A review of the development and application of soybean‐based diets ...

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Aquaculture Nutrition 2013 19; 441–448

doi: 10.1111/anu.12050

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Department of Fisheries and Allied Aquacultures, Auburn University, Auburn, AL, USA

The production of the Pacific white shrimp (Litopenaeus vannamei) has expanded to the point of being the most widely cultured species of shrimp. One of the advantages of this species is its acceptance of a wide variety of feed formulations including plant-based feeds. Given the increases in ingredient costs, particularly fish meal, there is considerable interest in the use of alternative feed formulations for cultured species. Given that soybean meal is one of the most widely available protein sources for which production can be expanded, the move to soy-based diets is inevitable. The successful use of alternative feed ingredients for shrimp production depends on a number of factors. This paper summarizes studies regarding the move towards high soy diets concerning manipulation of ingredients and nutrient profiles to maintain balanced feed formulations. KEY WORDS:

alternative feed, practical diets, soybean, van-

namei Received 9 September 2011, accepted 20 January 2013 Correspondence: D.A. Davis, Department of Fisheries and Allied Aquacultures, 203 Swingle Hall, Auburn University, Auburn, AL 36849-5419, USA. E-mail: [email protected]

Pacific white shrimp, Litopenaeus vannamei (Boone) is native to the eastern Pacific Ocean from Sonora, Mexico to Northern Peru. Currently, it is the most popular cultured shrimp species and has experienced a dramatic increase in aquaculture production from 186 113 tonnes in 1999 to 2 296 630 tonnes in 2007 (FAO 2009). The industry growth has been paralleled by an increase in shrimp feed production. The increases in demand and limitations of supply

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ª 2013 John Wiley & Sons Ltd

have resulted in some ingredients becoming less available and more costly, especially fish meal and fish oil. Fish meal and other marine ingredients are considered desirable ingredients in shrimp feed because of their nutrient content and palatability. In commercial feeds, fishmeal typically accounts for 200–300 g kg 1 of the shrimp feed formulation (Tacon & Metian 2008). The cost of fish meal and fish oil has generally increased over time as a result of the uncertainty of availability and large fluctuations in the price. Furthermore, there are growing social and environment concerns regarding the long-term sustainability of the use of marine ingredients. In addition to feed prices increasing, the market value for shrimp has declined because of increased production and limited demand. This has resulted in a reduction in the profit margin for shrimp farmers. When margins were good, feed manufacturers could afford to use expensive ingredients and over formulate a diet. However, as the margin decreased, feeds must become more cost-effective. Feed costs can account for as much as 40–60% of production costs (Hertrampf & Piedad-Pascual 2000). Feed costs and feed management both influence the investment in feeds. Reducing or removing costly protein sources through the use of a combination of less expensive and more economical protein and lipid sources could result in substantial saving in feed cost. Practical diets using plant-based ingredients to replace fish meal and fish oil have become an interesting alternative which could reduce these problems. The use of renewable plant protein sources has become the focus of protein substitution studies in shrimp feeds around the world because of their acceptable protein level, suitable amino acid content, economic opportunity and consistent quality (Watanabe 2002). Formulated diets are designed to contain sufficient levels of nutrients to meet requirements using plant-based protein sources for which production can be expanded and are often more

cost-effective. Feeding plant-based proteins to shrimp requires that the ingredients possess certain nutritional characteristics, such as low levels of fibre, starch (especially insoluble carbohydrates) and antinutrients. They must also contain a relatively high protein content, favourable amino acid profile, high nutrient digestibility and reasonable palatability (Gatlin et al. 2007; Naylor et al. 2009). Ten indispensable amino acids that are required for growth and maintenance of shrimp are arginine, histidine, isoleucine, leucine, lysine, methionine, phenylalanine, threonine, tryptophan and valine (Kanazawa 1989; Guillaume 1997). These amino acids should satisfy shrimp requirements to support optimum growth performance. Fish meal is considered ideal protein source for fish and shrimp feed production because of its high level of essential amino acids. On the other hand, plant protein sources contain lower levels of some essential amino acids (Tacon 1994). Thus, the balance of essential amino acids must be considered when diets are formulated to contain plant protein sources to replace fish meal. In general, the amino acid profile of soybean meal is comparable with that of fish meal, albeit is lower in sulphur amino acids, that is, methionine and cystine (Peres & Lim 2008). Soybean meal is often considered as the most reliable ingredient and cost-effective protein source in shrimp feed because of its high protein content, high digestibility, relatively well-balanced amino acid profile, reasonable price and steady supply (Davis & Arnold 2000; Amaya et al. 2007a,b). The protein digestibility was found higher in soybean protein than that in the marine animal meals (Akiyama 1989). Ezquerra et al. (1997, 1998) reported in vivo and in vitro protein digestibility by pH drop of feed using the white shrimp hepatopancreas ranged from 64%

to 91% where soybean protein showed greater APD than those in fish meal or crab meal (Table 1). However, the inclusion of soybean meal at high levels or as a sole protein source has resulted in reduced performance of the shrimp (Lim & Dominy 1990). This could be the results of imbalanced amino acid profiles or deficiencies of other dietary nutrients that were not taken into account. Fish meal is utilized as a protein source but it also provides lipids, essential fatty acids (EFAs), minerals and vitamins to the diet. Consequently, there will be most likely a need to use a variety of feed ingredients in association with soybean meal to provide a better balanced nutrient profile. Utilization of various potential protein sources in shrimp feeds such as animal by-product and other plant sources (listed in table 2) has been evaluated under different rearing conditions (Lim & Dominy 1990; Piedad-Pascual et al. 1990; Sudaryono et al. 1995; Cruz-Suarez et al. 2001; Amaya et al. 2007a,b; Ray et al. 2009). One of those ingredients that are considered a promising alternative for the substitution of fish meal in shrimp feeds is poultry by-product meal (Davis & Arnold 2000; Samocha et al. 2004; Amaya et al. 2007a; Markey 2007). Distiller’s dried grains with solubles (DDGS) is also a potential protein source for shrimp feed because of its low cost and consistent supply as a coproduct of the bio-ethanol production, which is expected to increase rapidly in the next decade. Several studies reported the successful use of DDGS as an alternative protein source in fish and crustacean feeds without causing negative impact on growth performance (Webster et al. 1991, 1992; Wu et al. 1994; Cheng & Hardy 2004; Coyle et al. 2004; Stone et al. 2005; Lim et al. 2007, 2009; Robinson & Li 2008; Thompson et al. 2008). Pea meal is also another widely used feed

Table 1 Chemical composition of the test ingredients and in vivo and in vitro protein digestibility of L. vannamei fed different protein sources. Results expressed as meanSD

Ingredients

Moisture1

Chilean anchovy Deboned white fish Langostilla Mexican tuna waste Menhaden fish meal A Menhaden fish meal B Soybean protein

104 54 59 49 78 83 93

      

3 3 10 4 5 2 5

Crude protein1 602 757 394 613 638 609 495

      

1 9 10 4 10 10 10

Crude fat1

Ash1

      

148 93 426 215 181 159 215

125 72 29 64 119 145 53

6 1 6 4 6 2 7

      

1 0.5 1 1 5 4 3

In vivo2 digestibility

In vitro3 digestibility

83.6 86.6 66.4 63.6 67.1 68.4 90.9

81.5 86.3 75.7 70.0 70.5 64.9 83.7

1

g kg 1 dry matter basis. Determined by chromic oxide method using 3.5–4 g shrimp, N = 3 tanks/treatment (from Ezquerra et al. 1997). 3 Determined by pH drop at 27 °C of feed using enzymes extracted from hepatopancreas of 10 to 12 g white shrimp (from Ezquerra et al. 1998). 2

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Aquaculture Nutrition 19; 441–448 ª 2013 John Wiley & Sons Ltd

Table 2 Chemical composition of the test ingredients (as-fed basis) (NRC 2011) Ingredient Fish meal Anchovy Menhaden White DDGS Pea meal Poultry by-product Soybean meal 44% CP 48% CP Soy protein concentrate

Dry matter (g kg 1)

Crude protein (g kg 1)

Crude fat (g kg 1)

Crude fibre (g kg 1)

Ash (g kg 1)

920 920 920 910 900 890

654 645 620 270 253 559

76 96 78 93 14 136

10 7 8 91 69 21

143 190 213 64 33 145

890 900 920

440 485 636

15 9 5

73 34 45

63 58 –

ingredient, mostly in livestock because of its high energy, moderate protein level (220–260 g kg 1 crude protein), amino acid profile and low cost (Borlongan et al. 2003). Several studies indicated that feed pea is another potential ingredient in fish and shrimp feeds (Gomes et al. 1995; Burel et al. 2000; Carter & Hauler 2000; Gouveia & Davies 2000; Booth et al. 2001; Cruz-Suarez et al. 2001; Davis et al. 2002; Bautista-Teruel et al. 2003; Borlongan et al. 2003). Because of the limitation in nutrient component of most ingredients, more than one ingredient is required for balanced feed formulations. Therefore, shrimp diets containing soybean meal as a main protein source should be combined with other alternative protein ingredients, that is, poultry by-product meal, DDGS and pea meal. Soybean and its products are acceptable protein sources with good digestibility for shrimp. However, soybean meal is deficient in the essential amino acids (EAAs) such as methionine, lysine and tryptophan as well as essential fatty acids and minerals (Lim & Dominy 1990). Methionine is one of the ten essential or indispensable amino acids that are dietary essential for shrimp (Millamena et al. 1996). Thus, supplementation of sulphur amino acids, that is, methionine or cystine, in soybean-based diets to meet the shrimp requirement is recommended to provide a good growth response (Akiyama 1989). Low levels of methionine found in soybean meal can also be countered by mixing with other protein sources and/or the supplementation of synthetic methionine. Several studies had reported successfully replacing fish meal with soybean meal with a methionine supplement in Milkfish (Davis et al. 1995; Shiau et al. 2007). Conversely, a diet containing only soybean protein with a methionine supplement was poorly utilized by red drum (Reigh & Ellis 1992). McGoogan & Gatlin (1997) suggested that diets containing soybean meal with low levels or no fish meal may have palatability problems. Thus,

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the inclusion of attractants or palatability enhancers, for example, fish solubles, may be considered. A reduction in feed intake was reported in largemouth bass fed diets with increased soybean meal levels (Cho et al. 1974; Kubitza et al. 1997). Similar results were observed in red drum (Reigh & Ellis 1992; Davis et al. 1995) and Pacific white shrimp (Lim & Dominy 1990). Along with protein, lipids constitute the major macronutrients that are required to provide the energy and cellular building blocks as well as maintain growth, health, welfare and reproduction in shrimp (Lim et al. 1997). Reducing or replacement of marine ingredients that are good sources of high quality oils from shrimp feed formulation may result in EFAs deficiencies. As we replace fish meal with alternative ingredients, for example soybean meal, we must ensure that we meet the shrimp EFAs requirements. Lipid content and the associated C18 PUFA (poly unsaturated fatty acids), linoleic (18:2n-6) and linolenic (18:3n-3) acids, as well as n-3 and n-6HUFA (highly unsaturated fatty acids), eicosapentaenoic acid (EPA, 20:5n-3), docosahexaenoic acid (DHA, 22:6n-3) and arachidonic acid (ARA, 20:4n-6) are required in shrimp and other crustacean feeds at levels between 5 and 10 g kg 1 (Akiyama et al. 1991; Gonz alez-Felix & Perez-Velazquez 2002). Generally, the primary protein and lipid sources used in practical shrimp feeds are fish meal and fish oil (Cheng et al. 2002). Lim et al. (1997) have reported that menhaden oil rich in n-3 HUFA (20:5n-3 and 22:6n-3) was better utilized by Penaeus vannamei than vegetable lipid sources such as linseed, sunflower, corn, soybean and coconut oil and stearic acid. Samocha et al. (2010) also suggested that the supplementation of HUFAs is a critical component to replace marine fish oil in shrimp feed. Their results demonstrated that the complete replacement of fish meal and fish oil using non-marine ingredients can be accomplished using supplementation of plant oils with DHA- and

ARA-rich oils from fermented products. Other studies have reported that partial or total placement of fish meal and fish oil with soybean meal and soy oil had no adverse effect on shrimp growth performance (Davis & Arnold 2000; Samocha et al. 2004; Gonzalez-Felix et al. 2010), but shrimp body crude fat and cholesterol concentration were reduced (Cheng & Hardy 2004). According to Gonz alezFelix et al. (2010), the substitution of fish oil up to 90% by plant-based oils in diets can be done without a significant reduction in growth performance, FCR, production yield and survival in L. vannamei. Apparently, this 10% of fish oil remaining in the diet supply enough of the essential fatty acids ARA, EPA and DHA for the proper development of this species, although the fatty acid composition of the muscular tissue of the animal reflected the lipid source fatty acid profile added to the diet followed by a reduction in HUFAs as fish oil was replaced. High levels of n-3 fatty acids can be obtained with the use of linseed oil, most of it comprised of the PUFA a-linolenic acid; yet, the levels of the essential HUFAs in linseed oil are found at low levels. Cholesterol is a vital component of cell membranes. It is the precursor of bile acids, steroids, and moulting hormones (Cheng & Hardy 2004). It is reported to be an essential nutrient for growth and survival of shrimp (Kanazawa et al. 1971; Gong et al. 2000; Morris et al. 2011). Gong et al. (2000) suggested that dietary cholesterol requirement of L. vannamei juveniles was affected by dietary phospholipids such as soybean lecithin and purified phosphatidylcholine. Phospholipids are considered another dietary necessity for optimum shrimp growth. Dietary cholesterol and phospholipids interact to improve growth as well as affect retention of total lipid and triglycerides in hepatopancreas and cholesterol in muscle of L. vannamei juveniles. Several studies have indicated a clear need for cholesterol supplementation in plant-based diets (Gong et al. 2000; Morris et al. 2011). Gong et al. (2000) suggested that optimal growth of L. vannamei was obtained with 3.5 g kg 1, 1.4 g kg 1, 1.3 g kg 1 and 0.5 g kg 1 supplemental cholesterol at dietary PL levels of 0, 15, 30 and 50 g kg 1, respectively, in dietary treatments containing no fish meal. Similar results reported by Morris et al. (2011) demonstrated that the cholesterol supplements in dietary treatments formulated with no fish meal and targeted crude protein levels of 350 g kg 1 for L. vannamei were between 0.2 and 0.4 g kg 1, thus containing a cholesterol level between 0.76 and 1.1 g kg 1 of diet. Shrimp are able to assimilate minerals directly from the aquatic environment (Montoya et al. 2000). In shrimp, minerals serve as structural components of hard and soft

tissues and metalloproteins as well as enzymatic cofactors and enzymatic activators (Davis & Lawrence 1997). Shrimp can utilize some soluble minerals such as calcium, copper, iron, magnesium, phosphorus, potassium, selenium, sodium and zinc from the water through the gill, epidermis or both. Generally, phosphorus is found at low concentration in natural water relative to its requirement by phytoplankton (Boyd 2007). When fish meal is replaced by soybean meal, the first limiting mineral in shrimp feed formulation is phosphorus as only 30–40% of total phosphorus content in soybean meal is available for L. vannamei (Hertrampf & Piedad-Pascual 2000). Therefore, supplemental phosphorus is essential for optimal shrimp growth as fishmeal was removed. The dietary phosphorus requirement for juvenile L. vannamei ranges from 3.4 to 20 g kg 1 (Davis et al. 1993) and 20.9 – 22.0 g kg 1 for postlarval L. vannamei (Niu et al. 2008). The dietary phosphorus requirement for shrimp is dependent on the calcium content in diet although a dietary calcium supplement is not required (Davis et al., 1993; Cheng et al. 2006). Compared with fish meal, soybean meal is found to have low availability of selenium, 48.0% and 17.5%, respectively (Gabrielsen & Opstvedt 1980). Selenium is an essential trace element that functions as a component of the enzyme glutathione peroxidase in shrimp, but it can be toxic (Davis & Gatlin 1996; Wang et al. 2006). Glutathione peroxidase converts hydrogen peroxide and lipid hydroperoxides into water and lipid alcohols, respectively, thus protecting the cell from the deleterious effects of peroxides (Davis & Gatlin 1996). Juvenile P. vannaemi was found to grow best when fed semi-purified diets supplemented with 0.2–0.4 mg Se kg 1 diet (Davis & Gatlin 1996). Supplemental selenium is not required in practical diets containing more than 150 g kg 1 fish meal. Therefore, selenium supplementation may be required in diets formulated with predominantly plant ingredients. Due to potentially toxic effects, selenium supplementation of 0.1 mg kg 1 is approved to be used with fish and crustaceans (Davis & Gatlin 1996). There are other issues of using soybean meal as an alternative to fishmeal besides nutritional factors such as the presence of nutrient inhibitors. Raw soybean contains antinutritional factors such as trypsin inhibitors, lectins, oligosaccharides, antigens and saponins that may affect the digestion and reduce nutrient availability to shrimp (Dersjant-Li 2002). However, the effect of some of these antinutrients can be reduced by heat process (New 1987). Clearly, the use of soybean meal in shrimp feed is feasible. However, there are several other plant protein sources that may be considered as alternative ingredients used in

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association with soybean meal to balance nutritional composition in feed formulations when non-fishmeal diets are formulated. It appears that fish meal can be partially or completely removed from shrimp formulations if suitable alternative sources of protein and lipids are provided to meet the nutritional requirements of the animal (Lim & Dominy 1990, 1992; Piedad-Pascual et al. 1990; Sudaryono et al. 1995; Cruz-Suarez et al. 2001; Smith et al. 2001; Davis et al., 2004; Samocha et al. 2004; Amaya et al. 2007a,b; Roy et al., 2009). Recently, National Research Council (NRC 2011) reported the minimum nutrient requirements for maximum performance of L. vannamei (Table 3). Yet, there is still limited information available on amino acid requirement data for L.vannamei, as well as fatty acids, vitamins and minerals are highly digestible; therefore, the values presented represent nearly 100% bioavailability. The use of complementary ingredients is a practice used to obtain a more balanced nutrient profile in the feeds (i.e. essential amino acids, fatty acids, minerals) and to increase nutrient utilization and facilitate feed processing (Amaya et al. 2007a). Sookying (2010) reported on a series of studies that demonstrated and developed a range of soy-based diets. This includes diets that demonstrate that fish meal Table 3 Nutrient requirements of L. vannamei (dry matter basis) (from NRC 2011)

Item Typical energy and protein concentrations Digestible energy (kcal kg 1 diet) Digestible protein (g kg 1) Amino acid (g kg 1) Lysine Fatty acids (g kg 1) n-3 LC-PUFA Cholesterol (g kg 1) Macrominerals (g kg 1) Magnesium Phosphorus Microminerals (mg kg 1) Copper Selenium Zinc Fat-soluble vitamins A (mg kg 1) E (mg kg 1) Water-soluble vitamins (mg kg 1) Vitamin B6 Vitamin C

1

Minimum requirement for L. vannamei

3000 300 16 2.5–5.0 g kg 1.3

1

2.6–3.5 3–7 16–32 0.2–0.4 15 1.4 100 80–100 50–100

1 These requirements have been determined with highly purified ingredients in which the nutrient composition has been defined.

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(100 g kg 1 diet) could be totally removed from diets for L. vannamei by a combination of plant and animal protein sources (soybean meal and poultry by-product meal) or all plant protein sources (soybean meal in combination with DDGS or pea meal with the inclusion of corn gluten meal and squid meal) when diets are formulated to contain acceptable nutrient levels and proper balanced nutrients without any apparent effect on survival, growth and feed palatability (Sookying & Davis 2011). They also demonstrated that up to 120 g kg 1 soy protein concentrate could be used in a high soy diet under outdoor production conditions without an effect on production performance of the shrimp (Sookying & Davis 2012). Alternative feed formulations for the pacific white shrimp seem to work across a number of culture technologies (clear water research systems, outdoor tank systems and research ponds) as well as across a range of densities in outdoor ponds (Sookying et al. 2011). Given the range of culture systems and densities, the use of alternative feed formulations for this species is warranted and appropriate for commercial production.

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Aquaculture Nutrition 19; 441–448 ª 2013 John Wiley & Sons Ltd

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