larval quality. Although research in this area is limited (nonexistent for haddock), there are clear examples in the literature and numerous examples under practical conditions of larval rearing that problems originate with broodstock nutrition. In species such as red sea bream (Watanabe 1995), problems with egg and larval quality have been traced back to low levels of polyunsaturated fatty acids (PUFA), phospholipids, astaxanthin and other carotenoids in the diet. Lipids, particularly DHA and EPA, are critical to broodstock nutrition and essential for the development of neural tissues. They are also important in stress response and adaptation.

Liver

lipid metabolism

The hepatosomatic index (HSI) or liver to body weight ratio of haddock may exceed 20 % when the diets contain high amounts of lipid (>12%) and this may affect the overall health of fish (Nanton 2001). Farmed haddock with fatty livers convert dietary energy into flesh less efficiently and this presents a major constraint for the commercial culture of haddock from juveniles to market size. The liver lipid content of juvenile haddock with higher HSI (12%) was up to 69% (wet weight), whereas the muscle lipid content remained low (~1%) (Nanton 2001). No pathological changes have been detected in liver of fish with excessive amounts of lipid. Although growth of these fish was not reduced, an increase in liver weight at the expense of somatic tissue growth was observed. Biochemical studies conducted on liver lipid transport mechanisms showed a low concentration of very low density lipoprotein (VLDL; <50 mg/dL) in the plasma of haddock after lipid absorption which suggests a low level of lipid transport out of the liver. Plasma VLDL is a major transporter of deposited lipid out of the liver (Sheridan 1988), and VLDL triacylglycerol (storage lipid) levels in the plasma were positively correlated with muscle lipid storage in marine teleost fish species (Ando 1993).

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The red muscle showed higher fatty acid catabolic activity (ß-oxidation of palmitic acid), on a per g wet tissue basis, compared with white muscle or liver. However, the white muscle appears to be the most important tissue in juvenile haddock for the overall catabolism of fatty acids because it com-prises a higher proportion of the total fish weight (Nanton 2003). These observations suggest that the transport of lipid as lipoprotein from the predominant storage organ, liver, to the catabolic site muscle, may be low in haddock (Nanton 2003). Current findings from our laboratory suggest that fatty liver condition or HSI in haddock is not significantly reduced by supple-mentation of lipotropic factors such as choline, methionine and inositol or an increase in chitin content or fibre content. The increase in HSI was mainly due to the increase in total lipid content of the diet.

Figure 2. Phosphorus deficiency in haddock. A— an X-radiograph of haddock showing deformation and diffusion of vertebrae (bar = 2 cm). B— an X-radiograph of haddock showing deformed vertebrae (bar = 1 cm).

C— fusion of vertebrae. Macerated spine. (bar = 2 cm).

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Practical

feeds

A “shotgun” approach has been applied to formulate grow out feeds for haddock and cod. Current knowledge of the nutrient requirement of gadoids and other marine fish suggests that a diet containing a high amount of protein (50-55%) and low levels of carbohydrates (<14%) and lipid (<15%) with sufficient amount of n-3 long chain HUFA (1.5 % EPA and DHA for juvenile fish) as well as being fortified with vitamins and trace elements would be suitable for initial feed formulations of haddock diets. Apparent energy digestibility coefficients measured with haddock are high or moderately high for herring meal, crab meal, soybean meal and corn gluten meal with values of 92.2, 82.5, 92.1 and 80.7%, respectively (unpublished data). The

digestibility value for herring meal (92%) is similar to those reported for other commercially important salmonid and marine species.

A proper balance of digestible protein (DP) and digestible energy (DE) is necessary to minimize excessive accumulation of lipid and glycogen in the liver and to keep feed-costs reasonable.This balance is referred to as the optimum DP/DE ratio, and for cold-water marine species like cod (Lie 1988) it is 33 g DP MJ DE-1and for halibut (Aksnes 1996) it is 27 g DP MJ DE-1. We have obtained the highest growth rate, best feed conversion efficiency and highest digestibility in growing haddock on a diet with a DP/DE ratio of 30 g DP MJ DE-1in a diet containing 55% crude protein, 11% lipid and 17 MJ/kg DE (unpublished data).

Moist and dry commercial and experimental feeds have been used for feeding gadoids, particularly cod. The processing and production of moist feeds show wide variations. Moist pellets contain a variable amount of fishery by-products (ground whole fish, fish and crustacean waste) or fish silage. Dry ingredients (fish, crab and shrimp meals, wheat by-products, corn gluten meal, soybean meal etc.), vitamins, minerals and binding agent etc. are first mixed and then incorporated with wet fishery products and extruded through a meat grinder or more elaborate cold extruders. Raw fish and fishery-products must be pasteurized to destroy pathogens and thiaminase enzyme found in fish tissues. The overall acceptability of moist diets by haddock remains high as compared with dry extruded feeds for fish reared in Eastern Canada where average annual seawater temperature is between 6-8°C (range 0-18°C).

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Feeding

of haddock

Feed attractiveness or palatability of the feed for haddock is important, particularly at lower water temperatures when food acceptability decreases. Due to the high level of energy (lipid) storage in the liver of gadoid fish fed formulated diets, less frequent feeding has been shown to improve feed utilization and reduce fatty liver to some extent in cod (Maeland 2001). However, growth of juvenile haddock held at 12°C under laboratory conditions was significantly reduced when they were fed either alternate days or once a day (unpublished data). Growth rate has been positively correlated with hepatosomatic index in gadoids (Dos Santos 1993), thus attempting to reduce the abnormally high hepatosomatic index or liver lipid storage levels by reducing feeding frequency can negatively affect growth in gadoids fed formulated diets. A better balance of protein and energy ratio may be a better strategy to reduce the fatty liver condition rather than to reduce feeding rate until investigations are undertaken to properly establish the feeding rate of haddock under farm conditions. Water temperature is an important factor that determines the ability of fish to grow and utilize feed. The optimum temperature for growth of juvenile and adult haddock has not been estab-lished. Studies conducted on cod show that large sexually mature cod have a lower optimal temperature for growth than smaller cod (50-1000 g) (Dos Santos 1993).

In summary, some progress has been made in recent years to develop grower diets for haddock and cod, however, there is a need to improve the larval, weaning and broodstock diets of haddock and also to better define the role of critical micronutrients such as vitamins E, D, K and C.

Literature cited

Aksnes A, Hjertnes T, Opstvedt J. 1996. Aquaculture 145:225-233.

Ando S, Mori Y. 1993. Nippon Suisan Gakkaishi 59: 565-1571.

Bromage NR, Roberts TJ. 1995. Pages XXXXX in: Broodstock management and egg

and larval quality. (NR Bromage and TJ Roberts, eds). Blackwell Science Ltd., Cambridge, Massachusetts, USA.

Dos Santos J, Burkow IC, Jobling M. 1993. Aquaculture 110:173-189.

Frantsi C, Lanteigne C, Blanchard B, Alderson R, Lall S, Johnson S, Leadbeater S,

Martin-Robichaud D, Rose P, 2002. Bull. Aquacult. Assoc. Canada 102-1:31-34.

Hemre G-I, Lie Ø, Lied E, Lambersten G, 1989. Aquaculture 80:261-270.

Kim JD, Lall SP, Milley JE. 2001. Aquacult. Res. 32:1-7.

Lall SP, Parazo MPM. 1993. Pages 157-186 in: Fish and fishery product:

Composi-tion, nutritive properties and stability, (A Ruiter, ed), Cab International, Oxon, UK.

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Lie Ø, Lied E, Maage A, Njaa LR, Sandnes K. 1994. Fiskeridir. Skr. Ser Ernæring

6:83-105.

Maeland KT. 2001. Int’l Aquafeed 1:39-42.

Nanton DA, Lall SP, McNiven MA. 2001. Aquacul. Res. 32:225-234.

Nanton DA, Lall SP, Ross NW, McNiven MA. 2003. Comp. Biochem. Physiol. B. (in

press).

NRC (National Research Council), 1993. Nutrient requirement of fish. National Academy Press, Washington DC, 114 p.

Roy PK, Lall SP. 2003a. Aquaculture (in press).

Roy PK, Witten PE, Hall BK, Lall SP. 2003b. J. Fish. Physiol. Biochem. (in press).

Sargent JR, Bell JG, Bell MV, Henderson RJ, Tocher DR. 1995. J. Appl. Ichthyol. 11:

183-198.

Sargent JR, Tocher DR, Bell JG. 2002. Pages 181-257 in: Fish Nutrition. 3rd Ed., (JE

Halver and RW Hardy, eds), Academic Press Inc., San Diego, CA,.

Sheridan MA. 1988. Comp. Biochem. Physiol. 90B:679-690.

Watanabe T, Kiron V. 1995. Pages 394-413 in: Broodstock management and egg

and larval quality. (NR Bromage and TJ Roberts, eds). Blackwell Science Ltd., Cambridge, Massachusetts, USA,.

Figure

Figure 1.  Fatty liver condition in haddock.
Figure 2. Phosphorus deficiency in haddock. A— an X-radiograph of haddock showing deformation and diffusion of vertebrae (bar = 2 cm)

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