Nutrient requirements of penaeid shrimps1

Aquaculture 164 Ž1998. 77–93

Nutrient requirements of penaeid shrimps Shi-Yen Shiau

1

)

Department of Marine Food Science, National Taiwan Ocean UniÕersity, Keelung, 202, Taiwan

Abstract Of the penaeid shrimps, Penaeus japonicus has been studied early and more thoroughly than the other species in terms of nutrient requirements. In recent years, more research has been carried out for Penaeus monodon and Penaeus Õannamei. Other penaeid shrimps lag behind in this aspect. However, even for P. japonicus, there are still many essential dietary nutrients that need to be quantified. Besides, evidence has shown that differences exist in nutrient requirements among penaeid species. This paper reviews nutrient requirements of penaeid shrimps based on available information in the literature. It is intended to serve as a guide for those who wish to have general knowledge on penaeid shrimp nutrition. q 1998 Elsevier Science B.V. All rights reserved. Keywords: Feeding and nutrition—crustacean; Nutrient requirements; Penaeids

1. Introduction As shrimp farms evolve from low to high stocking densities, the quality of feed become very important. Some extensive farms Žlow stocking densities. are not provided with artificial feed at all; shrimps feed on naturally occurring food organisms in the pond. Other extensive farms use small amounts of feed and fertilizer during a certain season or stage to stimulate the natural food chain. In semi-intensive farms where many shrimps scour the bottom of the ponds, most of the feed is consumed by the shrimp and less feed is available to serve as a stimulant to the natural food web. Hence, feed quality is more important because the shrimps get most of their nutrition from the feed. In intensive farms, shrimps depend mostly on artificial diets as source of their nutrients. Therefore, intensive farms require the best quality feed. )

Corresponding author. Tel.: q886-2-2462-2192 Ext. 5106; fax: q886-2-2462-1684; e-mail: [email protected] 1 This paper was presented at the Second International Conference on the Culture of Penaeid Prawns and Shrimps, 14–17 May 1996, Iloilo City, Philippines. 0044-8486r98r$19.00 q 1998 Elsevier Science B.V. All rights reserved. PII S 0 0 4 4 - 8 4 8 6 Ž 9 8 . 0 0 1 7 8 - 1

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Penaeid shrimps are among the most important and extensively cultured crustaceans in the world. Although diseases pose a serious threat to penaeid shrimp aquaculture, the production of these highly valued crustaceans continue to grow. In 1995, the world’s shrimp farmers produced an estimated 712,000 metric tons of whole shrimp ŽWorld Shrimp Farming, 1995. and feed mills around the world produced approximately one million metric tons of shrimp feed. Feeds can represent over 50% of the production cost in modern shrimp farms. The nutritional requirements of Penaeus japonicus were defined by the introduction of refined test diets in the early 1970s ŽKanazawa et al., 1970; Deshimaru and Kuroki, 1974a.. To a certain extent nutritional requirements of other species including P. monodon, P. Õannamei, P. aztecus, P. californiensis, P. indicus, P. merguiensis, P. setiferus, P. stylirostris, P. penicillatus, P. chinensis and P. duorarum have been studied. This paper presents an overview of the nutrient requirements of penaeid shrimps. 2. Protein and amino acids Proteins are indispensable nutrients for the structure and function of all living organisms including shrimps. Since proteins are continually being used by the animal for growth and repair of tissues, a continuous supply of proteins or its component amino acids is needed. Protein is a major and expensive component of feeds. Therefore, nutrition studies of shrimp often start with investigating the optimal dietary protein level. As a consequence, the most researched nutrient in terms of the number of penaeid shrimp species being studied is proteins. Table 1 lists a summary of the optimum dietary

Table 1 Summary of protein requirements for various species of penaeid shrimp Penaeus spp.

Requirement Ž%.

References

P. aztecus

40 51 35 43 50 52–57 45–55 34–42 45–50 40 40–50 40–44 36–40 28–32 30 35 30 ) 36

Venkataramiah et al., 1975 Zein-Eldin and Corliss, 1976 Colvin and Brand, 1977 Colvin, 1976 Deshimaru and Kuroki, 1975a Deshimaru and Yone, 1978a Teshima and Kanazawa, 1984 Sedgwick, 1979 Lee, 1971 Alava and Lim, 1983 Bautista, 1986 Shiau et al., 1991a Shiau and Chou, 1991 Andrews et al., 1972 Lee and Lawrence, 1985 Colvin and Brand, 1977 Colvin and Brand, 1977 Smith et al., 1985

P. californiensis P. indicus P. japonicus

P. merguiensis P. monodon

P. setiferus P. stylirostris P. Õannamei

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protein level for different penaeid shrimps. Generally, recommended dietary protein levels vary from 30% to 57% for various species. The reported estimate of protein requirement must be carefully examined because it is dependent on quality Žessential amino acid profile. of dietary protein, age or physiological state of crustaceans ŽD’Abramo and Sheen, 1994.. Recently, Shiau et al. Ž1991a. demonstrated that protein requirements of shrimp are influenced by the environment. The optimal dietary protein level was found to be lower in juvenile P. monodon when reared in seawater Ž40% protein. than those reared in 16 ppt brackish water Ž44% protein.. This may be due to the differential utilization of dietary protein as energy source when the shrimps are raised at varied salinity levels. P. monodon acclimated at low salinity showed higher ammonium-N excretion than those acclimated at high salinity ŽLei et al., 1989., thus indicating possible difference in protein utilization. Lei et al. Ž1989. suggested that shrimp raised in low salinity tend to use protein, not lipid, as energy source. The effect of salinity on protein digestibility may play a role in protein utilization. Shiau et al. Ž1992. determined the digestibility of fish meal, soybean meal, and casein by P. monodon in both brackish water Ž16 ppt. and seawater Ž32 ppt.. Casein was the most digestible protein, followed by soybean meal and fish meal. Salinity did not affect the digestibility of casein and fish meal but did affect digestibility of soybean meal. Protein digestibility was significantly lower when P. monodon was raised at 32 ppt than 16 ppt. Kanazawa and Teshima Ž1981. showed by tracer techniques using radioactive acetate that P. japonicus requires 10 amino acids, i.e., arginine, methionine, valine, threonine, isoleucine, leucine, lysine, histidine, phenylalanine, and tryptophan. These essential amino acid requirements have also been demonstrated for other penaeids such as P. monodon ŽColoso and Cruz, 1980. and P. aztecus ŽShewbart et al., 1972.. Juvenile and adult shrimps are incapable of efficiently utilizing free amino acids or hydrolyzed protein products in the diet. Deshimaru and Kuroki Ž1974c, 1975a,b. demonstrated that diets containing only amino acids instead of protein brought about very poor growth and high mortality in feeding trials of P. japonicus. In contrast, larval P. japonicus was observed by Teshima et al. Ž1986c. to be able to utilize crystalline amino acid in microparticulate diets as a partial protein substitute. The difference in the utilization of free amino acids or protein could be attributed to the difference in the developmental stage of the shrimp or the type of diets. The incapability of juvenile or adult shrimps to utilize dietary crystalline amino acids had made it a barrier in quantifying their essential amino acid requirements. Recently however, various methods have been used to overcome the problem. A study using microencapsulated L-arginine indicated that a level of 2.5 gr100 g diet Ž5.5 gr100 g protein. is required to achieve optimal growth for juvenile P. monodon ŽChen et al., 1992a,b.. By adjusting dietary pH to neutrality and increasing meal frequency to 5 times per day, Liou and Yang Ž1994. successfully incorporated crystalline methionine and other amino acids in the diet of juvenile P. monodon and estimated the methionine Žqcystine. requirement to be 1.4 gr100 g diet Ž4.0 gr100 g protein.. Millamena et al. Ž1996b. were able to show that threonine requirement for postlarval P. monodon growth is 1.4 gr100 g diet Ž3.5 gr100 g protein. using diets containing pure amino acid mixture. The amino acids were neutralized and coated with k-carboxymethylcellulose and feed pellets were bound with k-carrageenan. Using the same technique, Millamena

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et al. Ž1996a. also found the requirements of P. monodon for valine to be 3.75 gr100 g protein.

3. Carbohydrates The information on the carbohydrate nutrition of crustaceans is limited. Table 2 summarizes the studies on carbohydrate utilization by penaeid shrimps. 3.1. Utilization of sugars In general, simple sugars are poorly utilized by shrimp. Andrews et al. Ž1972. conducted two experiments to study the influence of protein, carbohydrate and lipid on P. setiferus. Glucose and starch were used as the carbohydrate sources. Results indicated that the addition of glucose to the diets resulted in depressed growth whereas supplemental starch did not cause a reduction in weight gain. Sick and Andrews Ž1973. reported that growth and survival of P. duorarum fed diets containing 40% starch were higher than those fed diets containing 40% glucose. Deshimaru and Yone Ž1978b. compared weight gain and feed efficiency of P. japonicus fed diets containing 10% of either glycogen, starch, dextrin, glucose or sucrose. P. japonicus fed the sucrose diet had the highest weight gain, whereas those fed the glucose diet had the lowest weight gain. Feed efficiency was highest for those fed the starch diet, followed by glycogen, sucrose and dextrin in decreasing order. Abdel-Rahman et al. Ž1979. compared the effect of a 19.5% dietary level of either glucose, galactose, sucrose, dextrin, soluble starch, potato starch and glycogen on the growth and the level of the hepatopancreatic glycogen and serum glucose of P. japonicus. Growth was poor in shrimp fed the monosaccharides, glucose and galactose. High hepatopancreatic glycogen concentrations were characteristic of shrimp fed glucose or galactose containing diets. Pascual et al. Ž1983. fed diets containing maltose, sucrose, dextrin, molasses, cassava starch, corn starch or sago palm starch to P. monodon at levels of either 10 or 40% and found no correlation between survival and the relative complexity of carbohydrates. While sucrose and maltose are both disaccharides, survival was significantly higher with the sucrose diet Ž10 vs. 40%.. The authors explained the response by noting that the end products of the digestion of sucrose are glucose and fructose while maltose is broken down into two units of glucose. Maltose is a reducing sugar whereas sucrose is not. A conclusion cannot be surmised because a treatment group fed a glucose containing diet was not included for comparison. Shiau and Peng Ž1992. demonstrated that corn starch was better utilized than glucose by P. monodon. Alava and Pascual Ž1987. fed diets containing either 10, 20, or 30% either trehalose, sucrose or glucose to P. monodon. Those shrimp fed diets containing trehalose and sucrose had higher weight gains and lower mortality than those fed the glucose diets. Trehalose is a sugar found in insect hemolymph and has similar properties to sucrose in that both are non-reducing sugars. However, like maltose, trehalose splits in two glucose units. The greater utilization of non-reducing sugars by shrimp is interesting and this

Table 2 Carbohydrate utilization by various species of penaeid shrimp Carbohydrate source

Resultsa

References

P. setiferus P. duorarum P. japonicus

glucoserstarch glucose, starch glycogen, starch, dextrin, glucose, sucrose glucose, starch, dextrin, potato starch, glycogen, galactose, fructose, sucrose maltose maltose, sucrose, dextrin, molasses, cassava starch, corn starch, sago palm starch trehalose, sucrose, glucose wheat flour, straight, first grade clear, second grade clear gelatinized bread flour glucose, dextrin, starch

) utilizationsstarch ) utilizationsstarch ) weight gainsssucrose, ) feed efficiency sstarch, - utilizations glucose - utilizations monosaccharides Žglucose, galactose.

Andrews et al., 1972 Sick and Andrews, 1973 Deshimaru and Yone, 1978b

)survivalssucrose

Pascual et al., 1983

) utilizations trehalose and sucrose utilized no difference

Alava and Pascual, 1987 Shiau et al., 1991b

- weight gain and feed conversion ratios 35% ) utilizationsstarch or dextrin

Catacutan, 1991a Shiau and Peng, 1992

P. japonicus

P. monodon

P. monodon P. monodon P. monodon P. monodon a

Abdel-Rahman et al., 1979

S.-Y. Shiaur Aquaculture 164 (1998) 77–93

Penaeus spp.

) s highest, - s lowest.

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area needs more research. Other related factors such as palatability, leaching, etc. also need to be addressed. Shiau and Peng Ž1992. investigated the utilization of different carbohydrate sources and the possible sparing effect of dietary protein by carbohydrate in P. monodon reared in seawater. In their study, three dietary protein levels Ž40, 35 and 30%. and three levels Ž20, 25, and 30%. of three carbohydrate sources Žglucose, dextrin, starch. were tested. Results indicated that shrimp fed starch or dextrin had significantly higher weight gain, feed efficiency ratio, protein efficiency ratio and survival than those fed glucose. It also appears that starch has a better protein-sparing effect than dextrin or glucose. Accordingly, the required dietary protein level for P. monodon is lower if starch, instead of glucose or dextrin, is used as carbohydrate source. 3.2. Poor utilization of glucose The mechanism responsible for the poor utilization of glucose by some species of penaeid shrimps studied is not yet fully understood. One possibility may simply be a higher rate of absorption across the digestive tract. In rainbow trout, Piefer and Pfeffer Ž1980. suggested that the poor performance relative to glucose containing diets might be the result of ‘negative physiological effects’ caused by glucose saturation. Glucose requires no digestion and is rapidly absorbed across the gut. However, more complex carbohydrates such as starch must undergo enzyme hydrolysis before assimilation. Thus, glucose arising from the enzymatic hydrolysis of starch appears at gut absorption sites slower than free glucose. Furuichi and Yone Ž1982b. and Murai et al. Ž1983. suggested that the rapid absorption of glucose across the intestine combined with insufficient levels of insulin secretion to metabolize the rapid increase are responsible for the poor utilization of glucose by carp. Rapid absorption of free glucose results in a considerable amount of glucose entering the body tissue before sufficient elevation of the activities of carbohydrates metabolizing enzyme can occur ŽFuruichi and Yone, 1982a.. A similar situation may exist in penaeid shrimps. Abdel-Rahman et al. Ž1979. reported that the level of plasma glucose in P. japonicus increased rapidly after they were fed a diet containing glucose and remained at high levels for 24 h. In contrast, plasma glucose was found to increase to a maximal level after 3 h and then decrease to a low level when the diet contained disaccharides and polysaccharides. These authors suggested that dietary glucose was quickly absorbed from the alimentary canal and released into the hemolymph, resulting in a physiologically abnormal elevation of plasma glucose levels, thereby impairing its utilization as an energy source. Shiau and Peng Ž1992. also reported that plasma glucose levels in P. monodon fed glucose containing diets peaked earlier than in shrimp fed dextrin or starch containing diets. Another possible explanation for the poor growth performance of shrimp fed glucose containing diets is the possible inhibition of amino acid absorption in the intestine due to the presence of glucose ŽAlvarado and Robinson, 1979.. Hokazeno et al. Ž1979. reported that the presence of 10 mM of glucose reduced the uptake of L-lysine from 26.64 to 12.34% in the mid-intestine and from 23.24 to 5.40% in the posterior intestine, of the rainbow trout. However, this interaction has not been studied in penaeid shrimps.

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3.3. Utilization of complex polysaccharides 3.3.1. Starch Shiau et al. Ž1991b. studied the utilization of three types of wheat flour Že.g., straight flour, first grade clear flour, second grade clear flour. in diets fed to P. monodon. Starch content of the flours tested is about 70%. At a dietary level of 35%, no differences in body weight gain, protein digestibility, dry matter digestibility and carbohydrate digestibility were found, suggesting that all three types of wheat flour have the same nutritional value. Catacutan Ž1991a. used gelatinized bread flour as a carbohydrate source at levels of 5, 15, 25, and 35% in diets fed to P. monodon. Weight gain and specific growth rate were lowest and the feed conversion ratio was poorest in shrimp fed the diet containing 35% of this carbohydrate source. 3.3.2. Chitin Results of studies on the effect of supplemental dietary glucosamine on growth and survival of P. japonicus are inconclusive. Kitabayashi et al. Ž1971. demonstrated that addition of 0.52% glucosamine to diets improved growth but growth was retarded if chitin was added to the diet. Deshimaru and Kuroki Ž1974b. stated that a dietary source of glucosamine is unnecessary for P. japonicus juveniles and that the presence of glucosamine inhibits the growth-promoting effect of cholesterol. Akiyama et al. Ž1992. indicated that chitin is the major structural component of the exoskeleton of shrimp. They recommended a minimum level of 0.5% chitin in shrimp feeds because dietary chitin is believed to have a growth promoting effect.

4. Lipids Penaeid shrimp may not have a definite lipid requirement. Recommended lipid levels for commercial shrimp feeds range from 6% to 7.5% and a maximum level of 10% was suggested ŽAkiyama et al., 1991.. Sheen et al. Ž1994a. found no difference in weight gain of juvenile P. monodon fed isoenergetic and isonitrogenous diets containing between 4 to 11% mixture of cod liver oil and corn oil. The unique aspect of lipid nutrition in penaeid shrimps is the requirement of polyunsaturated fatty acids, phospholipids and sterols. A series of feeding experiments conducted by Kanazawa et al. Ž1977, 1978, 1979b,d. demonstrated that there are four fatty acids that are considered essential for P. japonicus: linoleic Ž18:2 n y 6., linolenic Ž18:3n y 3., eicosapentaenoic Ž20:5n y 3, EPA., and decosahexaenoic Ž22:6n y 3, DHA. acids, and the latter two n y 3-highly unsaturated fatty acids ŽHUFA. are the most indispensable. Qualitative n y 3 HUFA requirements have been reported for P. indicus ŽRead, 1981., P. stylirostris ŽLeger et al., 1985., and P. monodon ŽCatacutan, 1991b.. The optimum levels of EPA and DHA for P. japonicus juveniles were found to be about 1% in the diet ŽKanazawa et al., 1979a.. Shewbart and Mies Ž1973. also revealed that growth of P. aztecus was improved by the addition of 18:3n y 3 to the diet, and that optimum growth was attained with diets containing 1% 18:3n y 3. Growth experiments of early postlarval P. monodon with alginate-encapsulated diets indicated the requirement for HUFA to be

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0.5–1% of diet ŽChen and Tsai, 1986.. Rees et al. Ž1994. concluded that while P. monodon postlarvae can grow well on an Artemia diet containing low amounts of n y 3 HUFA, a high dietary supply of HUFA Ž12 to 22 mgrg dry weight of lipid. would considerably enhance their ability to sustain stress, and would eventually improve their survival. However, excessive dietary n y 3 HUFA ŽG 31.2 mgrg dry weight. may lead to detrimental effects on both the growth and survival of the postlarvae. Cholesterol is essential for penaeids and this is probably the most unique aspect of lipid nutrition in crustaceans. A feeding experiment using artificial diets conducted by Kanazawa et al. Ž1971. first demonstrated that P. japonicus requires sterols for growth and survival. The level of cholesterol required by P. japonicus is still not certain. Kanazawa et al. Ž1971. reported a value of 0.5% dietary cholesterol for good growth whereas other researchers have obtained the best growth of P. japonicus with diets containing 0.2% ŽShudo et al., 1971. and 2.1% ŽDeshimaru and Kuroki, 1974b. cholesterol. These contradictory results with P. japonicus are considered to be due to differences in the composition of the test diets used. The interaction between cholesterol and phosphatidylcholine on weight gain of P. penicillatus and P. monodon was investigated by Chen and Jenn Ž1991. and Chen Ž1993., respectively. Both studies used three levels of dietary cholesterol Ž0, 0.5, and 1%.. These workers concluded that 0.5% dietary cholesterol is required for good growth of P. penicillatus and P. monodon. Sheen et al. Ž1994b. found no difference in weight gain of P. monodon fed diets containing 0.2–0.8% cholesterol and suggested that a 1% addition has an adverse effect on growth. In the study of Chen Ž1993. however, no detrimental effects on growth were detected when the diets contained 1% cholesterol. A requirement for dietary phospholipids ŽPL. and particularly phosphatidylcholine ŽPC. has been demonstrated in various penaeid species, including larval ŽTeshima et al., 1982; Kanazawa et al., 1985. and postlarval P. japonicus ŽKanazawa et al., 1979c; Teshima et al., 1986a,b., juvenile P. penicillatus ŽChen and Jenn, 1991., P. monodon ŽPiedad-Pascual, 1986; Chen, 1993. and P. chinensis ŽKanazawa, 1993.. Inclusion levels of dietary PL reported for various species of penaeids range from 0.84% of PC for P. chinensis ŽKanazawa, 1993. to 1.25% of PC for P. penicillatus and P. monodon ŽChen and Jenn, 1991; Chen, 1993..

5. Vitamins So far, only P. monodon, P. japonicus, P. chinensis, P. Õannamei and P. californiensis have been studied to some extent for their vitamin requirements, of which a great deal of research have been carried out in P. monodon and P. japonicus. Table 3 lists the vitamin requirements of these five penaeid shrimps. The dietary requirements of P. monodon for thiamin, riboflavin and niacin requirements are much lower than those of P. japonicus. This species variation can also be seen in the vitamin K and choline requirements. Shiau and Liu Ž1994a. reported that 30–40 mg vitamin Krkg diet is required for P. monodon, whereas the same authors observed that P. chinensis requires 185 mg vitamin Krkg diet ŽShiau and Liu, 1994b.. A choline requirement of 4000 mgrkg diet was reported for P. chinensis ŽLiu et al., 1993., whereas, 600 mgrkg diet

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Table 3 Vitamin requirements of Penaeus monodon, P. japonicus, P. chinensis, P. Õannamei and P. californiensis Vitamins

Requirement Žmgrkg diet. P. monodon

Thiamin Riboflavin Pyridoxine Vitamin B12 Niacin Biotin Folic acid Inositol Choline

13–14 22.5 b y 0.2 c 7.2 d y 2–8 e y y

a

P. japonicus l

60–120 80 m 120 l y 400 m y y 2000–4000 n 600 n dispensable l Pantothenic acid y y Ascorbic acid 2000 ŽC1. f 3000 ŽC1. o 210 ŽC2PP. g 10,000–20,000 ŽC1. p 100–200 ŽC2PMg. h 215–430 ŽC2PMg. q 40 ŽC2MP. i 157 ŽC2S. i A y y D 0.1 ŽD 3 . j y E y y K 30–40 k y

P. chinensis P. Õannamei

P. californiensis

y y y y y y y 4000 r 4000 r

y y 80–100 t y y y y y y

y y y y y y y y y

y y

y y 90–120 ŽC2PP. u 2000 ŽC1. v

y y y 185 s

y y 99 u y

y y y y

a

Chen et al., 1991. Chen and Hwang, 1992. c Shiau and Lung, 1993. d Shiau and Suen, 1994. e Lung, 1991. f Shiau and Jan, 1992. g Chen and Chang, 1994. h Catacutan and Lavilla-Pitogo, 1994. i Shiau and Hsu, 1994. j Shiau and Hwang, 1994. k Shiau and Liu, 1994a. l Deshimaru and Kuroki, 1979. m NRC, 1983. n Kanazawa et al., 1976. o Deshimaru and Kuroki, 1976. p Guary et al., 1976. q Shigueno and Itoh, 1988. r Liu et al., 1993. s Shiau and Liu, 1994b. t He and Lawrence, 1991. u He and Lawrence, 1993. v Lightner et al., 1979. b

was reported for P. japonicus ŽKanazawa et al., 1976.. The choline requirement for P. japonicus was studied by Deshimaru and Kuroki Ž1979., using a choline-deficient diet showed a similar growth to those fed on choline-supplemented diets. They also found

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that body choline concentration was high regardless of the level of dietary choline chloride and concluded that this vitamin is dispensable for the growth of P. japonicus. This result was contradictory to the findings of Kanazawa et al. Ž1976. in which the choline-deficient diet resulted in poor growth and high mortality of the shrimp. The reason for such discrepancy between the two studies is still not known. No further study has yet been done to clarify this. L-Ascorbic acid ŽC1. is the only source of vitamin C traditionally used in shrimp feed. Lightner et al. Ž1977. demonstrated for the first time that penaeid shrimp Ž P. californiensis. developed the ‘black death syndrome’ when fed a C1-free diet. Later work by Magarelli and Colvin Ž1978. added P. stylirostris to the list of penaeids that require dietary ascorbic acid and develop ‘black death syndrome’ if deficient in vitamin C. The Kuruma shrimp, P. japonicus, was shown to require 3000 mgrkg diet ŽDeshimaru and Kuroki, 1976. or 10,000–20,000 mgrkg diet ŽGuary et al., 1976. of C1 for optimal growth. The C1 requirement for maximum growth of P. californiensis is 2000 mgrkg diet ŽLightner et al., 1979.. For P. monodon, a requirement of 2000 mgrkg diet was suggested; scorbutic shrimp suffered reduced growth rate and ‘black death syndrome’ ŽShiau and Jan, 1992.. C 1 is unstable, and practical diets have been shown to lose ascorbic acid during processing and storage due to exposure to high temperature, oxygen and light ŽHilton et al., 1977; Lovell and Lim, 1978; Soliman et al., 1987.. Shiau and Hsu Ž1993. found that about 75% of the initial amount of supplemental C 1 in shrimp feeds can be lost during processing at ambient temperature. Attempts have been made to increase retention of vitamin C activity in shrimp feeds by using alternative forms of ascorbic acid. L-Ascorbic acid derivatives with sulfate and phosphate moieties at the unstable C-2 position in the lactone ring are highly resistant to oxidation ŽTolbert et al., 1975.. Studies have shown beneficial effects of using alternative forms of ascorbic acid in meeting penaeid shrimp requirements. For example, the requirement for P. monodon significantly decreased from 2000 mgrkg diet for L-ascorbic acid ŽC1; Shiau and Jan, 1992. to 210 mgrkg diet for polyphosphated form ŽC2PP. ŽChen and Chang, 1994., to 100–200 mgrkg for phosphated Mg form ŽC2PMg. ŽCatacutan and Lavilla-Pitogo, 1994. or to 40 mgrkg for monophosphated form ŽC2MP. ŽShiau and Hsu, 1994.. The requirement of P. japonicus for C2PMg was 215–430 mgrkg ŽShigueno and Itoh, 1988. and of P. Õannamei for C2PP was 90-120 mgrkg ŽHe and Lawrence, 1993.. The potency of each alternative form for the penaeid shrimp is critical in calculating the supplemental level. Shiau and Hsu Ž1994. suggested that L-ascorbyl-2-sulfate ŽC2S. is about 25% as effective as C2MP in meeting vitamin C requirement for P. monodon.

6. Minerals For both human and animal nutrition, it is important to establish minimum requirement and maximum tolerance for an element to secure optimal growth and health. In shrimp, essential minerals may be obtained from water by exchange across the gill membrane or ingestion and by absorption across the gut. Nevertheless, it is generally considered that a dietary source of some minerals for growth is necessary because of the

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Table 4 Mineral requirements of juvenile P. japonicus Minerals

Ca P K Mg Mn Fe Cu

Requirement Ž%. Deshimaru and Yone Ž1978c.

Kanazawa et al. Ž1984.

dispensable 2.0 1.0 dispensable y dispensable y

1.0 1.0 0.9 0.3 dispensable dispensable 0.6

repeated losses of certain minerals during molting. Requirement studies normally involve experiments where animal responses or performance characteristics are studied relative to feeding graded levels of an essential mineral over a wide range, from zero to levels far beyond optimal. Because many minerals are required in small quantities, it is difficult to formulate basal diets and maintain environments that are free of the test minerals to be able to conduct requirement studies. Thus, investigations on mineral requirements of shrimps are still very limited. The requirements for several minerals have been quantified for P. japonicus and P. Õannamei. Other than these two species, studies on the mineral nutrition of P. aztecus and P. californiensis have also been done but to a lesser extent. Tables 4 and 5 summarize the mineral requirements of P. japonicus and P. Õannamei, respectively. Deshimaru et al. Ž1978. have shown that P. japonicus takes in calcium from seawater and does not require calcium, magnesium and iron from dietary source. Kanazawa et al. Ž1984. have reported that addition of calcium to diets would be necessary to maintain the 1:1 ratio of calcium–phosphorus in diets, although growth of P. japonicus on diets with and without calcium supplementation is comparable. Kitabayashi et al. Ž1971. have also pointed out the importance of the CarP ratio, indicating an optimum ratio of 1:1 for P. japonicus. Huner and Colvin Ž1977. have shown that a CarP ratio of 2.2:1 is optimum for growth of P. californiensis. The necessity of phosphorus has been manifested in P. japonicus ŽKitabayashi et al., 1971; Deshimaru and Yone, 1978c; Kanazawa et al., 1984.. Deshimaru and Yone Ž1978c.

Table 5 Mineral requirements of juvenile P. Õannamei Minerals

Requirement Ž%. Davis et al. Ž1993a.

Davis et al. Ž1993b.

Ca P

dispensable 0.35 Ž0% Ca. 0.5–1.0 Ž1% Ca. 1.0–2.0 Ž2% Ca. y

y y

Cu

0.0032

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have reported that P. japonicus requires phosphorus Ž2.0%., potassium Ž1.0%., and trace metals Ž0.2%.. Kanazawa et al. Ž1984. have shown that this species requires calcium Ž1.0%., phosphorus Ž1.0%., magnesium Ž0.3%., potassium Ž0.9%. and copper Ž0.6%. in dry diets. Shewbart et al. Ž1973. indicated that calcium, potassium and sodium chloride are not necessary for P. aztecus, but phosphorus may be essential. Davis et al. Ž1993a. demonstrated that a dietary calcium supplement is not required for P. Õannamei and the dietary phosphorus requirement for this species is dependent upon the calcium content of the diet. In the absence of calcium supplement, the basal diet containing 0.35% phosphorus is adequate to maintain good growth and survival of shrimp. In the presence of 1.0 and 2.0% supplemental calcium, supplementation of 0.5–1.0% phosphorus and 1.0–2.0% phosphorus, respectively, are required to maintain normal growth of the shrimp. The dietary copper requirement of P. Õannamei has been reported to be 0.0032% ŽDavis et al., 1993b..

7. Conclusion The nutrition of penaeid shrimps has been reviewed by Kanazawa Ž1984. for the First International Conference on the Culture of Penaeid PrawnsrShrimps. Since then, progress in many aspects of penaeid nutrition has been achieved. So far, P. japonicus, P. monodon and P. Õannamei are the three penaeid species that have been studied thoroughly for their nutrient requirements. Despite many progress in recent years, knowledge on the nutritional requirements of penaeid shrimps is still far from complete. Among the nutrients that are indispensable to the penaeids, the essential amino acid and mineral requirements are the two areas that need more research work. Furthermore, variation in nutrient requirement among the penaeid species, such as vitamin K, niacin, riboflavin, choline and others may imply that more research work is needed on dealing the individual species. Nutrient interactions such as one nutrient affecting the requirement for another nutrient as has been demonstrated in a few examples in the literature certainly need to be explored further. Studies to examine the relationships between nutrition and disease, and the environmental factors affecting nutrient requirements should be given more attention.

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