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F: Formulation
Minerals
hrimp is one of the most important internationallytraded commodities, in terms of value and according to the latest figures available, from the FAO (2009), world shrimp production from aquaculture represents 3.7 million tonnes per year.
and utilise nutrients from these sources, for metabolic and physiological processes. Fishmeal has traditionally been one of the primary protein sources in shrimp diets and in addition to providing protein and essential amino acid it also serves as an attractant and provides a rich source of essential fatty acids, fat soluble nutrients and minerals. The requirement for sustainable diets demands a reduction in fishmeal usage It is the most valuable fishery export in and its replacement by vegetable sources. many tropical developing countries providA number of studies have investigated this ing significant employment in these regions. (Forster et al. 2003, Tan et al. 2005). Feed represents more than 50 percent However, a major concern is the incomof the production costs of intensive shrimp plete nutrient composition of these alternaproduction (Tan et al. 2005) with protein tive ingredients, which may not only reduce sources accounting for at least 30 percent growth, but also increases the complexity of of the compounds found in commercial feed formulation as more ingredients must diets. be utilised to satisfy nutrient requirement. Volatility in the availability and pricing This shift in formulation raises numerous of fishmeal, fish oil and vegetable protein concerns, such as an increase in indigestible sources, such as soymeal and grains etc, has components (fibre and non-starch polysacencouraged flexibility in the formulation of charides), an increase in phytic acid (with its diets to minimise cost. negative interaction on mineral availability), This flexibility requires an in-depth available phosphorous, phospholipids and understanding of the nutritional requiregeneral digestibility of the protein content. ment of the species and its ability to digest As a result, current research is now focused on the effect of phytochemical components, present in these diets, on the metabolic and physiological response in shrimp. In addition, the identification of genes responsible for such changes, at the molecular level, is also being investigated and is an area of growing imporFigure 1: A comparison of the essential amino acid requirements for four shrimp species tance. Information
from these studies are providing further knowledge on the nutritional requirements of shrimp and enabling rapid advances in shrimp nutrition (Gillies & Faha 2003, Match et al.2005).
Proteins
Shrimp, like other fish and animals, have a requirement for a well-balanced mixture of essential and non-essential amino acids from which to construct their own protein tissues and therefore the balance of essential amino acids in the diet is crucial. The amino acid requirements of four commercially important shrimp species are shown in Figure 1 The figure indicates that the amino acid requirements of carnivorous species like P. japonicus are higher than the amino acid requirements of herbivorous species like P. vannamei. Some authors thought that their protein utilization was less effective than in the other species and therefore additional dietary protein supplementation was required. However it has been proven that the source and nature of the protein its self is important in determining the protein levels required by each species. The assimilation of amino acids by shrimps is most effective when they are bonded; as opposed to pure free essential amino acids and in this form they have performed better under the same experimental conditions.
Fatty Acids
Crustaceans have long been recognized as having a limited ability to synthesize highly unsaturated fatty acids (HUFA) de novo and having no ability to synthesize sterols de novo. Shrimp, therefore, do not have a definitive dietary lipid requirement but rather require sufficient lipid to meet
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F: Formulation
pathways such as zinc activation of alkaline phosphatase. Mineral supplementation is considered even more important for the freshwater prawns, like M. Rosenbergii since their environment contains less available minerals (Muir & Roberts 1982). The macro minerals calcium (Ca), phosphorous (P), sodium (Na) and potassium (K) have a special role to play in osmoregulation, mineral and water equilibrium, and the maintenance of the acid-base balance. Davis et al. (1993) demonstrated that Ca and P are essential components of hard tissues such as the exoskeleton of the shrimp and that calcium is necessary for impulse transmission, osmoregulation and muscle functions. A dietary requirement for Na has not been demonstrated for marine shrimps, but in the case of pond grown shrimps, supplementation with salt (NaCl) has resulted in increased growth. Both freshwater and seawater probably contain sufficient concentrations of Na and P to satisfy the requirements of shrimp, however Kanazawa et al. (1984) reported that diets with 0.9 percent of P improved growth in P. japonicus. Trace minerals such as manganese (Mn), copper (Cu), iron (Fe), zinc (Zn) and selenium (Se) have important roles to play in shrimp biochemistry and physiology. The determination of mineral dietary requirements is very difficult for aquatic animals because of their ability to absorb minerals directly from their environment. Copper is important in the oxygen transfer agent in crustaceans, hemocyanin, and it is has been estimated that around 40 percent of the total Cu weight found in shrimps is contained within hemocyanin. Copper is also required to achieve maximum growth and tissue mineralization, deficiency of Cu in P. vannamei has been characterized by poor growth and reduced concentrations of Cu in the carapace. Excess Fe can produce toxic effects in shrimps, which can lead to decreasing growth in P. japonicas and zinc has been shown to have an important role in normal tissue mineralization in P. vannamei (Davis et al. 1993). A comparison of the macro and trace mineral requirements of 4 shrimp species is given in Figures 3 and 4. use and increase the quality of the final product.
References
FAO (2009). The state of world fisheries and aquaculture 2008. Rome, FAO Fisheries and Aquaculture Department, Rome. 176pp DAbramo LR (1998) Nutritional requirements of the freshwater Prawn Macrobrachium rosenbergii: Comparisons with species of penaied shrimp. Reviews in Fisheries Science 6: 153-163. DAbramo LR, Conklin DE (1992) New developments in the understanding of the nutrition of penaeid and caridean species of shrimp. In: Browdy CL, Hopkins SJ Eds, Swimming Through Troubled Water. Proceedings of the Special Session on Shrimp Farming, Aquaculture 95, World Aquaculture Society, Baton Rouge, LA, USA, pp. 95107. Davis DA, Lawrence AL et al. (1993) Evaluation of the dietary zinc requirement of Penaeus vannamei and effects of phytic acid on zinc and phosphorous bioavailability. J of the World Aquaculture Society 24: 40-47. Forster I, Dominy W et al. (2003). Rendered meat and bone meals as ingredients of diets for shrimp Litopenaeus vannamei (Boone, 1931). Aquaculture 219: 655670. Gillies JP, Faha PD (2003) Nutrigenomics: the Rubicon of molecular nutrition. J American Dietetic Association 103: s50s55. He H, Lawrence AL et al. (1992) Evaluation of dietary essentiality of fat soluble vitamins A, D, E and K for penaeid shrimp (Penaeus vannamei) Aquaculture 103: 177-185. Kanazawa A, Teshima S, Sasaki M (1984) Requirements of the juvenile prawn for calcium, phosphorus, magnesium, potassium, copper, manganese, and iron. Mem Fac Fish Kagoshima Univ 33:6371. Match MD, Wahli W, Williamson G (2005) Nutrigenomics and nutrigenetics: the emerging faces of nutrition. The Federation of American Societies for Experimental Biology publishes The FASEB Journal 19: 16021614. Muir JF, Roberts RJ (1982) Recent advances in aquaculture. London, Croom Helm. Tan B, Mai K et al. (2005) Replacement of fish meal by meat and bone meal in practical diets for the white shrimp Litopenaeus vannamei (Boone). Aquac Res 36: 439444. Teshima S I (1997). Phospholipids and sterols. In DAbramo LR, Conklin DE and Akiyama DM (Eds) Crustacean nutrition, advances in world aquaculture. World Aquaculture Society, pp 85-107.
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Conclusion
Current feeding methods are taking advantage of shrimp omnivorous habits by incorporating plant-based ingredients in their diet. Alternative sources of protein require supplementation of some essential amino acids, lipids and minerals in order to meet the nutritional requirements of the shrimp species being cultured. By using the new research area of nutrigenomics it is possible to develop a more detailed understanding of how different components of the diet , e.g. cholesterol, can influence molecFigure 3: A comparison of the macro mineral ular mechanisms requirements for four shrimp species which in turn will help in understanding their role in shrimp physiology and metabolism. These future studies will enable strategies to be developed that will help us to better understand Figure 4: A comparison of the trace element requirements shrimp nutrition, for four shrimp species optimise nutrient
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