IntroductionGood nutrition in animal production systems is essential to economically produce a healthy, high quality product. In fish farming, nutrition is critical because feed represents 40-50% of the production costs. Fish nutrition has advanced dramatically in recent years with the development of new, balanced commercial diets that promote optimal fish growth and health. The development of new species-specific diet formulations supports the aquaculture (fish farming) industry as it expands to satisfy increasing demand for affordable, safe, and high-quality fish and seafood products.
Prepared (artificial) Diets
Prepared or artificial diets may be either complete or supplemental. Complete diets supply all the ingredients (protein, carbohydrates, fats, vitamins, and minerals) necessary for the optimal growth and health of the fish. Most fish farmers use complete diets, those containing all the required protein (18-50%), lipid (10-25%), carbohydrate (15-20%), ash (< 8.5%), phosphorus (< 1.5%), water (< 10%), and trace amounts of vitamins, and minerals. When fish are reared in high density indoor systems or confined in cages and cannot forage freely on natural feeds, they must be provided a complete diet.
In contrast, supplemental (incomplete, partial) diets are intended only to help support the natural food (insects, algae, small fish) normally available to fish in ponds or outdoor raceways. Supplemental diets do not contain a full complement of vitamins or minerals, but are used to help fortify the naturally available diet with extra protein, carbohydrate and/or lipid.
Fish, especially when reared in high densities, require a high-quality, nutritionally complete, balanced diet to grow rapidly and remain healthy.
Because protein is the most expensive part of fish feed, it is important to accurately determine the protein requirements for each species and size of cultured fish. Proteins are formed by linkages of individual amino acids. Although over 200 amino acids occur in nature, only about 20 amino acids are common. Of these, 10 are essential (indispensable) amino acids that cannot be synthesized by fish. The 10 essential amino acids that must be supplied by the diet are: methionine, arginine, threonine, tryptophan, histidine, isoleucine, lysine, leucine, valine and phenylalanine. Of these, lysine and methionine are often the first limiting amino acids. Fish feeds prepared with plant (soybean meal) protein typically are low in methionine; therefore, extra methionine must be added to soybean-meal based diets in order to promote optimal growth and health. It is important to know and match the protein requirements and the amino acid requirements of each fish species reared.
Protein levels in aquaculture feeds generally average 18-20% for marine shrimp, 28-32% for catfish, 32-38% for tilapia, 38-42% for hybrid striped bass. Protein requirements usually are lower for herbivorous fish (plant eating) and omnivorous fish (plant-animal eaters) than they are for carnivorous (flesh-eating) fish, and are higher for fish reared in high density (recirculating aquaculture) than low density (pond aquaculture) systems.
Protein requirements generally are higher for smaller fish. As fish grow larger, their protein requirements usually decrease. Protein requirements also vary with rearing environment, water temperature and water quality, as well as the genetic composition and feeding rates of the fish. Protein is used for fish growth if adequate levels of fats and carbohydrates are present in the diet. If not, protein may be used for energy and life support rather than growth.
Proteins are composed of carbon (50%), nitrogen (16%), oxygen (21.5%), and hydrogen (6.5%). Fish are capable of using a high protein diet, but as much as 65% of the protein may be lost to the environment. Most nitrogen is excreted as ammonia (NH3) by the gills of fish, and only 10% is lost as solid wastes. Accelerated eutrophication (nutrient enrichment) of surface waters due to excess nitrogen from fish farm effluents is a major water quality concern of fish farmers. Effective feeding and waste management practices are essential to protect downstream water quality.
Lipids (fats) are high-energy nutrients that can be utilized to partially spare (substitute for) protein in aquaculture feeds. Lipids supply about twice the energy as proteins and carbohydrates. Lipids typically comprise about 15% of fish diets, supply essential fatty acids (EFA) and serve as transporters for fat-soluble vitamins.
A recent trend in fish feeds is to use higher levels of lipids in the diet. Although increasing dietary lipids can help reduce the high costs of diets by partially sparing protein in the feed, problems such as excessive fat deposition in the liver can decrease the health and market quality of fish.
Simple lipids include fatty acids and triacylglycerols. Fish typically require fatty acids of the omega 3 and 6 (n-3 and n-6) families. Fatty acids can be: a) saturated fatty acids (SFA, no double bonds), b) polyunsaturated fatty acids (PUFA, >2 double bonds), or c) highly unsaturated fatty acids (HUFA; > 4 double bonds). Marine fish oils are naturally high (>30%) in omega 3 HUFA, and are excellent sources of lipids for the manufacture of fish diets. Lipids from these marine oils also can have beneficial effects on human cardiovascular health.
Marine fish typically require n-3 HUFA for optimal growth and health, usually in quantities ranging from 0.5-2.0% of dry diet. The two major EFA of this group are eicosapentaenoic acid (EPA: 20:5n-3) and docosahexaenoic acid (DHA:22:6n-3). Freshwater fish do not require the long chain HUFA, but often require an 18 carbon n-3 fatty acid, linolenic acid (18:3-n-3), in quantities ranging from 0.5 to 1.5% of dry diet. This fatty acid cannot be produced by freshwater fish and must be supplied in the diet. Many freshwater fish can take this fatty acid, and through enzyme systems elongate (add carbon atoms) to the hydrocarbon chain, and then further desaturate (add double bonds) to this longer hydrocarbon chain. Through these enzyme systems, freshwater fish can manufacture the longer chain n-3 HUFA, EPA and DHA, which are necessary for other metabolic functions and as cellular membrane components. Marine fish typically do not possess these elongation and desaturation enzyme systems, and require long chain n-3 HUFA in their diets. Other fish species, such as tilapia, require fatty acids of the n-6 family, while still others, such as carp or eels, require a combination of n-3 and n-6 fatty acids
Carbohydrates (starches and sugars) are the most economical and inexpensive sources of energy for fish diets. Although not essential, carbohydrates are included in aquaculture diets to reduce feed costs and for their binding activity during feed manufacturing. Dietary starches are useful in the extrusion manufacture of floating feeds. Cooking starch during the extrusion process makes it more biologically available to fish.
In fish, carbohydrates are stored as glycogen that can be mobilized to satisfy energy demands. They are a major energy source for mammals, but are not used efficiently by fish. For example, mammals can extract about 4 kcal of energy from 1 gram of carbohydrate, whereas fish can only extract about 1.6 kcal from the same amount of carbohydrate. Up to about 20% of dietary carbohydrates can be used by fish.
Vitamins are organic compounds necessary in the diet for normal fish growth and health. They often are not synthesized by fish, and must be supplied in the diet.
The two groups of vitamins are water-soluble and fat-soluble. Water-soluble vitamins include: the B vitamins, choline, inositol, folic acid, pantothenic acid , biotin and ascorbic acid (vitamin C). Of these, vitamin C probably is the most important because it is a powerful antioxidant and helps the immune system in fish.
The fat-soluble vitamins include A vitamins, retinols (responsible for vision); the D vitamins, cholecaciferols (bone integrity); E vitamins, the tocopherols (antioxidants); and K vitamins such as menadione (blood clotting, skin integrity). Of these, vitamin E receives the most attention for its important role as an antioxidant. Deficiency of each vitamin has certain specific symptoms, but reduced growth is the most common symptom of any vitamin deficiency. Scoliosis (bent backbone symptom) and dark coloration may result from deficiencies of ascorbic acid and folic acid vitamins, respectively.
Minerals are inorganic elements necessary in the diet for normal body functions. They can be divided into two groups (macro-minerals and micro-minerals) based on the quantity required in the diet and the amount present in fish. Common macro-minerals are sodium, chloride, potassium and phosphorous. These minerals regulate osmotic balance and aid in bone formation and integrity.
Micro-minerals (trace minerals) are required in small amounts as components in enzyme and hormone systems. Common trace minerals are copper, chromium, iodine, zinc and selenium. Fish can absorb many minerals directly from the water through their gills and skin, allowing them to compensate to some extent for mineral deficiencies in their diet.
Energy and Protein
Dietary nutrients are essential for the construction of living tissues. They also are a source of stored energy for fish digestion, absorption, growth, reproduction and the other life processes. The nutritional value of a dietary ingredient is in part dependant on its ability to supply energy. Physiological fuel values are used to calculate and balance available energy values in prepared diets. They typically average 4, 4, and 9 kcal/g for protein, carbohydrate and lipid, respectively.
To create an optimum diet, the ratio of protein to energy must be determined separately for each fish species. Excess energy relative to protein content in the diet may result in high lipid deposition. Because fish feed to meet their energy requirements, diets with excessive energy levels may result in decreased feed intake and reduced weight gain. Similarly, a diet with inadequate energy content can result in reduced weight gain because the fish cannot eat enough feed to satisfy their energy requirements for growth. Properly formulated prepared feeds have a well-balanced energy to protein ratio.
Commercial fish diets are manufactured as either extruded (floating or buoyant) or pressure-pelleted (sinking) feeds. Both floating or sinking feed can produce satisfactory growth, but some fish species prefer floating, others sinking. Shrimp, for example, will not accept a floating feed, but most fish species can be trained to accept a floating pellet.
Extruded feeds are more expensive due to the higher manufacturing costs. Usually, it is advantageous to feed a floating (extruded) feed, because the farmer can directly observe the feeding intensity of his fish and adjust feeding rates accordingly. Determining whether feeding rates are too low or too high is important in maximizing fish growth and feed use efficiency.
Feed is available in a variety of sizes ranging from fine crumbles for small fish to large (1/2 inch or larger) pellets. The pellet size should be approximately 20-30% of the size of the fish species mouth gape. Feeding too small a pellet results in inefficient feeding because more energy is used in finding and eating more pellets. Conversely, pellets that are too large will depress feeding and, in the extreme, cause choking. Select the largest sized feed the fish will actively eat.
Feeding Rate, Frequency, and Timing
Feeding rates and frequencies are in part a function of fish size. Small larval fish and fry need to be fed a high protein diet frequently and usually in excess. Small fish have a high energy demand and must eat nearly continuously and be fed almost hourly. Feeding small fish in excess is not as much of a problem as overfeeding larger fish because small fish require only a small amount of feed relative to the volume of water in the culture system.
As fish grow, feeding rates and frequencies should be lowered, and protein content reduced. However, rather than switching to a lower protein diet, feeding less allows the grower to use the same feed (protein level) throughout the grow-out period, thereby simplifying feed inventory and storage.
Feeding fish is labor-intensive and expensive. Feeding frequency is dependent on labor availability, farm size, and the fish species and sizes grown. Large catfish farms with many ponds usually feed only once per day because of time and labor limitations, while smaller farms may feed twice per day. Generally, growth and feed conversion increase with feeding frequency. In indoor, intensive fish culture systems, fish may be fed as many as 5 times per day in order to maximize growth at optimum temperatures.
Many factors affect the feeding rates of fish. These include time of day, season, water temperature, dissolved oxygen levels, and other water quality variables. For example, feeding fish grown in ponds early in the morning when the lowest dissolved oxygen levels occur is not advisable. In contrast, in recirculating aquaculture systems where oxygen is continuously supplied, fish can be fed at nearly any time. During the winter and at low water temperatures, feeding rates of warmwater fishes in ponds decline and feeding rates should decrease proportionally.
Feed acceptability, palatability and digestibility vary with the ingredients and feed quality. Fish farmers pay careful attention to feeding activity in order to help determine feed acceptance, calculate feed conversion ratios and feed efficiencies, monitor feed costs, and track feed demand throughout the year.
Published feeding rate tables are available for most commonly cultured fish species. Farmers can calculate optimum feeding rates based on the average size in length or weight and the number of fish in the tank, raceway, or pond (see Hinshaw 1999, and Robinson et al. 1998). Farmed fish typically are fed 1-4% of their body weight per day.
Fish can be fed by hand, by automatic feeders, and by demand feeders. Many fish farmers like to hand feed their fish each day to assure that the fish are healthy, feeding vigorously, and exhibiting no problems. Large catfish farms often drive feed trucks with compressed air blowers to distribute (toss) feed uniformly throughout the pond.
There are a variety of automatic (timed) feeders ranging in design from belt feeders that work on wind-up springs, to electric vibrating feeders, to timed feeders that can be programmed to feed hourly and for extended periods. Demand feeders do not require electricity or batteries. They usually are suspended above fish tanks and raceways and work by allowing the fish to trigger feed release by striking a moving rod that extends into the water. Whenever a fish strikes the trigger, a small amount of feed is released into the tank. Automatic and demand feeders save time, labor and money, but at the expense of the vigilance that comes with hand feeding. Some growers use night lights and bug zappers to attract and kill flying insects and bugs to provide a supplemental source of natural food for their fish.
Feed Conversion and Efficiency Calculations:
Because feed is expensive, feed conversion ratio (FCR) or feed efficiency (FE) are important calculations for the grower. They can be used to determine if feed is being used as efficiently as possible.
FCR is calculated as the weight of the feed fed to the fish divided by the weight of fish growth. For example, if fish are fed 10 pounds of feed and then exhibit a 5 pound weight gain, the FCR is 10/ 5 = 2.0. FCRs of 1.5-2.0 are considered „good¾ growth for most species.
FE is simply the reciprocal of FCRs (1/FCR). In the example above, the FE is 5/10 = 50%. Or if fish are fed 12 pounds of feed and exhibit a 4 pound weight gain, the FE = 4/12 = 30%. FEs greater than 50% are considered „good¾ growth.
Fish are not completely efficient (FEs of 100 %, FCRs of 1.0). When fed 5 pounds of feed, fish cannot exhibit 5 pounds of growth because they must use some of the energy in feed for metabolic heat, digestive processing, respiration, nerve impulses, salt balance, swimming, and other living activities. Feed conversion ratios will vary among species, sizes and activity levels of fish, environmental parameters and the culture system used.
Feed Care and Storage
Commercial fish feed is usually purchased by large farms as bulk feed in truckloads and stored in outside bins. Smaller farms often buy prepared feed in 50-pound bags. Bag feed should be kept out of direct sunlight and as cool as possible. Vitamins, proteins, and lipids are especially heat sensitive, and can be readily denatured by high storage temperatures. High moisture stimulates mold growth and feed decomposition. Avoid unnecessary handling and damage to the feed bags which may break the pellets and create „fines¾ which may not be consumed by fish.
Feed should not be stored longer than 90 to 100 days, and should be inventoried regularly. Bags should not be stacked higher than 10 at a time. Older feed should be used first, and all feed should be regularly inspected for mold prior to feeding. All moldy feed should be discarded immediately. Mice, rats, roaches and other pests should be strictly controlled in the feed storage area, because they consume and contaminate feed and transmit diseases.
Medicated FeedsWhen fish reduce or stop feeding, it is a signal to look for problems. Off-feed behavior is the first signal of trouble such as disease or water quality deterioration in the fish growing system. Relatively few therapeutic drugs are approved for fish by FDA (see Helfrich and Smith 2001), but some medicated feeds for sick fish are available. Although using medicated feeds is one of the easiest ways to treat fish, they must be used early and quickly because sick fish frequently will stop feeding.
Managing Fish Wastes
The most important rule in fish nutrition is to avoid overfeeding. Overfeeding is a waste of expensive feed. It also results in water pollution, low dissolved oxygen levels, increased biological oxygen demand, and increased bacterial loads. Usually, fish should be fed only the amount of feed that they can consume quickly (less than 25 minutes). Many growers use floating (extruded) feeds in order to observe feeding activity and to help judge if more or less feed should be fed.
Even with careful management, some feed ends up as waste. For example, out of 100 units of feed fed to fish, typically about 10 units of feed are uneaten (wasted) and 10 units of solid and 30 units of liquid waste (50% total wastes) are produced by fish. Of the remaining feed, about 25% is used for growth and another 25% is used for metabolism (heat energy for life processes). These numbers may vary greatly with species, sizes, activity, water temperature, and other environmental conditions.
Food Intake in Fish. 2001. Houlihan, D., Bouiard, T. and Jobling, M., eds. Iowa State University Press. Blackwell Science Ltd. 418 pp.
Fish Kills: Their Causes and Prevention. 2001. Helfrich L., and S. Smith. Viginia Cooperative Extension Service Publication 420- 252. Website: http://pubs.ext.vt.edu/420-252/
Feeding Catfish in Commercial Ponds. 1998. E. Robinson, M. Li, and M. Brunson. Southern Regional Aquaculture Center, Fact Sheet # 181. Web Site: http://www.msstate.edu/dept/srac/fslist.htm
Nutrient Requirements of Fish. 1993. Committee on Animal Nutrition. National Research Council. National Academy Press. Washington D.C. 114 pp.
Nutrition and Feeding of Fish. 1989. Tom Lovell. Van Nostrand Reinhold, New York. 260 pp.
Principles of Warmwater Aquaculture. 1979. Robert R. Stickney. John Wiley and Sons, New York. 375 pp.
Standard Methods for the Nutrition and Feeding of Farmed Fish and Shrimp. 1990. Albert G.J. Tacon. Volume 1: The Essential Nutrients. Volume 2:Nutrient Sources and Composition. Volume 3: Feeding Methods. Argent Laboratories Press. Redmond, WA.
Reviewed by Michelle Davis, Research Associate, Fisheries and Wildlife
Virginia Cooperative Extension materials are available for public use, reprint, or citation without further permission, provided the use includes credit to the author and to Virginia Cooperative Extension, Virginia Tech, and Virginia State University.
Issued in furtherance of Cooperative Extension work, Virginia Polytechnic Institute and State University, Virginia State University, and the U.S. Department of Agriculture cooperating. Edwin J. Jones, Director, Virginia Cooperative Extension, Virginia Tech, Blacksburg; M. Ray McKinnie, Administrator, 1890 Extension Program, Virginia State University, Petersburg.
May 1, 2009