LARVAL NUTRITION

Nutrient requirements of all animals vary throughout their life-cycle. The changes that occur in the morphology and physiology of aquatic animals between hatching and maturity lead to a number of important variations in feeding and nutritional requirements through the larval, fingerling and adult stages. These variations occur in the morphology of the digestive organs, the digestive processes, the nutrient requirements and the feeding behaviour. Five periods in the lifespan of a fish are as follows Embryonic Larval Juvenile From fertilization to first exogenous nutrition When active swimming and feeding is taking place Beginning when the fins are fully developed, the scales have appeared and most organs have been formed and ending with the beginning of gametogenesis Beginning with the onset of gametogenesis Following the last spawning event. This stage is most clear in those salmonids which spawn only once and may not occur in some species.

Adult Senescent

The transition from an endogenous to an exogenous food supply which marks the onset of the larval stage is one of the most critical phases of the cycle and is the period when much of the mortality of hatchery reared stock occurs. MORPHOLOGY The morphology of adult digestive tracts varies markedly Between species, and Development stages, Such as morphological changes i.e. increase in mouth size and the differentiation of the digestive tract. Changes in mouth size are extremely important in considering larval nutrition as they affect the capacity of the fish to ingest food organisms. Larval fish provided with food particles that are too large will be unable to consume them and result in starvation. Whereas, if the particles are too small the energy required together sufficient quantities of food will be high and will result in poor growth. Therefore, one of the most important features of larval nutrition is correct relation of mouth size with the particle size provided to the fish. It is also important to increase the size of the food particle with changes in mouth- size as the larvae grow. The morphology of the digestive tract is generally very simple in swim-up larvae. Teeth are often absent. The gut is relatively short, about half the standard body length and the epithelial cells lining the digestive tract show no regional differentiation. The majority of

these epithelial cells are absorptive enterocytes with many microvilli on their luminal surface. Secretory cells are present in small numbers. After first exogenous feeding, a number of changes occur. Mucosal folds develop and the intestine develops regional differentiation. Absorption of nutrients in these animals is characterized by pinocytosis of macromolecules (e.g. protein and lipid) in which the surface membrane of the epithelial cells engulfs the molecule. Material-absorbed in this way is then broken down intra-cellularly. Later in larval development, teeth appear on the jaws, the stomach and pyloric caecae develop and the intestine lengthens relative to body length begins to coil. With the development of the mucosa, greater amounts of digestive enzymes are produced, which facilitate digestion within the gastrointestinal tract.

The developmental changes are related to the feeding frequency and prey size for larval animals: Prey size should match with the mouth size. In addition, the lack of teeth on the jaw and the absence of a stomach mean that the food items must be easily captured and digested independently of these mechanisms. The short relative gut length of younger fish means that a limited number of food items can be taken at one time and these items cannot be retained for a long period of time in the gut. As a result their digestion must be rapid and feeding must be done to provide sufficient nutrients. DIGESTIVE PROCESS It has been shown that the digestive mechanisms are poorly developed in larval life and diet provided must take account of this deficiency. The development of the digestive processes of larval aquatic animals corresponds to morphological complexity. Majority of the epithelial cells of the digestive tract are absorptive in swim-up larvae thus indicating limited secretion of digestive enzymes in early development. Four of the enzymes, trypsin, chymotrypsin, carboxypeptidase and pepsin, are protein-degrading enzymes, and their activities can be correlated with developmental changes in the gastro-intestinal tract associated with the change from intracellular to intra-gastrointestinal digestion of protein. Trypsin is most active at neutral pH. It increases through larval development to age 12 days, after that it decreases to day 16. However, by day 25 it again increases. Pepsin is active at low pH and present only in the stomach, increases from day 16. This increase is due to the development of the stomach and the glands that secrete the enzyme and acid to produce the low- pH environment. Chymotrypsin shows no great variation in activity over development and Carboxypeptidase shows a decline to day 16, but like trypsin carboxypeptidase activities increase at day 25. Amylase activity also remains largely unchanged through larval development.

In a study of the digestive processes in a stomach less fish, it was found that relative gut length, gut passage rate, trypsin activity and the ability to reabsorb digestive enzymes in the hind gut increased with age from larvae to adults. FEEDING BEHAVIOUR AND ITS ROLE IN LARVAL NUTRITION Despite the clear importance of nutrition in larval survival, growth and development, relatively little is known about the absolute nutrient requirements of the various larval aquatic animals due to number of factors such as Difficulty in working with small animals Sensitivity to handling, which make measuring and weighing larvae problematic. Difficulty experienced in inducing larvae to take artificial diets

Some examples of species whose larvae are not known to take an artificial diet include turbot (Scophthalmus. maximus), milkfish (Chanos chanos), striped bass (Marone spp.), black bass (Micropterus spp.), barramundi (Lates calcarifer), Murray cod (Maccullochella peelii), golden perch (Macquaria ambigua) and silver perch (Bidyanus bidyallus). Of other cultured species that will take an artificial diet, it is necessary to supplement such diets with live zooplankton. Species in this group include yellow tail (Senoia quinqueradiata), gilthead sea bream (Sparus aurata) and puffer fish (Fugu rubripes). In the case of puffer fish a complete diet of live food is not necessarily optimal, however, provision of microparticulate diets in addition to live food enhances both growth rate survival rate. The species that require live food during their larval stage are induced to feed by the movement of the prey. The eyes develop early in aquatic animal larvae, and thus larvae can capture the prey. This capacity varies between species, and some fish, such as the sturgeon (Acipenser sp.) are capable of catching zooplankton without any light. In most cyprinid species such as Chinese carp, feeding in larvae is known to be by passive filtration. ** Development of artificial diets for larvae is extremely difficult process as live feeds provide full nutrition to the larval stages. NUTRIENT REQUIREMENT OF LARVAE Larval fish require diets having a higher protein and fatty acid content. Therefore, the nutrient quality of live foods is increased by enrichment of these foods for the larvae. The live foods that have been most-intensely used for larval rearing are brine shrimp and rotifers. In brine shrimp, Artemia, the main factor affecting its quality is its content of the essential fatty acid i.e. eicosapentaenoic acid, C20:5 (n-3) or EPA and Docosahexaenoic acid, C22:6 (n-3) or DHA. These fatty acids can be incorporated into brine shrimp nauplii prior to offering it as prey for fish larvae. This bioencapsulation is known as Artemia enrichment or boosting. Increasing the essential fatty acid content of live foods which are then fed to larvae has been shown to be beneficial for fish larvae.

It is generally considered that EPA and DHA are important fatty acids in the nutrition of larval fish, the particular fatty acid required varies between species. The problems associated with nutrient quality of live feeds do not occur when rearing of larvae is done in outdoor ponds, where planktons are being grown with the fertilization as is carried out in semi-intensive culture of carps. In such conditions, availability of number of suitable natural food organisms enables the cultured organism to obtain all its nutrient requirements without supplementation. Studies have shown that fish larvae are susceptible to dietary deficiency in greater extent than juvenile or adult fish. Thus, in the formulation of dry compound diets for larval rearing more account should be taken of zooplankton inclusion.

Major portion of larval feed development should be involved with more precise control of the following nine factors 1. nutritional balance of the formulation 2. retention of nutritional components 3. homogeneity of particles 4. particle size and distribution 5. density of particles 6. water stability 7. water solubility 8. storage stability and 9. packaging requirements. It is clear that the majority of these parameters is physical in nature and shows the importance of matching the format of the feed to the morphology in larval fish nutrition.