Metabolism (M) can be subdivided to accountfor the energy losses representing different physiological processes ; maintaining basic bodily

functions :activity and the digestion ,absorption and processing of food. It is also usually to differentiate between energy retained as somatic (body) growth and energy chenneled into the production of gametes(reproductive growth). Experimental studies are aimed at quantifying the different components of the energy budget equation to present as complete a picture as possible of the physiological transformations and pathways of energy partitioning occuring within the fish. In the studies of fish energetics it is usual to express these energy transformations in terms of physiological rates, but other forms of expression have also been used. It si impiortant that all the component of the energy budget be expressed in the same units, with SI energy units being considered the best choice. The majority of researchers endeavour to provide information in trems of joules or kilojoules(kJ). But the caloric energy units are still quite frequently used(1 cal = 4.184J). Th e energy content of a sample of food, feaces of fish tissue can be determined directly using bomb calorimetry. An alternaltive, approach is to undertake a chemical analysis of the samples with respect to protein,lipid, and carbohydrate. And thereafter calculated the energy content using series af conversions factors. The conversion factors most commonly employed are 24kj, 38kj, 17 kj for protein, lipid, and carbohydrate respectively. Thus in bioenergetic terms, the rates of a physiological process will be expressed in SI units of power 1Js = 1Watt The metabolic energy(heat) losses of fish are difficult to measure using direct methods and it is therefore more usual to use determinations of oxygen consumption as an indirect measure of energy metabolism. Metabolic rates in terms of energy units can be estimnated from rates of oxygen consumption using conversion factors. The conversion factor to be used will depent upon the type of metabolic substrate(protein, lipid, and carbohydrate). For fish metabolize primarily protein and lipid, a conversion factor of 19.4 kJ O2 (or 13.6 kJo2) has frequently been used. In very few studies have all the components of the energy budget been meansured. And it is usual to find that one or more of the major components has been estimated by difference(substraction). Since they form a balnaced equation. Dterminations of any four of the five main components- ingestion. Faecal loss, excrection, metabolism and production allow the fifth to be estimated by difference. Further problems may arise because it is not possible to determine all the components simultaneously. For example, production is ideally studied as the

change in energy content in the fish body with time , but since the energy content can only be determited after the fish has been slaughtered , it is not possible to obtain information about the energy content of a particular fish both before the start, and on completion, of an experiment. In this instance the researher must rely on information obtained from sampling different individuals and must thereforetake care to ensure that the fish sampled before the start of the experiment are as similar as possible to those used in the experiment with respect to generic background. Nutritional history and physiological status. Whilst there are considerable advantages to be gained by expressing the components of the budget in terms of common energy units, it is also possible to find budgets constructed solely in terms of cahnges in biomass (wet or dry weight). Weight, rather than energy gain is particularly widely used for the expression of the results of growth studies. In order for weight gain, as an expression of growth, to be a realistic reflection of energy deposition. It si essential that the relative composition of the fish tissue remains unaltered under differnt growth condition. This prerequisite is rerely fulfilled and there may be large variations in the composition of fish tissues. Both with season and with age. The precentage liipid content of fish usually increases with increasing fish size , so that the energy density of large fish is often greater than that of small fish of the same species. In addition, fish that are feeding well, and growing conspecifics. Thus if growth is expressed solely in terms of weight gain. And there is a failure to recognize that there may be significant changes in body composition during the course of a growth study, incorrect consclusion may be drawn about the energy utilization and partitioning by the fish. `FACTORS INFLUENCING INGESTION (R) When conduction laboratory studies on fish energetics and growth. The fish objective will usually be to abtain information abaout maximum growth rates of the fish under different environmental , and thereafter conduct additional experiments in wicth optima can be determined. Fish will therefore often be fed according to regimes that aim to ensure that rates of ingestion are maximized, and there are a number of feeding options open to the experimenter. Feeding to excess can be employed both when natural prey and when formulated pellets are used as food. Pelleted feeds can be provided from automatic feeder timed to distributed portions of food at short intervals. This ensures that food is avialable more or less continously and therefore gives the fish opportunity to

consume maximum rations and grow maximally under the given set of experimental conditions. The disadventages of this method are that it is wasteful of food and unless care in taken. The excess food may disintegrate and lead to a deterioration of water quality in the experimental tanks. Furthermore , allthough the fish are feeding maximally , it si impossible to determine a value for this maximum rate of feeding without applying special techniques. Talbot and Higgins describe d a radiographic method for the quantitative determination of the gut content fo fish. And application of this method allows estimations of food intake to be made on groups of fish that have been fed in excess. The method has the advantages that fish need not be deprived of food prior to the start of a measurement period, disturbance of the fish can be kept to a minimum during feeding and the method is non-invasive so repeated measurements may be made on individual fish. Thus, this X-radiographic method can be a valuable tool for the study of food intake and feeding behaviour of fish exposed to a variety of conditions. Satiation feeding is defined as the maximum amount of food a fish will consume, and can therefore only be achieved by having food continously available (i.e. under conditions of excess feeding). Thus, satiation feeding as practiced in laboratory growth and energetics studies usually represents a compromise, and the term has been used to described feeding regimes employed in a wide range of experimental protocols. Most usually, satiation feeding has come to mean the maximum amount of food a fish will consume when presented with food two or three times per day. It is questionable how accuratelythis approximates to true satiation feeding, but it is suggested that under the experimental conditions most often employed with salmonids ( ) the estimate obtained is close to maximum ingestion. Evidence for this is given by Grayton and Beamish(1977) who studied food intake and growth of small (10 kg) rainbow trout(Oncorhyncus mykiss) fed at different frequencies at 10oC . Feeding frequency had no significant effects on food intake and growth at frequencies above two times per day. Additional information is given by elliot (1975), who in a comprehensive study, examined the effect of feeding frequency, fish size and temperature on foofd intake in brown trout, Salmo trutta. There was no significant difference between intermeal times of various sizes of fish, but increasing the temperature brought about an increase in the number of meals consumed per day (meal frequency). The result suggest that two or three feeding per day should be adequate to ensure the consumption of maximum rations under normal rearing conditions. Even though the presentation of food to the fish two or three times per day should be sufficient

To ensure maximum rates of ingestion, the experimenter will be the problem of deciding when to present foofd in order to achieve maximum feeding. Fish may not show equal propensity to feed all times of the day. And when allowed to feed voluntarily , fish of many species display peak and troughs of feeding activity during the course af a 24 cycle: some species are most active at night, others during the day, and several predatory fish species feed most actively in the periods around dusk and dawn. Furthermore, the time of day at which the fish feed most avidly may also changewith season. For example, during spring autumn both rainbown trout and Arctic Charr, Salvelinus alpinus , show peaks of feeding activity around dusk and dawn, but during winter the majority of food ingestion occurs during the hours of darkness. In summer, feeding activity is greatest during the daylight hours(Landless, 1976, jorgensen and Jobling 1989). By contrast, juvenil Atlantic salmon, Salmon salar, appear to feed almost exclusively during the hours of daylight, irrespective of season( ) Restricted feeding if the aim of an experiment is the investigation of growthrations relationship (Fig.1.1) fish will have to be fed on a range of rations varying from zero(starvation) up to maximum. In other words, various

Fig. 1.1 Influence of rates of ingestion on growth rates in fish species. Rmaint(maintenance ration) is the ingestion rate required for fish to maintain body weight:Ropt(optimum ration) is the ingestion rate at which gross conversion effeciency(weight gain per unit food intake) is maximized;Rmax is maximum ration. Horizontal dashed line indicates the growth rate Gmax to be expected at Rmax
Restricted feeding regimes will have to be employed . This will usually entail the feeding af a fixed amount of food to the fish at some predetermined time of the day. Within medical and agricultural sciences, the effects of different feeding and rearing practices on a number of physiological variables have been investigated, but there is comparatively little information avialable for fish species. There is, however evidence that both graowth and fat deposition may be influenced by the time of day at which food is provided to the fish ( ). Thus by arbitrarily adopting a given feeding rountine (feeding once or twice per day at fixed time) the experimenter may inadvertenly impose a growth restriction on the fish, whilst at the same time influencing patterns of fat deposition. When fish are held together of groups, interaction between individuals will often lead to the establishment of sosial (dominance) hierarchies. Once established, a

heirarchy can remain stable for an extended period of time, with the social rank of the dominant fish being reinforced by performanceof threat displays of overt acts of aggression(chase and bite) againts subordinate individuals. Once consequence of the social heirarchy is that resources(e.g. space, feeding site and food supplies ) are not equally devided among all members of the group and it sis usually be dominant individuals that secure first acces to any limited resourse. Thus ,when food supply is restricted , the food will nnot be eqully apportioned, and this may lead to large differences in food consumption and growth among individuals fish within the group. Acts of aggression may increasein frequency during feeding, and severe restriction af food supply will also, generally, lead to increases in agonistic interactions between the fish has they complete for the limite resource(Magnuson, 1962). When the behavioural interactions between fish have been observed, it has usually been found that the largest fish within the group are the dominants, and this has given rise to the concept of the size-related dominance heirarchy. Aviablable evidence suggests that it is the competitively superior fish whish become dominant, and, in ,turn, grow to being the largest individuals, either as a consequance of securing a major proportion of the food supplied or by suppression of the growth of subordinate individuals(A ) .Whatever the causes and the effects leading to the formationof dominancesubordinate relationships, it is clear that the establishment of social heirarchies may influence the result obtained in a feeding trial, and the effect of heirarchy formation are likely to become increasingly important with decreasing food supply. Influence of body size on Ingestion The majority of biological traits, including physiological rates, are size dipendent. The relationship between a biological variable and body weigght can usually be described by a equation of the form f(W)=aWb and b is usually <1 so that the biological variable or function [f(W)] increases allometrically