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The effect of cycle period, ration level and repetitive cycling on the compensatory growth response in… Quinton, John Chadwick 1989

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T H E EFFECT OF C Y C L E PERIOD, RATION L E V E L A N D REPETITIVE CYCLING ON T H E COMPENSATORY G R O W T H RESPONSE IN RAINBOW TROUT, Salmo gairdneri Richardson By JOHN CHADWICK QUINTON B.Sc, The University of British Columbia, 1981 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF T H E REQUIREMENTS FOR T H E D E G R E E OF MASTER OF SCIENCE in T H E F A C U L T Y OF G R A D U A T E STUDIES D E P A R T M E N T OF Z O O L O G Y We accept this thesis as conforming to the required standard T H E UNTVERSITY OF BRITISH COLUMBIA May 1989 © John Chadwick Quinton, 1989 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of & / o rp r The University of British Columbia Vancouver, Canada DE-6 (2/88) Abstract The Effect of Cycle Period, Ration Level and Repetitive Cycling on the Compensatory Growth Response in Rainbow Trout, Salmo gairdneri Richardson Compensatory growth is the phase of rapid growth, greater than normal or control growth, which occurs upon adequate refeeding following a period of undernutrition. The effect of cycle period (length of the starvation and following refeeding periods), ration level and repetitive cycling (repetition of cycle periods) on the compensatory growth response in rainbow trout, Salmo gairdneri Richardson were evaluated in two experiments. A cycle period of three weeks produced better results in terms of average percentage changes in weight and length and in specific growth rate than either one or two week cycle periods. There was no significant difference between the cyclically fed fish and a constantly fed control group. Three ration levels were compared using a three week cycle period and the only effect of increased ration was to decrease conversion efficiency. There were no significant differences in the average weight of control and experimental groups after six or twelve weeks of continuous cycling thought the controls had been fed more than twice as much food. Carcass analysis of moisture, fat, protein and ash showed no significant differences between the controls and experimental group after one complete cycle. Possible mechanisms underlying the compensatory growth response are discussed. ii Table of Contents Abstract ii Table of Contents iii List of Tables iv List of Figures v Acknowedgements vi Introduction 1 Early References 1 Recent Agricultural Work 3 Metabolic Energetics and the Compensatory Growth 10 Response in Fish Materials and Methods 23 Results 30 Discussion 57 Conclusions 64 References 66 Appendix 72 iii List of Tables Table I: Group Treatments for Experiments 1 and 2 27 Table II: Summary of Results of Experiment 1 31 Table HI: Summary of Results of Experiment 2 43 Table IV: Carcass Composition: Average Values (%) 47 Table V : Carcass Composition Changes in a 100 g Fish 56 iv List of Figures Fig. 1 Experimental Facilities at South Campus 25 Fig. 2 Group 1A 32 Fig. 3 Group IB 33 Fig. 4 Group 1C 34 Fig. 5 Group ID 35 Fig. 6 Group IE 36 Fig. 7 Group IF 37 Fig. 8 Average % Change in Weight, Experiment 1 38 Fig. 9 Average % Change in Length, Experiment 1 39 Fig. 10 Specific Growth Rate, Experiment 1 40 Fig. 11 Conversion Efficiency, Experiment 1 41 Fig. 12 Average Weight, Experiment 2 44 Fig. 13 Total Feed, Experiment 2 45 Fig. 14 Carcass Composition, Moisture 48 Fig. 15 Carcass Composition, Fat 49 Fig. 16 Carcass Composition, Protein 50 Fig. 17 Carcass Composition, Ash 51 Fig. 18 Carcass Composition, Dry Fat 52 Fig. 19 Carcass Composition, Dry Protein 53 Fig. 20 Carcass Composition, Dry Ash 54 Fig. 21 Average % Change in Carcass Composition 55 Fig. 22 Average % Change in Weight, Experiment 2 61 v Acknowledgements I would like to thank my supervisor, Dr. Robert W. Blake and Dr G.E. Scudder and the Department of Zoology of the University of British Columbia for financial support. vi Introduction The purpose of this study is to determine the effect of cycle period, ration level and repetitive cycling on the compensatory growth response in rainbow trout, Salmo gairdneri Richardson. Compensatory growth is a phase of rapid growth, greater than normal or control growth rates associated with adequate refeeding of animals following a period of weight loss caused by undernutrition (Dobson and Holmes, 1984). Cycle period is the length of the starvation and following refeeding periods. A three week cycle period would be three weeks of starvation followed by three weeks of feeding. The ration level is the amount of feed fed to the fish per day. This is calculated as a percentage of the total body weight of all fish in an experimental group and is expressed in percent body weight per day. Repetitive cycling means the cycles of starvation and refeeding are repeated, alternating periods of feeding with periods of starvation. Early References Compensatory growth has been observed in agricultural animals since the turn of the century. Waters (1908, 1909) showed that undernourished beef steers, if adequately fed, could recover and reach normal mature size and weight. He felt this was an essential trait for any animal subject to periods of undernutrition or starvation. Osbourne and Mendel (1915a,b) found that rats, when given unrestricted food, could recover and reach normal mature size after being held at a constant weight for up to 500 days 1 with restricted diet. Work with dairy heifers (Swett and Eckles, 1918) showed that there was a general tendency for animals to recover from periods of undernutrition but growth could be permanently stunted if the restriction was too severe. This permanent stunting was also found in rats held at constant weight by dietary restriction for 1000 days (McCay et al, 1939). Brody (1927) suggested that the increased rate of growth following a period of restriction was proportional to that required to achieve normal adult size. This idea was expressed by Bohman (1955) when he defined compensatory growth as abnormally rapid growth relative to age. This long term compensatory growth has been shown to occur in human children (Sternes and Moore, 1931). Children given a balanced diet after a period of malnutrition showed up to a nine times normal increase in weight and a four times normal increase in height in the first three months. Growth rate can also be affected by other stresses. Crichtion and Aitken (1954) showed that the decline in growth rate which occurs when heifers are mated too early is completely made up during the following lactations if the cattle are adequately fed. Palsson (1955) noted that generally any part, organ or tissue of an ariimal whose growth has been retarded by nutritional restriction can recover if the restriction has not been too severe. This long term type of compensatory growth response is clearly shown in the negative correlation between winter and summer weight gains of pastured animals. Those animals which lost the most weight over the winter gained the most in the spring and summer when feed became plentiful (Black et al, 1940, Pearson-Hughs et al, 1955). In their review of compensatory growth Wilson and Osbourn 2 (1960) state that compensatory growth, after a period of undernutrition, is a constant feature among higher animals. Their conclusions are summarized below: i) The growth rate following period of undernutrition is usually enhanced. ii) Too severe restriction can cause permanent stunting, iii) Five main factors influence the extent of compensatory growth: a) nature of restricted diet. b) degree of severity of restriction. c) length of restriction period. d) relative rate of maturity of the species. e) the pattern of refeeding. iv) Recovery from protein and/or carbohydrate restriction is usually complete. v) The rate of compensatory growth immediately following refeeding increases with severity of restriction, vi) The pattern of refeeding may effect the carcass composition, vii) Recovery may occur either by prolonging the time to reach mature weight or by increasing growth rates during refeeding, especially when refeeding has just begun. Recent Agricultural Work There has been a great deal of interest in the compensatory growth response of sheep and cattle. Recent work has focused on the changes in body composition which accompany compensatory growth and attempts to describe some of its underlying mechanisms. Meyer and Clawson's 1964 study looked at undernutrition and 3 refeeding in both rats and sheep. They found that the maintenance ration was about 52% of what was eaten ad libitum in both animals. Any less resulted in weight loss, which was similar in proportion to weight gains when fed above this level The alimentary tract decreased in size during starvation, contrary to finding by Wilson and Osbourn (1960). Compensatory growth occurred in both rats and sheep. The total energy content and protein levels of the compensating rats, when given equal feed, were less than that of the controls. There was no significant difference in the total energy content between the refed and control sheep but the protein levels was lower in refed sheep than in the controls. The weight gain of the compensating animals was more fat and less protein than for the animals fed ad libitum in both sheep and rats. There was no depression of the metabolic rate during undernutrition or refeeding, nor was there any increase in appetite in either sheep or rats. Increased efficiency of food utilization above maintenance was largely responsible for the compensatory growth in both species. The study of mature sheep body composition and efficiency during loss and regain of live weight by Keenan et al (1969) gave different results. During weight loss the tissue was inefficiently mobilized. A 16% loss of weight resulted in a 30% loss of total body energy. The sheep were maintained at the reduced weight for eight weeks and then fed ad libitum for five weeks. Only 75% of the energy deficit was recovered. The regained tissue had a high water and low fat content compared to the continuously grown sheep. This contrasts with the results of Meyers and Clawson (1964) who found that the refed sheep has a greater fat content 4 than their controls. Walker and Garrett (1970) subjected male rats to prolonged undernutrition and examined the effects of refeeding. The energy intake required for maintenance decreased as the duration of food restriction increased. This reduced maintenance level continued into the feeding period. During both restriction and refeeding there was an increase in the efficiency of utilization of energy. As the refeeding period continued the increased efficiency of utilization of energy declined to the level of the controls. McMannus et al (1972) examined compensatory growth in five to six month old sheep. Uninterrupted growth for 58 days (36.2 % gain) was compared to undernutrition for 27 days (21.7% loss) followed by refeeding for 52 days (62.2% gain). During restriction the sheep used the feed more efficiendy that the controls. During refeeding they were less efficient than the controls. The compensating animals drank more water, ingested more food per unit body weight, laid down less fat and more protein and retained more water. There was no significant difference in the THS output from the thyroid between the compensating sheep and the controls. Compensating sheep had significantly lower plasma somatotrophin potency per unit body weight than the underfed sheep and no A C T H activity was detected. With severe undernutrition the anterior pituitary gland decreased in size but not cell number and still elaborated somatotrophin. Decrease in body size with underfeeding resulted in an increase in the ratio of circulating somatotrophin per unit body size. During compensatory growth there was hypertrophy of the anterior pituitary gland and evidence of enhanced synthesizing capacity. 5 Little and Sandland (1975) studied the distribution of body fat in sheep during continuous growth and after feed restriction and refeeding. Restricted animals had the same proportion of fat per unit wool-free empty body weight as the continuously grown animals. Refeeding sheep accumulated less fat and more protein and water than the continuously grown sheep. There was a relatively greater loss of fat from subcutaneous deposits than from the body. Deposition of fat on the skeleton continued during restriction. Weaner sheep (Graham and Searle, 1975) had greater voluntary food intake during refeeding. The basal metabolic rate was reduced during weight stasis. The suppressed metabolic rate rose during the first month of recovery but to levels less than that of the controls. In the first week of recovery the net energetic efficiency was higher while the maintenance requirements were lower. The gross efficiency was higher as intake was high relative to maintenance. Nitrogen utilization was found to be more efficient in the first two weeks of compensatory growth. The body composition of the weaner sheep was then examined (Searle and Graham, 1975). After weight stasis the sheep had less protein, more water and equal fat composition compared to the constantly fed controls. With partial and complete recovery the body composition was the same as the controls. In immature sheep (Drew and Reid, 1975 a,b,c) underfeeding to and empty body weight (E.B.W.) loss of 25% generally produced changes in body composition similar to reversal of normal growth. The level of body fat however did not decrease during the first half of the restriction period and did not increase for the first two weeks of refeeding. Refed sheep at 45 kg E.B.W. contained more 6 protein and water and less fat than the continuously fed sheep. These effects were greater in the carcass and so the carcasses of the refed sheep were heavier with less fat and more lean than those continuously fed. Sheep fed at 70% ad libitum produced carcasses with more protein than those fed ad libitum for either continuously fed and refed sheep. The reduction of bone water and accumulation of bone fat during severe underfeeding was rapid. Upon refeeding the bone fat was rapidly mobilized and bone water returned to normal. Initial weight loss was due mostly to loss of water from the bone and fat utilization. In early regrowth there was a stimulation of protein synthesis and a depression of fat synthesis. The ratio of muscle to fat gain in sheep from 30 to 40 kg E.B.W. was 2.23:1 for refed sheep and 1.08:1 for continuously grown sheep. There was a 46% increase in the rate of gain following refeeding with no increase in intake per day compared to continuously grown sheep. Much of this could be due to the rapid accumulation of water (Drew and Reid, 1975c). The total feed cost to reach 45 kg E.B.W. was 27% higher for the refed sheep than those fed ad libitum to the same E.B.W.. There was no significant difference between normal growth and refeeding in efficiency of energy retention above maintenance. Rats showed a significant difference in the compensatory growth response between young and old animals (Miller and Wise, 1976). Young refed animals were 39% more efficient and old animals were 21% more efficient than the controls in food conversion. In gross energetic efficiency the young refed animals were 29% and the old refed animals were 17% more efficient than the continuously fed animals. This was probably due to differences in 7 metabolism between the younger and older animals. The "catch up" growth was associated with an increased food uptake and metabolic adaptations that gave higher efficiencies. Miller and Wise (1976) postulated that the difference between young and old animals was due to either a higher efficiency of synthesis or lower metabolic costs for the younger animals. Thornton et al (1979) was the first work on compensatory growth in sheep to incorporate two periods of weight loss and refeeding on both immature (below 23 kg) and mature (above 43 kg) sheep. Immature sheep were depleted of fat during weight loss. The loss of fat from the meat was associated with both atrophy and hypoplasia of the subcutaneous adipose cells. The meat of mature sheep showed an increase in fat during weight loss but only atrophy with no hypoplasia of the adipose cells. This difference was probably due to the much higher initial fat content of the mature sheep. The greatest loss of fat in both the mature and immature sheep was from the meat but there was a proportionally higher loss from the offal, especially in mature sheep. The amount of protein in the carcass was similar for control, starved or refed sheep of the same body weight. During the first few days of refeeding, the food consumption of the sheep was three to four times as great as during the starvation period. The apparent digestibility coefficient of the food went from 53-68% to 80-90% and the live weight gains were 500 to 600 g per day. The refed sheep showed increased protein, water and fat in their meat. Sheep which were starved and refed, either once or twice, quickly reached the same live weights as the continuously grown animals and were similar in body and meat composition. 8 The findings regarding compensatory growth in sheep are equivocal for changes in basal metabolic rate, appetite and carcass composition. Thornton et al (1979) suggests that although the results cannot be reconciled, they possibly suggest the true variety of results which can result from separate experiments on limited numbers of animals with variable body composition under differing conditions of nutritional restriction and refeeding. The conclusions of many of these studies are that the advantages of the rapid growth following restriction is outweighed by the energy cost of maintenance during restriction (Thornton et al, 1979). This may be a problem caused by the experimental design of the studies. All of the previously examined studies used fairly long periods of weight loss and reduced weight stasis in their design. It is during this period that maintenance costs become important. If the period of weight loss and stasis was shortened, the gains made in the compensatory growth phase could outweigh the maintenance costs during the starvation stage. Even a small overall increase greater than constantly fed controls could be of economic importance. The reduced cycle time would allow for a greater number of cycles, the effect being cumulative. The success of this type of feeding would depend on the effect starvation period had on the compensatory growth response. The optimum cycle period for body weight would have to be determined. Some studies showed that body composition can be altered by the pattern of starvation and refeeding to produce increased protein and reduced fat levels in the carcass (Keenan et al, 1969, McMannus et al, 1972, Drew and Reid, 1975a,b,c, Little and Sandland, 1975). This could be manipulated to produce leaner carcasses in meat producing 9 animals. Compensatory growth is already an important economic factor of livestock production in Australia's pastoral zones (Thornton et al, 1979). Studies such as those by Bennett et al (1970) on the effects of grazing cattle and sheep together show that compensatory growth is very important to stock which overwinter on poor pasture. Animals which lost weight heavily in the winter tended to gain weight faster in the spring than those which had lost less weight. Compensatory gains were important for herd management. Supplemental feeding in winter, except for survival, is not only expensive to the farmer but may also deprive him of the benefits of compensatory gains and greater pasture use the following spring. The existence of compensatory growth in agricultural animals has been known and noted for a long time. In some areas it is of fundamental economic importance while in others, such as sheep nutrition, the understanding of its mechanisms is incomplete to the extent that it is not presently possible to incorporate it into agricultural practices. Metabolic Energetics and The Compensatory Growth Response in Fish In order to examine compensatory growth in fish, we must examine their metabolic energetics. A great deal of work has been done on the nutrition of salmonids. Two excellent reviews on energy partitioning and feeding study techniques are available, Cho et al (1982) and Jobling (1983). The utilization of dietary energy is the basis of the study of fish nutrition. The 10 metabolizable energy intake (ME) is equal to the energy retained as new tissue (RE) and the energy dissipated as heat (HE). The gross energy intake (IE) is the product of food consumption and its heat of combustion. The standard value for carbohydrates is 17.2 kJ/g, for protein is 23.4 kJ/g, for fat is 39.2 kJ/g and for ash is 0 kJ/g. The ash content of a feed can thus greatly affect the IE. The digestible energy (DE) is the energy digested and absorbed to be used as fuel. The fecal loss (FE) is the energy value of feed components which are not digested. The feces are made up of both undigested food (FE) and unreabsorbed residues of body origin (FmE). Thus apparent digestible energy = IE - F E while the corrected digestible energy is IE - (FE-FmE). The major loss of ingested gross energy has been found to be fecal energy loss. Metabolizable energy (ME) is the energy of the absorbed amino acids, fatty acids and sugars to be used less the by-products of the catabolism of the amino acids. The excretion of ammonia in the form of urea is a loss of combustible energy for the fish. The loss of ammonia through the gills (ZE) or kidney (VE) means the digestible energy of the diet is an overestimate of its fuel value to the fish. Thus : M E = I E - ( F E + V E + ZE) The loss of combustible energy in the feces depends on the digestibility of the feed components. There appears to be little interaction between the diet components that affect absorption. The loss of energy through the gills and urine depends on the level and digestibility of the protein in the diet. This is influenced by the proportion of other components in the diet, especially the level and type of fat (Cho et al, 1982). 11 The metabolizable energy is the energy available to the fish. The metabolic rate of the fish is the rate at which heat is liberated. Heat is produced by the transformation of food into tissue, tissue turnover and physical activity. The basal metabolic rate is the minimum rate of metabolic activity needed to sustain the structure and function of the fish. Any form of activity increases the metabolic rate. The heat increment of feeding is the increase in metabolic rate caused by ingestion, digestion and utilization of food. This energy is not then available for growth. Growth is only possible if the energy from food (ME) is greater than the total heat loss. The energy required for maintenance is basal metabolism (HeE), thermoregulation in homeotherms (HcE) and involuntary resting activity (HjE). The specific dynamic action (SDA) is the heat produced by the chemical work of the glands. However the SDA is often more broadly defined as the heat or energy required for digestion of food, the heat increment of feeding. The duration of the SDA depends on the quality and quantity of food and on water temperature. The cost of protein deamination for use as an energy source is a major factor contributing to the heat increment of feeding. The heat increment can be 8% to 12% of IE in fish. The heat increment is quite small when compared to the metabolic work however. The physiological basis of SDA is the post-absorptive process related to ingested food, especially protein rich food. It is mainly the metabolic work to form proteins and fats from amino acids and fatty acids, plus the formation of excretory nitrogen products. There is contradictory information regarding the biochemical to the physical/mechanical ratio of heat loss. The variety of results is 12 regarded to be due to the differences in experimental techniques and changes in activity levels (Cho et al, 1982). The effect of temperature on the fasting heat production is very marked. An increase in temperature from 3 to 18°C resulted in a doubling of the heat production of Atlantic salmon and rainbow trout (Smith et al, 1978a,b). The heat production rates for Atlantic salmon and rainbow trout increased more slowly than for either brook or lake trout as the water temperature increased. Cho and Slinger (1980) measured the heat production of rainbow trout weighing from 47 to 139 g at 7.5, 10, 15 and 20°C. The largest effect occurred between 7.5 and 10°C when heat production doubled. From 10 to 15°C there was a 50% increase in heat production and from 15 to 2 0 ° C there was no further increase. These results strongly support the finclings of other researchers discussed earlier which suggest that basal metabolic rate and maintenance costs increase with temperature. Basal metabolism has traditionally been calculated by extrapolation of activity levels back to zero. The basal metabolic rate for rainbow trout was found to be 59 to 63 kJ/kg/day at 15°C for 96 to 145g trout (Cho et al, 1975) and 54 to 139 kJ/kg/day for .85 to 57g trout (Smith et al, 1978a,b). The following equation relating the body weight to heat production for rainbow trout of 1 to 59g body weight is proposed (Smith et al, 1978a,b). Heat Production(kJ/kg/day) = 204 W ° J 5 (r=0.92) This relation between heat production and the fractional coefficient of body weight would appear to indicate that surface area rather than body weight may be the important factor in basal metabolism. 13 Growth and energy retention (RE) is made up of the metabolizable energy not dissipated as heat but retained in the body as new tissue. This retained energy may be stored as fat or protein. As fish increase in maturity a higher proportion of the energy is stored as fat (Cho et al, 1982). The relative importance of fat and protein depends on the maturity of the fish, the balance of available amino acids in the dietary protein and the amount by which the dietary energy intake exceeds the energy expended as heat. Proteins of higher biological value promote greater protein deposition (Cho et al, 1982). If there is a marginal excess of energy intake a greater proportion of it is retained as protein. As the energy excess increases the total amount of protein deposition increases but the proportion retained as fat increases at a greater rate. Increasing energy levels lead to an overall increase in both total fat and in the fat to protein ration (Cho et al, 1982). Temperature has been found to affect energy retention. An increase in temperature from 7.5 to 2 0 ° C increased energy retention from 44 to 58% of digestible energy intake (Cho et al, 1982). Watanabe et al (1979) found that the maximum protein retention and an optimum protein/fat ration was achieved with a diet of 35% protein and 15 to 20% fat. A number of other factors influence growth and conversion efficiency. The stocking density for a farmed group of fish affect the variability of the growth rate of the fish. L i and Brocksen (1977) found that the metabolic rate of rainbow trout increased with increasing density and attributed this to i) starvation, ii) increased exercise levels and iii) higher levels of excitation. The variance of routine metabolism, growth rate and consumption 14 rate increased with density, due to intraspecific competition. The dominant trout grew faster and more efficiently with a higher lipid content at all densities. At higher densities dominance gave less benefits than at lower ones. Trezeviatowski et al (1981) showed that fish production, weight gain per cubic meter of water and the feed conversion rate all increased with stock densities. High stocking levels may produce pollution problems however (Clark et al, 1985). The genetic component of growth rate and body composition is difficult to assess as trout are so sensitive to environmental factors which tend to mask genetic effects. Ayles et al (1979) suggest that the lipid content of rainbow trout can be significantly different between strains and that breeding programs are viable. They also suggest that any evaluation of a stock's performance must be done under production conditions or the results are confounded by environmental factors. Refstie (1980) found that heritability is higher for length than for weight but genetic variability is much greater for weight than length. The heritability for growth of fingerlings was low, suggesting that individual selection would not be very efficient. They suggest that a combination of family and individual selection would likely give some improvement in growth rate. Muscle growth plays an important role in compensatory growth. Three different muscle types have been histochemically identified (Gill et al, 1982, Hoyle et al, 1986): white, red and pink. White muscle constitutes the greater portion of the swimming musculature. It functions anaerobically during contraction and so has reduced myoglobin and mitochondrial content, increased 15 glycolytic enzyme content and is less vascularized (Hoyle et al, 1986). Red muscle occurs as a thin superficial layer of triangular cross section below the skin which parallels the lateral line. It is geared for aerobic metabolism with a higher myoglobin and mitochondrial content, more lipid and lipolytic enzymes and is highly vascularized (Hoyle et al, 1986). Pink muscle has intermediate properties. White myotomal muscle forms the bulk of the market portion of the fish. The growth dynamics of muscle fibre was examined by Weatherly et al (1979) for yearling rainbow trout. The fish grew faster when fed ad libitum at 12°C than at 16°C and that both groups grew faster than those on a restricted ration at 12°C. The growth of the myotomal muscle mass was characterized by an increase in mean muscle fibre diameter, though most of the bulk increase resulted from the increases in fibre number. The ratio of fibre diameter to fish length was lowest for the fastest growing trout, which indicated a greater ability to add new fibres compared to those growing more slowly. The fibre diameter range increased in trout larger than 18 cm but small fibres persisted in dimMshing numbers even in the largest fish. In slower growing fish muscle growth was more influenced by mean fibre diameter. In their 1980a work Weatherly et al examined the relationship between mosaic muscle fibres and size in rainbow trout form 2.1 to 61.3 cm fork length (FL). In trout < 5 cm all the muscle fibres were < 40 Lim in diameter. From 5 to 20 cm the fibres were all in the 0 to 39.9 nm diameter class though the range was extended. The mosaic muscle bulk increased mainly by the recruitment of new small fibres. At > 20 cm the mode of the muscle fibre diameter was in 16 the 40 to 79.9 um class. Larger fibres appeared,( > lOOum), but the subsequent overall diameter frequency distribution changed very little until 50 cm. The increase in muscle mass was partly due to increases in fibre diameter but was largely the result of the continued recruitment of small fibres. At 55cm the recruitment of new fibres ceased and increases were due to the gains in diameter of the existing fibres. This would seem to place an upper limit on fish size. In fingerling trout (Weatherly 1980b) 2.3 to 5cm no fibres were > 40am but at > 5cm fibres of > 40nm appeared, ranging up to 100 um. Trout 5 to 18 cm were dominated by fibres < 40 Jim, in the 20 to 39.9 urn class with nothing above 100 |im. This was true for fish with either fast or slow growth rates. There was a marked decrease in the number of fibres in the 0 to 19.9 um class. Differences in the condition factor, dry weight and, from inference, protein did not significantly affect the fibre diameter frequency. In trout of 18 to 20 cm the fibre diameter mode shifted to the 40 to 59.9 um class in most growth rate groups and hatchery reared trout. There were a small number of large fibres up to 120 um and the 0 to 19.9 um class was further reduced. Above 20 cm the growth by fibre recruitment decreased. Between 20 and 25 cm the increase in cross-sectional area of the muscle was due mainly to gains in fibre diameter. Fish with very rapid growth rates (12°C, ad libitum) had smaller fibre diameter to fish length ratios than slower growing fish. This indicated a greater rate of recruitment of new fibres in the faster growing fish and a potential for larger ultimate size. Comparison of the growth dynamics of rainbow trout with bluntnose minnow (Pimephales notatus Rafinesque) (Weatherly and Gill, 1984) concluded that the main mechanism of 17 myotomal growth in large fast growing fish was the input of new fibre while for small slow growing fish increase in fibre diameter was of greater relative significance. During starvation different tissues are utilized sequentially as the energy source (Denton and Yousef, 1976, Elliott, 1975, Smith, 1981, Weatherly and Gill, 1981, Black and Love, 1986) and there are differential rates of mobilization of similar substrates in different organs (Love, 1980). Red muscle is less affected by starvation than white (Loughna and Goldspink, 1984). The mean fractional rate of synthesis of red muscle is 2.5 times greater than that of white. Prolonged starvation causes a significant decrease in the rate of synthesis in both muscle types (Loughna and Goldspink, 1984). The white muscle tissue responds very quickly during starvation and its mean rate of synthesis is halved during the first week while that of the red muscle remains unchanged. After two weeks the rate of synthesis in the red muscle is also halved (Loughna and Goldspink, 1984). Significant protein utilization does not occur for seven or eight weeks in Salmo gairdneri (Denton and Yousef, 1976, Elliott, 1975, Weatherly and Gill, 1981). In prolonged starvation the degradation rate was only slightly above normal (Loughna and Goldspink, 1984). The protein synthesis and degradation rates reach relatively constant values with the degradation rates exceeding the synthesis rates (Loughna and Goldspink, 1984). The reduced synthetic rate is related to both reduced RNA concentration and activity (Loughna and Goldspink, 1984). The energy source during short term starvation in S. gairdneri is adipose fat (Weatherly and Gill, 1981) and muscle lipid (Parker 18 and Vanstone, 1966, Smith, 1981) which is proportionally replaced with water (Idler and Bitners, 1959). In Gadus morhua liver lipid, liver glycogen and white muscle glycogen are utilized (Black and Love, 1986). Full recovery from short term starvation and very high growth rates are possible (Bilton and Robins, 1973; Smith, 1981; Weatherly and Gill, 1981; Dobson and Holmes, 1984; Kinkschi, 1988). If the starvation continues there is a point past which full recovery upon refeeding does not occur and the ability to catch up to constantly fed control fish is lost. High mortality begins to occur, especially upon refeeding (Bilton and Robins, 1973; Love, 1970; Love, 1980). The fish may be unable to recover because the ability to utilize feed (Bilton and Robins, 1973) is reduced by gut atrophy (Salmo gairdneri: Weatherly and Gill, 1981) and reabsorption of the microvilli, especially in the middle section of the intestine (Cyprinus carpio: Love, 1980). Long term starvation results in decreases in the length, weight and diameter of the intestine (Love, 1980). White muscle tissue is then utilized as the energy source (Johnston, 1981, Johnston and Goldspink, 1973, Moon, 1983, Moon and Johnston, 1980). Mammalian studies have produced similar results (Swett and Eckles, 1918, McCay et al, 1939; Wilson and Osbourn, 1960; Thornton et al, 1979). The presence of compensatory growth in fish is far less well documented than for mammals. Bilton and Robbins (1973) examined the effects of starving and feeding on the survival and growth of sockeye salmon fry. The fry were capable of withstanding three to four weeks of starvation with less than a 10% mortality but many 19 were incapable of recovery. Beyond four weeks of starvation mortality increased sharply to 90% at seven weeks. The pattern of mortality was similar in all experimental groups with a sharp increase after 30 days. The mortality continued when the fish were offered food. The length and weight of the fry starved up to seven weeks decreased significantly. The decrease in length may have been due to reabsorption of cartilaginous material from the skeletal system. There was an accelerated growth rate among some groups of fish which survived to the end of the eight week feeding period following starvation. Those fish which were starved from one to three weeks caught up in length and weight to the control group when fed. It appeared that these fry utilized feed more efficiently. Survivors of four weeks starvation and eight weeks feeding did not catch up to the controls in either length or weight. Starvation of up to three weeks did not prevent the sockeye fry from reaching the size of others in the population which had not been starved. Prolonged starvation, longer than three weeks, inhibited the fry's ability to utilize the feed when offered and resulted in permanent stunting or death. Weatherly and Gill (1981) compared the starvation response and subsequent recovery of fingerling rainbow trout (Salmo gairdneri Richardson) on restricted rations for 16 weeks, starved for 3 weeks (14.5% weight loss) and starved for 16 weeks (32.5% weight loss). The visceral fat was completely utilized in both the long and short term starvation groups. The gut was significantly reduced in the long term group. Subsequent recovery at full rations produced growth rates that were approximately equal to those of the controls with respect to wet body weight and 20 condition factor. The recovery fish surpassed the controls in percent dry weight, heart, liver, gonad and gut and visceral fat weight. This indicated an overcompensative response. The gut, skin and dry carcass weight were less in the controls and in the three week starved group than in the reduced ration and severely starved group. This indicated that the slow growth from limited rations resembled severe starvation rather than short term starvation. Dobson and Holmes (1984) examined the effects of starvation and feeding in farmed rainbow trout (Salmo gairdneri Richardson). Fish were divided into three groups: group A was fed for three weeks then starved for three weeks; group B was starved for three weeks then fed for three weeks; and group C, the control, was fed constantly for the six week period. The fish were all fed Omega pelleted trout food at the manufacturer's recommended level of 5% of body weight per day. The experiment was repeated five times. The fish gained weight when fed and lost weight when starved. Comparisons of subgroups with controls showed that in four of the five periods the total percentage weight gain of subgroup B (starved then fed) was equal or greater than the control, group C. Thus fish starved then fed for three weeks gained as much weight as fish fed throughout the six weeks of the experiment, though fed half as much feed. Comparison of weight gain prior to starvation with weight gain after starvation showed a significant increase in weight gain if feeding is preceded by a period of starvation. Comparison of overall weight gain for starvation and refeeding with the weight gain for the first three weeks of group A (feeding only) shows the mean weight gain for starvation and feeding is greater than that of feeding only for three weeks. Length changes 21 were measured and showed that starving and refeeding produced greater overall length increases than feeding and starving. This indicated that the weight gains made after the starvation period were associated with increases in length and could be considered as growth and not just gut fat deposits or water uptake. Unfortunately no data was presented comparing the length increases of the starved then fed group (B) with the controls (C). Figures for weight loss during starvation showed a reduction in the rate of weight loss over the three week period. The experiments performed here were designed to determine the effect of different short starvation and refeeding periods, feeding levels and repetitive cycles on the compensatory growth response and the carcass composition. The first experiment compares the average percentage increases in weight and length, the conversion efficiency and specific growth rate of a constantly fed control group with experimental groups. These experimental groups were starved then fed for one, two or three week cycles or were starved then fed for three weeks cycles at ration levels higher or lower than the control. The second experiment compares the average percentage increases in weight and length, conversion efficiency and specific growth rate of the constantly fed control group with the experimental group which was starved then fed for alternate three week periods. The ration level was the same for both groups. Samples from both the control and experimental groups were analyzed for moisture, fat, protein, and ash composition at the start, at three weeks and at six weeks in order to determine if the compensatory growth response altered carcass composition. 22 Materials and Methods Rainbow trout, Salmo gairdneri Richardson were purchased from the Sun Valley Trout Farm of Mission, British Columbia and delivered to the Animal Care Centre of the University of British Columbia. The layout of the experimental facilities is shown in Fig. 1. The experiments were performed in circular fibreglass tanks eight feet in diameter and four feet in depth with a central standpipe to control water level and a standpipe sleeve to improve water circulation and flushing. Water was supplied from the general service of the University, mixed with hot water to maintain a constant temperature of 12 to 14°C and run through two large activated charcoal filters (Triton model TR-140, capacity 140 gallons per minute) to remove particulate matter and chlorine. A thiosulfate injection system was used (Mec-o-matic Powermatic II continuous injection pump) to further reduce chlorine levels. The water was supplied to the tanks through aerator bars mounted on the sides of the tanks to insure normal dissolved oxygen levels. For the first experiment the fish (length: mean: 13.27 cm, range: 9.5 to 19.1 cm, weight: mean: 36.24 g, range: 10 to 110 g) were acclimated for a two week period and then divided into experimental groups as follows. Fish were netted from the holding tank and placed in a temperature controlled anesthetic tank containing 2-phenoxy-ethanol (0.4 ml per litre, Syndel Laboratories, Vancouver, B.C.). Once anesthetized, the fish were removed from the anesthetic, weighed (Mettler balance, P1200 ± 23 Figure 1: Legend A: Hot Water B: Cold Water C: Air Compressor D: Carbon Filter E : Thiosulphate Injection System F: Air Line G: Water Line H : Experimental Tanks I: Water Supply Valves J: Air Supply Valves K: Tank Drains L : Gutter Scale: 1/4" = 1.0' 24 25 0.01 gm.), measured for standard length using calipers which were then compared to a steel rule (± 0.5 cm), and tagged with individually numbered fingerling tags (Floy Tag Co., Seattle, Washington, U .S .A.) sutured through the dorsal musculature just posterior to the dorsal fin. Following measurement and recovery from the anesthetic the fish were placed in one of six experimental tanks. There were a total of 40 fish in each experimental group. The group treatments are summarized in Table I. Groups were sampled by netting 10 fish out of each tank, anesthetizing and measuring them and then returning them to their experimental groups. Al l groups were sampled once per week for the six week duration of the experiment (Appendix). Experiment 2 fish (length: mean: 18.98cm, range: 13.3 to 22.5 cm, weight: mean: 120.22 g, range:42 to 189 g) were treated as per those in experiment 1. Shortly after arrival the fish were taken out of the holding tank, anesthetized, measured for standard length and weight and tagged with the individually numbered fingerling tags. They were then divided into two groups of 50 fish each. The treatments of the two groups are summarized in Table I. The length and weight of five fish from the holding tank were recorded, then these fish were killed and immediately frozen and stored in the freezer (Bel-Par Industries, -20^ C). Both groups were sampled at three week intervals, tag number, weight and length were recorded for each fish in the sample. After three weeks 15 fish were sampled from each tank , five of which were 26 Table I: Group Treatments for Experiments 1 and 2 Group Cycle Period 1A Constant IB 1 Week 1C 2 Weeks ID 3 Weeks IE 3 Weeks IF 3 Weeks 2A Constant 2B 3 Weeks Ration Level 5% 5% 5% 3% 5% 7% 5% 5% 27 killed, frozen and stored. After six weeks 30 fish from each tank were sampled, five of which were killed, frozen and stored. The percentage of moisture, fat, protein, and ash were assessed for the 25 frozen samples (General Testing Laboratories, Vancouver, B.C.; Official Methods of Analysis of the Association of Official Analytical Chemists, Tests # 24.003 (moisture), 24.005 (fat), 24.009 (ash) and 24.027 (protein as nitrogen)). The groups were sampled again after 9, 12, 15 and 18 weeks (Appendix). All groups of fish were fed once per day when fed. Moore Clarke Extruded New Age Salmon Feed of appropriate size was the feed used. The tag number, weight and length of each fish sampled in both experiments were recorded and the average percentage changes in weight, length and condition factor were calculated as : Average % Weight Change = ^ 1 - ^ ^ — i x 100 j -s- n Average % Length Change = I | - L - x lOoJ + n r ( W / L ^ M W . / L 3 ) . Average % Condit ion Factor = Z * * — * 100 + n t ( W . / L f ) > (Black and Love, 1986) Where W is sample weight, W- is initial weight, L c is sample o I S length, L^ is initial length and n is number of individual samples. Conversion efficiency (C.E.), specific growth rate (S.G.R.) and percentage average change in body composition were also calculated. 28 Conversion E f f i c i e n c y = Z F S p e c i f i c Growth Rate (%/ d a y) = I C . / n - i C / n % Average Change in Body C o m p o s i t i o n = x 100 where F is food fed per day, is the final weight, is the initial weight, T is the time in days, C^ is the final composition and C i s the initial composition. Non parametric statistical analysis is used to examine the average percentage change data (Kruskal-Wallis single factor analysis of variance by ranks, Zar, 1984). 29 Results The results for experiment 1 are summarized in Table II. The individual measurements of all the samples are found in appendix 1 as are the calculations for percentage change in weight and length. The changes in average percentage weight and length for groups 1A through IF are shown in Figs. 2 through 7. The results for group E, those starved for three weeks then fed for three weeks were anomolous (Fig.6) due to a mechanical breakdown which resulted in an interruption in the water supply to the tank. The treatment of group 2B for the first six weeks of the second experiment was identical to that for group IE and are substituted into the results for experiment 1. The average percentage change in weight for each of the groups is shown in Fig. 8 A Kruskal-Wallis analysis of variance showed no significant differences in the average percentage change in weight (q=11.0346, cc(75 d.f.) > 0.05) (Fig. 8), length (q = 10.0122, a(75 d.f.) > 0.074) (Fig. 9) and specific growth rate (q = 8.78718, a(75 d.f.) > 0.11)(Fig. 10) though the average values for ID, IE and IF are greater than that of the controls while those of IB and 1C are less (Table II). The greatest differences between the groups are seen in the conversion efficiency (Table 2, Fig. 11). All cyclicly fed group values except IB were higher than that of the controls. The trend in groups ID, IE and IF was that increased ration level did not increase percentage change in weight (Fig. 8) or length (Fig. 9) but decreased conversion efficiency (Fig. 11). The pattern of weight loss and gain during starvation and refeeding is shown for group ID in Fig. 5. There is a significant difference in 30 Table U: Summary of Results of Experiment 1 Group % change weight 1A 18.68 IB 7.99 IC 7.61 ID 27.24 IE 27.91 IF 24.91 % change % change length cond. fact. 3.69 2.92 3.50 -4.26 3.65 -3.66 7.36 2.46 6.59 4.58 6.04 8.42 conver. spec, gr e f f i c . rate 0.074 0.407 0.069 0.183 0.108 0.175 0.527 0.571 0.299 0.583 0.188 0.527 31 4) CJl c o £ o K o> cn o i _ 4) 3 F i g u r e 2 Group 1A 14 21 28 35 (771 Weight Time (Doys) IXXI Length 8 F i g u r e 3 Group 1B 1 7 -6 -5 -4 -3 -2 -1 -0 •4 ~ -5 -•6 0.5 1 I I r 0 1.5 I 2 -nr-2.5 3 3.5 ~ r 4 4.5 5 ~~r~ 5.5 6 Time (Weeks) IZ71 Weight (XXI Length A v e r a g e % C h a n g e I I I I I I C T ) t r i > O J r O - ' 0 - » r O O J - N t r i 0 1 > J C O o cn f I D 3 a ? o ST 3 I D J L i i I I I I L A 3 ro -ro 04 -cn 3 m m w w w \ s °> cn cn -cn cn ZZ1 34 Average % Change Average % Change 37 F i g u r e 8 Average % Change in Weight 1A IB 1C 1D 1E 1F Group F i g u r e 1 0 Specific Growth Rote 0.6 - i a. K 1A IB IC 10 1E 1F Group F i g u r e 11 Conversion Efficiency 0.6 -i 1A 1B 1C 1D 1E 1F Group the average percentage weight change with time (q = 30.1266, tx(45 d.f.)<0.05). The large weight loss during the first week of starvation is typical of all experimental groups. The weight loss continues during the starvation period but at a reduced rate. In the first week of refeeding there is a moderate increase in average percentage weight gain of approximately eight percent. This gain is of the same order as that found in all experimental groups (Figs. 3,4,5,7). The gain during the second week is also approximately eight percent, typical of groups 1C, ID, and IF (Figs. 4,5,7). It is during the third week of refeeding (groups ID, IE, IF, Figs. 5,7) that the greatest average percentage increases in weight and specific growth rate occur. Over this one week period the conversion efficiency for group D (Fig 5) was 1.289 and the specific growth rate was 4.04 percent per day. The results of the second experiment are shown in Table HI. The average weights of the control (2A) and experimental (2B) groups are shown in Fig. 12. There is no significance difference in the means initially (t = 0.975837, a (100 d.f.) >0.33), after three weeks (t = 1.89186, a (29 d.f.) >0.07) or after six weeks (t = -0.36416, a(58 d.f.) >0.72) though the control group had been fed 230 % more feed (Fig. 13). After nine weeks the experimental group was significantly smaller than the control (t = 2.1933, a(21 d.f.) < 0.04) but after refeeding (12 weeks) there was again no significant difference (t = 0.786878, a(29 d.f.) > 0.43) and the control had been fed 264 % more feed (Fig. 13). At weeks 15 and 18 the control group was significantly heavier than the experimental group ( t = 3.87415, a (28 d.f.) < .001; t = 3.67697, a (32 d.f.) < 0.001 respectively) but 294% more feed. The differences in the 42 Table UJ: Summary of Results for Experiment 2 wk wt .(%chg) In. (%chg) con.fac. C . E . S . G . R . 2A 2B 2A 2B 2A 2B 2A 2B 2A 2B 3 16 .7 -11.2 2.9 0.2 4.7 -11.6 0.16 1.05 6 39 .5 27.9 7.3 6.6 8.3 4.6 0.17 0.30 0.72 0.67 9 118 .6 17.1 18.9 7.2 26.2 -5.4 0.32 1.06 12 165 .2 102.3 28.1 18.6 22.0 20.7 0.27 0.47 0.98 0.81 15 207 .6 64.8 36.5 19.1 16.5 -2.8 0.24 0.98 18 254 .6 123.7 42.9 24.9 18.2 14.1 0.21 0.32 0.90 0.61 450 400 -350 -300 -3 250 -200 150 -100 -50 -F i g u r e 1 2 Average Weight V, 2 :^ 12 Vs 1 31 \ 15 18 Time (Weeks) ZZ} 2A K 3 2B amount fed are reflected in the greater conversion efficiencies of the experimental group (Table HI). The carcass composition analysis results are shown in Table IV and Figs. 14 through 20. The only significant differences occurred in the samples taken after three weeks during which the experimental group had been starved and the control group fed. The experimental group was significantly higher in moisture (t= -2.92, a(8 d.f.) <0.02) (Fig. 14), and dry protein (t = -3.96, a(8 d.f.) <0.005) (Fig. 19) and lower in fat (t = 3.21, a(8 d.f.)<0.02) (Fig 15). After six weeks there were no significant differences between the control and experimental groups at a < 0.05. The average percentage change in carcass composition (Fig. 21) illustrates the large changes in fat content with starvation. These average percentage changes were then used to calculate the body composition of a hypothetical 100 g trout in each of the control and experimental groups initially, after three weeks , and after six weeks of growth at the average percentage change in weight for each group (Table V). This shows that during the three weeks of starvation the weight loss in the experimental group consists mainly of water (6.86 g) and equal amounts of fat (2.09 g) and protein (2.19 g). In the following three week feeding period the experimental fish gained 39.10 g in weight of which 26.88g is water, 5.62 g is fat and 6.02 g is protein. After six weeks the control group fish is 11.60 g heavier of which 7.28 g is water, 2.16 g is fat and only 1.43 g is protein at a cost of 134.3 g more feed than the experimental fish received (based on 5% body weight per day feeding level for both groups). 46 Table IV: Carcass Composition: Average Values (%) Week Group Moi sture Fat Protein Ash other Total 0 i n i t i a l 71.24 8.54 17.20 2.02 1.00 100 3 2A 70.51 10.22 16.40 2.22 0.65 100 2B 72.50 7.26 16.90 2.44 0.90 100 6 2A 70.64 10.20 16.12 2.40 0.64 100 2B 71.36 9.44 16.44 2.38 0.38 100 47 F i g u r e 1 4 Moisture Time (Weeks) Z Z l 2A E l 2B % Wet W e i g h t 49 F i g u r e 1 8 Dry Fat 35 F i g u r e 1 9 Dry Protein 70 T — X Dry W e i g h t 55 Table V : Changes in a 100 g fish Week Group Moisture Fat Protein Ash Other Total 0 I n i t i a l 71.24 8.54 17.20 2.02 1.00 100.0 3 2A 82.29 11.93 19.14 2.59 0.76 116.7 2B 64.38 6.45 15.01 2.17 0.80 88.8 6 2A 98.54 14.23 22.49 3.35 0.89 139.5 2B 91.26 12.07 21.03 3.04 0.49 127.9 56 Discussion Experiment 1 shows that the cycle period has a great effect on the C.G.R.. Groups ID, IE and IF all showed greater average percentage weight (Fig. 8) and length (Fig. 9) gains, higher specific growth rates (Fig. 10) and better conversion efficiencies (Fig. 11) than the control or the two other cyclically fed groups, though the differences were not statistically significant at the a = 0.05 level. Thus groups ID, IE and IF did at least as well if not better than the controls though they were fed for half as long. The control group's specific growth rate (0.407 % per day) is comparable to that of other constantly fed groups in the literature (Houlihan and Laurent, 1987, Davidson and Goldspink, 1977, Elliott, 1975) which were fed to satiation. Experiment 1 shows that during starvation, the greatest weight loss occurs during the first week of starvation as was found by Dobson and Holmes (1984). This large initial loss is probably due to the emptying of the gut (Elliott, 1972). As starvation continues the carcass composition results show a decrease in fat and an increase in moisture (Tables IV and V, Fig.21). This reflects the utilization of the visceral fat deposits and muscle lipids (Parker and Vanstone, 1966, Smith, 1981; Weatherly and Gill, 1981, Jezoerska et al, 1982) and the replacement with water of the muscle lipids (Idler and Bitners, 1959). The starvation periods used here are shorter than those required to initiate significant protein utilization (Denton and Yousef, 1976, Elliott, 1975, Weatherly and Gill, 1981) although changes in the metabolic rate 57 of the muscle tissues occur shortly after starvation begins (Loughna and Goldspink, 1984). The protein turnover rate is reduced during moderate starvation (Smith, 1981, Loughna and Goldspink, 1984). This reduction in protein metabolism may be the physiological mechanism underlying the C.G.R. and determining the ideal cycle periods for maximal growth and conversion efficiency. Upon starvation the basal metabolic rate drops and activity levels are reduced (Love, 1970, Love, 1980, Loughna and Goldspink, 1984). One of the main factors in reducing the basal metabolic rate is the reduction of the protein turnover rate in the white muscle tissue which comprises 70 percent of the fish's total wet body weight (Loughna and Goldspink, 1984). The protein turnover rate is determined by two factors, the degradation rate and the synthesis rate. In short term starvation both are reduced (Love, 1980, Smith, 1981, Loughna and Goldspink, 1984). As the starvation period increases the degradation rate increases in order to utilize the muscle tissue as an energy source through gluconeogenesis (Moon and Johnston, 1980). The very high growth rates associated with the C.G.R. would be possible if during the refeeding period the degradation rate remained low but the synthesis rate increased, allowing much more protein to be retained as growth. The inability of the shorter starvation periods (Groups IB, 1C, Table II) to facilitate a greater C.G.R. may be the result of the protein turnover rate not having decreased sufficiently. The three week period could allow the synthesis and degradation rates to fall but not be long enough to 58 cause the degradation rate to r ise as the prote in i s not yet an energy source ( S m i t h , 1981, Weather ly and G i l l , 1981). T h i s r ise i n the degradation rate m a y be the point at w h i c h a f u l l recovery f r o m the starvation p e r i o d i s not poss ible . T h e mechanics for the r a p i d g r o w t h i n the last w e e k o f refeeding is not k n o w n . In r a i n b o w trout r a p i d g r o w t h i s n o r m a l l y achieved through the recruitment o f n e w s m a l l musc le f ibres , as opposed to increas ing the diameter o f ex is t ing ones (Weatherly et al, 1979, W e a t h e r l y et al, 1980a,b, W e a t h e r l y and G i l l , 1984). T h e prote in synthetic rates i n trout are m u c h l o w e r than m a m m a l i a n rates and a greater propor t ion o f musc le tissue prote in synthesis i s retained as g r o w t h i n f i s h epaxia l musc le ( S m i t h , 1981). T h i s i s due to the l o w e r basal metabol ic demands o n the p o i k i l o t h e r m i c f i s h c o m p a r e d to the homeotherms ( S m i t h , 1981). D u r i n g starvation f i s h have m u c h l o w e r energy demands than m a m m a l s as they d o not mainta in a constant b o d y temperature different than the environment . T h e effect o f ra t ion l e v e l o n the C . G . R . i s minimal at the ra t ion leve ls used i n these experiments . T h e o n l y effect that ra t ion levels greater than three percent o f b o d y weight per day had was to decrease convers ion e f f i c i ency (Table II, F i g . 11). T h i s indicates that the f i s h f e d at the higher ra t ion levels were over fed . T h i s i s supported b y observations made d u r i n g the experiment that some o f the feed i n the higher ra t ion l e v e l groups w a s washed out o f the tanks and col lec ted at the o u t f l o w . It was not poss ib le to quanti tat ively measure this w i t h the setup used f o r these experiments . A l l groups were f e d a l l their ra t ion at one 59 time once per day. This would result in rapid filling of the gut and decrease residence time and assimilation efficiency during digestion (Jobling, 1981). If feeding were to be spread throughout the day even greater conversion efficiencies may be possible (Wurtsbaugh and Davis, 1977). Starvation periods of greater than seven days reduce the gastric evacuation rate and, the longer the starvation, the greater the reduction (Elliott, 1972). This reduced evacuation rate may increase assimilation efficiency during refeeding and contribute to the C.G.R. by making more energy available from the feed consumed. The second experiment shows the effect of repetitive cycles on the C.G.R.. The control group showed a very high specific growth rate and conversion efficiency compared to the control in the first experiment and other sources in the literature. The experimental group showed great gains in average percentage weight during the three refeeding periods (39.1%, 85.2%, 58.9%, Table Ul , Fig. 22). A mechanical breakdown in the system severely reduced water quality during the last two weeks of the experiment. This is reflected in the reduced growth and conversion efficiency during the last three week period for both the control and experimental groups. The experimental group was more severely affected because it occurred during the final week of the feeding period when most of the compensatory growth occurs (Figs. 5,7). The average weights of the control and experimental groups were not significantly different after six or twelve weeks (Fig. 12) even thought the control had received 230% more feed at six weeks 60 F i g u r e 22 Weight 260 - i and 260% more feed at 12 weeks (Fig. 13). The growth rate and conversion efficiency increased from the first cycle to the second and the average percentage increase in weight and length and the conversion efficiency were greater during the third cycle than the first (Table UJ). This indicates that the C.G.R. may increase with repetitive cycling. The carcass composition analysis (Tables IY, V, Figs. 14 to 21) show that there is no significant difference between the experimental and control fish after six weeks in moisture, fat, protein or ash. The experimental group tended to have slightly more protein and less fat than the controls. This indicates that the compensatory growth response does not affect the tissue qualities of the fish. The effect of fish size on the compensatory growth response can be inferred by comparing experiment 1 and the first six weeks of experiment 2. The average fish size in experiment 1 was 36.24 g while that for experiment 2 was 120.22 g. As fish increase in size their growth rate declines (Brett, 1979, Houlihan et al, 1986) but the results for experiment 1 and for the first 6 weeks of experiment 2 show very similar performance. Compensatory growth may increase the growth rate as well as the conversion efficiency in larger fish. The effect of fish size on the optimal cycle period is probably determined by the amount of lipid present to serve as the energy source during the starvation period and so determine the period before protein utilization occurs. Metabolic rate scales inversely with fish size (Wurtsbaugh and Davis, 1977). 62 Since the utilization of fat reserves is proportional to the metabolic rate it is probable that cycle period scales with fish size. The results presented here indicate that the compensatory growth response can be utilized to grow fish of comparable size to fish fed daily with far less feed. Application of these techniques to commercial aquaculture operations could lead to a considerable saving in feed cost, the major operating expense of most fish farms (Dr. F. Ming, Pers.Comm). 63 Conclusions The experiments show that equal or better growth rates can be achieved with less than half the feed through compensatory growth. The cycle period is critical. The three week cycles gave much higher average percentage changes in weight (27.24%, 27.91%, 24.91%) than either the control ( 18.68%) or the one (7.99%) and two (7.61%) week cycles. The results for average percentage change in length and specific growth rate show the same pattern. Most of the compensatory growth occurred in the last week of the three week feeding period. Group ID, which had a three week cycle period and a three percent of body weight per day ration level, produced a specific growth rate of over four percent at a conversion efficiency of 1.23 during the last week of its feeding period. The ration levels tested, three, five and seven percent of body weight per day, did not affect the growth rate. The higher ration levels only decreased the conversion efficiency (0.527 for 3%, 0.299 for 5% and 0.188 for 7%). This indicates that groups fed at greater than 3% were overfed. There were no significant differences between average weights of the control and experimental groups after six and twelve weeks though the control group had been fed 230 percent and 264 percent more feed respectively. After eighteen weeks the control group was significantly heavier than the experimental group but it had received 294 percent more feed. Carcass composition analysis of moisture, protein, fat and 64 ash show that compensatory growth has no significant effect on the overall body composition after a complete cycle. The effect of compensatory growth is to increase the growth rate and conversion efficiency during the refeeding period if the starvation and refeeding periods have been long enough. Further research in this area should focus on the effect of independently varying the starvation and refeeding periods, including longer refeeding periods to detennine when the increased growth rate due to compensatory growth rate falls to control levels. The underlying mechanisms of compensatory growth should also be examined by determining the changes in protein synthesis and degradation during compensatory growth. Histological studies to determine if muscle growth occurs through new fibre recruitment or increasing diameter of existing fibres would also be worthwhile. 65 References Ayles, G.B., Bernard, D. and Hendzel, M . 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Biostatistical Analysis, Second Edition, Prentice-Hall, New Jersey, 718pp. 71 Appendix: O r i g i n a l Data Group 1A/ I n i t i a l Values Dec. 6, 1987 Tag Weight Length New Tag 236 36.15 13 .8 242 17.81 10 .8 243 27.4 12 .8 288 22.83 12 .1 292 18.8 11 .3 297 17.85 10 .6 200 21.8 11 .7 201 17.65 10 .7 202 46.83 13 .8 204 24.74 12 .5 205 45.4 14 .2 206 44.68 15 .1 207 32.43 13 .8 209 19.45 11 .1 211 28.08 13 212 60.71 15 213 30.91 13 .3 214 26.3 12 .8 215 30.9 13 .4 216 21.23 12 217 12.05 9 .8 218 58.79 16 .3 223 37.94 14 .1 224 30.1 13 .7 225 25.58 12 .8 227 29.16 13 228 13.45 10 .4 230 24.85 11 .9 250 48.75 15 .1 251 18.7 10 .9 254 19.43 10 .8 258 28.28 13 .2 259 12.91 9 .5 260 31.75 13 .2 262 33.5 13 .9 263 26.52 12 .7 267 12.98 10 .2 268 36.3 14 282 38.8 14 .6 318 28.59 15 .1 28.301 12.414 avg sum 1160.3 count 40 fee d 58.019 g/day 469 515 458 236 72 Group 1A, Samples Sample 1, Dec. 10, 1987 Tag i n i t i a l sample % i n i t i a l sample % 200 21.8 21.5 -1.37614 11.7 11.7 0 214 26.3 25.98 -1.21673 12.8 12.3 -3.90625 215 30.9 30.14 -2.45954 13.4 13.4 0 216 21.42 21.23 -0.88702 12 12.1 0.833333 217 12.05 12.51 3.817427 9.8 9.65 -1.53061 225 25.58 25.71 0.508209 12.8 12.7 -0.78125 228 13.45 13.88 3.197026 10.4 10.45 0.480769 258 28.28 29.22 3.323903 13.2 12.8 -3.03030 268 36.3 36.15 -0.41322 14 14 0 288 22.83 23.13 1.314060 12.1 12.1 0 23.891 23.945 0.580795 12.22 12.12 -0.79343 Sample 2, Dec. 11 Tag i n i t i a l sample % i n i t a i l sample % 211 28.08 27.84 -0.85470 13 12.7 -2.30769 214 26.3 26.55 0.950570 12.8 12.5 -2.34375 216 21.42 21.31 -0.51353 12 12 0 217 12.05 12.95 7.468879 9.8 9.8 0 223 37.94 38.34 1.054296 14.1 13.6 -3.54609 228 13.45 13.5 0.371747 10.4 10.4 0 258 28.28 28.61 1.166902 13.2 12.7 -3.78787 260 31.75 31.58 -0.53543 13.2 13.2 0 268 36.3 36.29 -0.02754 14 13.9 -0.71428 288 22.83 21.35 -6.48269 12.1 12.1 0 25.84 25.832 0.259847 12.46 12.29 -1.26997 Sample 3, Dec. 14 Tag i n i t i a l sample % i n i t i a l sample % 204 24.74 24.17 -2.30396 12.5 12.6 0.8 214 26.3 25.7 -2.28136 12.8 12.6 -1.5625 217 12.05 12.28 1.908713 9.8 9.8 0 224 30.1 30.4 0.996677 13.7 13.5 -1.45985 227 29.16 28.61 -1.88614 13 12.7 -2.30769 228 13.45 13.47 0.148698 10.4 10.4 0 258 28.28 28.13 -0.53041 13.2 12.8 -3.03030 260 31.75 31.16 -1.85826 13.2 13.2 0 267 12.98 12.88 -0.77041 10.2 10.3 0.980392 268 36.3 35.78 -1.43250 14 13.9 -0.71428 24.511 24.258 -0.80089 12.28 12.18 -0.72942 73 Sample 4, Dec. 15 Tag i n i t i a l sample % i n i t i a l sample % 200 21.8 20.38 -6.51376 11.7 11.8 0.854700 204 24.74 24.62 -0.48504 12.5 12.5 0 207 32.43 32.22 -0.64754 13.8 13.4 -2.89855 212 60.71 67.79 11.66199 15 14.9 -0.66666 228 13.45 13.31 -1.04089 10.4 10.5 0.961538 258 28.28 27.92 -1.27298 13.2 13 -1.51515 260 31.75 31.04 -2.23622 13.2 13.3 0.757575 262 33.5 33.82 0.955223 13.9 13.8 -0.71942 267 12.98 12.71 -2.08012 10.2 10.4 1.960784 268 36.3 35.63 -1.84573 14 14 0 29.594 29.944 -0.35050 12.79 12.76 -0.12651 Sample 5/ Dec. 17 Tag i n i t i a l sample % chg i n i t i a l sample % chg 214 26.3 25.58 -2.73764 12.8 12.7 -0.78125 215 30.9 30.39 -1.65048 13.4 13.3 -0.74626 216 21.23 20.46 -3.62694 12 12 0 217 12.05 12.06 0.082987 9.8 9.7 -1.02040 223 37.94 36.43 -3.97996 14.1 13.8 -2.12765 225 25.58 24.78 -3.12744 12.8 12.8 0 227 29.16 28.32 -2.88065 13 12.9 -0.76923 258 28.28 27.84 -1.55586 13.2 12.9 -2.27272 262 33.5 33.96 1.373134 13.9 13.8 -0.71942 268 36.3 35.7 -1.65289 14 14 0 28.124 27.552 -1.97557 12.9 12.79 -0.84369 Sample 6, Dec. 18, 1987 I n i t i a l Sample Tag i n i t i a l sample % chg % chg 214 26.3 25.61 -2.62357 12.8 12.5 -2.34375 223 37.94 36.72 -3.21560 14.1 13. 6 -3.54609 228 13.45 13.49 0.297397 10.4 10.5 0.961538 243 27.4 24.84 -9.34306 12.8 12.6 -1.5625 258 28.28 27.79 -1.73267 13.2 12.9 -2.27272 262 33.5 33.93 1.283582 13.9 13.7 -1.43884 267 12.98 12.57 -3.15870 10.2 10.3 0.980392 268 36.3 35.55 -2.06611 14 13.9 -0.71428 469 18.8 15.9 -15.4255 11.3 11.3 0 515 19.45 20.47 5.244215 11.1 11.1 0 25.44 24.687 -3.07400 12.38 12.24 -0.99362 74 Sample 7, Dec. 22 Tag i n i t i a l sample % chg i n i t i a l sample % chg 202 46.83 57.01 21.73820 13.8 15.2 10.14492 207 32.43 35.07 8.140610 13.8 13.6 -1.44927 213 30.91 30.21 -2.26463 13.3 13.2 -0.75187 214 26.3 24.76 -5.85551 12.8 12.6 -1.5625 215 30.9 29.61 -4.17475 13.4 13.3 -0.74626 224 30.1 29.85 -0.83056 13.7 13.5 -1.45985 258 28.28 27.68 -2.12164 13.2 12.8 -3.03030 260 31.75 30.26 -4.69291 13.2 13.2 0 262 33.5 32.49 -3.01492 13.9 13.9 0 263 26.52 25.79 -2.75263 12.7 12.7 0 267 12.98 12.59 -3.00462 10.2 10.4 1.960784 268 36.3 34.96 -3.69146 14 13.9 -0.71428 318 28.59 27.988 -2.10563 15.1 13.1 -13.2450 30.41461 30.636 -0.35619 13.31538 13.18461 -0.83489 Sample 8, Dec. 25 Tag i n i t i a l sample % chg i n i t i a l sample % chg 204 24.74 22.91 -7.39692 12.5 12.3 -1.6 206 44.68 71.05 59.01969 15.1 16.9 11.92052 217 12.05 11.28 -6.39004 9.8 9.7 -1.02040 223 37.94 35.7 -5.90405 14.1 13.7 -2.83687 236 36.15 42.54 17.67634 13.8 14.2 2.898550 258 28.28 27.94 -1.20226 13.2 12.8 -3.03030 260 31.75 29.66 -6.58267 13.2 13.1 -0.75757 262 33.5 31.41 -6.23880 13.9 13.8 -0.71942 469 18.8 17.2 -8.51063 11.3 11.2 -0.88495 515 19.45 24.99 28.48329 11.1 11.5 3.603603 28.734 31.468 6.295392 12.8 12.92 0.757313 Sample 9, Dec. 29 Tag i n i t i a l sample % chg i n i t i a l sample % chg 204 24.74 22.81 -7.80113 12.5 12.4 -0.8 207 32.43 36.49 12.51927 13.8 13.7 -0.72463 211 28.08 26.96 -3.98860 13 12.4 -4.61538 212 60.71 79.18 30.42332 15 15.4 2.666666 217 12.05 11.53 -4.31535 9.8 9.7 -1.02040 224 30.1 29.03 -3.55481 13.7 13.4 -2.18978 227 29.16 26.92 -7.68175 13 12.8 -1.53846 258 28.28 27.53 -2.65205 13.2 12.7 -3.78787 260 31.75 29.62 -6.70866 13.2 13.2 0 262 33.5 31.21 -6.83582 13.9 13.6 -2.15827 31.08 32.128 -0.05955 13.11 12.93 -1.41681 75 Sample 10, Jan. 1 Tag i n i t i a l sample % chg i n i t i a l sample % chg 204 24.74 22.68 -8.32659 12.5 12.5 0 206 44.68 77.48 73.41092 15.1 17.6 16.55629 215 30.9 29.55 -4.36893 13.4 13.2 -1.49253 217 12.05 11.43 -5.14522 9.8 9.8 0 224 30.1 28.68 -4.71760 13.7 13.4 -2.18978 225 25.58 25.03 -2.15011 12.8 12.6 -1.5625 260 31.75 29.47 -7.18110 13.2 13.3 0.757575 268 36.3 35.1 -3.30578 14 14.1 0.714285 458 58.79 67.29 14.45824 16.3 15.8 -3.06748 469 18.8 17 -9.57446 11.3 11.3 0 31.369 34.371 4.309932 13.21 13.36 0.971584 Sample 11, Jan 5, 1988 Tag i n i t i a l sample % chg i n i t i a l sample % chg 207 32.43 38.24 17.91551 13.8 13.8 0 217 12.05 11.28 -6.39004 9.8 9.8 0 224 30.1 28.63 -4.88372 13.7 13.4 -2.18978 227 29.16 27.38 -6.10425 13 12.8 -1.53846 254 19.43 30.11 54.96654 10.8 12.3 13.88888 260 31.75 29.68 -6.51968 13.2 13.2 0 262 33.5 31.57 -5.76119 13.9 13.7 -1.43884 263 26.52 25.42 -4.14781 12.7 12.5 -1.57480 318 28.59 28.39 -0.69954 15.1 12.9 -14.5695 458 58.79 68.81 17.04371 16.3 16.2 -0.61349 30.232 31.951 5.541951 13.23 13.06 -0.80360 Sample 12, Jan. 8, 1988 Tag i n i t i a l sample % chg i n i t i a l sample % chg 211 28.08 48.09 71.26068 13 12.8 -1.53846 217 12.05 11.3 -6.22406 9.8 9.7 -1.02040 223 37.94 40.4 6.483921 14.1 14 -0.70921 224 30.1 28.61 -4.95016 13.7 13.4 -2.18978 227 29.16 26.8 -8.09327 13 12.8 -1.53846 236 36.15 51.57 42.65560 13.8 15 8.695652 254 19.43 29.48 51.72413 10.8 12.4 14.81481 258 28.28 26.93 -4.77369 13.2 12.8 -3.03030 263 26.52 25.87 -2.45098 12.7 12.6 -0.78740 318 28.59 28.68 0.314795 15.1 12.9 -14.5695 27.63 31.773 14.59469 12.92 12.84 -0.18731 76 Sample 13, Jan. 15 i n i t i a l sample % chg i n i t i a l sample % chg 206 44.68 82.31 84 .22112 15.1 18.2 20.52980 211 28.08 27.53 -1 .95868 13 12.7 -2.30769 212 60.71 89.11 46 .77977 15 16.1 7.333333 217 12.05 11.18 -7 .21991 9.8 9.7 -1.02040 225 25.58 24.51 -4 .18295 12.8 12.8 0 227 29.16 26.53 -9 .01920 13 12.8 -1.53846 230 24.85 48.91 96 .82092 11.9 14 17.64705 258 28.28 26.39 -6 .68316 13.2 12.8 -3.03030 260 31.75 29.29 -7 .74803 13.2 13.2 0 268 36.3 34.77 -4 .21487 14 13.9 -0.71428 32.144 40.053 18 .67949 13.1 13.62 3.689904 77 Group IB, I n i t i a l Values Dec. 4, 1987 Tag Weight Length New Tag 210 30.61 13.4 233 31.2 13.1 234 30.44 13.1 235 34.38 13.8 237 39.68 14 238 23.78 12 239 20.22 11.1 240 35.5 14.1 241 19.49 11.1 244 20.61 11.2 281 19.06 11.4 284 34.16 13.5 285 24.85 12.5 286 55.97 15.1 287 20.24 11.9 289 37.75 14.9 290 20.81 11.5 291 42.28 13.4 301 35.24 14.5 307 33.75 13 308 24.43 11.7 309 40.55 13.4 321 21.5 11.6 324 13.9 10 328 18.38 10.6 331 77.32 15.4 334 21.35 11.7 335 30.9 13.2 336 37.08 13.8 337 19.69 11.5 338 22.78 11.6 354 49.3 14.8 367 25.53 12.4 369 19.29 11.1 374 12.31 10 383 23.62 12.1 384 26.19 12.5 385 54.75 15.5 386 80.05 16.1 387 32.59 12.8 count 40 avg 31.538 12.76 sum 1261.5 feed 63.076 g/day 261 258 521 78 Group IB, Samples Dec. 8, 1987 Tag I n i t i a l Sample % I n i t i a l Sample % 281 19.06 18.3 -3.98740 11.4 11.3 -0.87719 284 34.16 32.35 -5.29859 13.5 13.6 0.740740 285 24.85 23.92 -3.74245 12.5 12.7 1.6 289 37.75 38.57 2.172185 14.9 14.8 -0.67114 301 35.24 34.47 -2.18501 14.5 14.1 -2.75862 307 33.75 32.23 -4.50370 13 13.2 1.538461 308 24.43 25.37 3.847728 11.7 12 2.564102 331 77.32 61.43 -20.5509 15.4 15.4 0 336 37.08 37.43 0.943905 13.8 13.8 0 374 12.31 12.37 0.487408 10 9.9 -1 33.595 31.644 -3.28169 13.07 13.08 0.113635 Dec. 11, 1987 Tag I n i t i a l Sample % I n i t i a l Sample % 210 30.61 30.53 -0.26135 13.4 13.1 -2.23880 238 23.78 21.91 -7.86375 12 11.7 -2.5 261 30.44 30.35 -0.29566 13.1 13.2 0.763358 281 19.06 18.05 -5.29905 11.4 11.5 0.877192 289 37.75 38.17 1.112582 14.9 14.7 -1.34228 291 42.28 35.82-15.2790 13.4 13.6 1.492537 301 35.24 34.17 -3.03632 14.5 14.1 -2.75862 336 37.08 36.78 -0.80906 13.8 13.9 0.724637 384 26.19 25.92 -1.03092 12.5 12.7 1.6 387 32.59 32.29 -0.92052 12.8 12.8 0 31.502 30.399 -3.36831 13.18 13.13 -0.33819 Dec. 15, 1987 Tag I n i t i a l Sample % I n i t i a l Sample % 238 23.78 25.06 5.382674 12 11.9 -0.83333 261 30.27 13.3 -56.0621 13.1 13.3 1.526717 281 19.06 18.98 -0.41972 11.4 11.4 0 285 24.85 23.82 -4.14486 12.5 12.6 0.8 289 37.75 36.92 -2.19867 14.9 14.9 0 309 40.55 46.18 13.88409 13.4 13.7 2.238805 331 77.32 70.4 -8.94981 15.4 15.9 3.246753 383 23.62 22.95 -2.83657 12.1 12 -0.82644 384 26.19 25.35 -3.20733 12.5 12.6 0.8 387 32.59 32.18 -1.25805 12.8 13 1.5625 33.598 31.514 -5.98103 13.01 13.13 0.851499 79 Dec. 18, 1987 Tag I n i t i a l Sample % I n i t i a l Sample % chg. 210 30.61 49.65 62.20189 13.4 13.3 -0.74626 261 30.44 30.17 -0.88699 13.1 13.2 0.763358 281 19.06 19.31 1.311647 11.4 11.4 0 286 55.97 65.28 16.63391 15.1 16.1 6.622516 289 37.75 36.55 -3.17880 14.9 14.8 -0.67114 301 35.24 32.92 -6.58342 14.5 14.2 -2.06896 308 24.43 24.22 -0.85959 11.7 11.9 1.709401 374 12.31 12.45 1.137286 10 10 0 383 23.62 23.28 -1.43945 12.1 12 -0.82644 521 80.05 54.41 -32.0299 16.1 16.4 1.863354 34.948 34.824 3.630647 13.23 13.33 0.664580 Dec. 22, 1987 Tag I n i t i a l Sample % I n i t i a l Sample % 210 30.61 29.1 -4.93302 13.4 13.5 0.746268 235 34.38 33.92 -1.33798 13.8 13.9 0.724637 238 23.78 25.84 8.662741 12 12.1 0.833333 281 19.06 18.8 -1.36411 11.4 11.4 0 285 24.85 22.99 -7.48490 12.5 12.8 2.4 286 55.97 57.65 3.001608 15.1 16.4 8.609271 289 37.75 36.18 -4.15894 14.9 14.8 -0.67114 301 35.24 32.58 -7.54824 14.5 14.1 -2.75862 328 18.38 19.18 4.352557 10.6 10.9 2.830188 354 49.3 49.32 0.040567 14.8 15.6 5.405405 32.932 32.556 -1.07697 13.3 13.55 1.811934 Dec. 25, 1987 Tag I n i t i a l Sample % I n i t i a l Sample % 258 25.53 22.82 -10.6149 12.4 12.7 2.419354 261 30.44 29.5 -3.08804 13.1 13.1 0 281 19.06 18.46 -3.14795 11.4 11.5 0.877192 286 55.97 55.85 -0.21440 15.1 16.4 8.609271 289 37.75 35.72 -5.37748 14.9 14.7 -1.34228 301 35.24 32.45 -7.91713 14.5 14.2 -2.06896 309 40.55 43.8 8.014796 13.4 14.8 10.44776 328 18.38 18.89 2.774755 10.6 10.9 2.830188 354 49.3 48.29 -2.04868 14.8 15.6 5.405405 383 23.62 22.63 -4.19136 12.1 12 -0.82644 33.584 32.841 -2.58104 13.23 13.59 2.635148 80 Dec. 29, 1987 Tag Tag Tag I n i t i a l Sample % I n i t i a l Sample % 210 30.61 28.43 -7.12185 13.4 13.2 -1.49253 235 34.38 34.66 0.814426 13.8 13.9 0.724637 237 39.68 38.23 -3.65423 14 14.2 1.428571 238 23.78 28.71 20.73170 12 12.2 1.666666 285 24.85 22.48 -9.53722 12.5 12.8 2.4 289 37.75 35.32 -6.43708 14.9 14.8 -0.67114 291 42.28 42. 62 0.804162 13.4 14 4.477611 308 24.43 23.15 -5.23945 11.7 11.8 0.854700 336 37.08 35.31 -4.77346 13.8 13.9 0.724637 374 12.31 11.98 -2.68074 10 10 0 30.715 30.089 -1.70937 12.95 13.08 1.011314 1/ : 1988 I n i t i a l Sample % I n i t i a l Sample % 210 30.61 28.69 -6.27245 13.4 13.4 0 235 34.38 34.35 -0.08726 13.8 13.8 0 237 39.68 38.02 -4.18346 14 14.4 2.857142 281 19.06 20.1 5.456453 11.4 11.6 1.754385 285 24.85 22.74 -8.49094 12.5 12.7 1.6 289 37.75 35.02 -7.23178 14.9 14.8 -0.67114 308 24.43 23.68 -3.06999 11.7 11.9 1.709401 309 40.55 59.44 46.58446 13.4 14.9 11.19402 354 49.3 59.24 20.16227 14.8 15.8 6.756756 383 23.62 22.5 -4.74174 12.1 11.9 -1.65289 32.423 34.378 3.812552 13.2 13.52 2.354768 5, : 1988 I n i t i a l Sample % chg I n i t i a l Sample % chg 210 30.61 28.23 -7.77523 13.4 13.2 -1.49253 235 34.38 33.56 -2.38510 13.8 14 1.449275 238 23.78 29.6 24.47434 12 12.6 5 261 30.44 29.58 -2.82522 13.1 13.3 1.526717 285 24.85 23.2 -6.63983 12.5 12.6 0.8 289 37.75 35.28 -6.54304 14.9 14.8 -0.67114 308 24.43 23.46 -3.97052 11.7 11.8 0.854700 309 40.55 53.71 32.45376 13.4 15.2 13.43283 331 77.32 82.01 6.065700 15.4 17 10.38961 336 37.08 35.1 -5.33980 13.8 13.9 0.724637 383 23.62 22.45 -4.95342 12.1 11.9 -1. 65289 34.98272 36.01636 2.051053 13.28181 13.66363 2.760109 81 Jan. 8, 1988 Tag Tag I n i t i a l Sample % I n i t i a l Sample % 210 30.61 28.43 -7.12185 13.4 13 .4 0 237 39.68 37.06 -6.60282 14 14 .4 2.857142 261 30.44 29.72 -2.36530 13.1 13 .2 0.763358 281 19.06 19. 65 3.095487 11.4 11 .5 0.877192 284 34.16 42.91 25.61475 13.5 14 .8 9.629629 289 37.75 22.91 -39.3112 14.9 12 .7 -14.7651 331 77.32 79.72 3.103983 15.4 17 .1 11.03896 336 37.08 34.72 -6.36461 13.8 13 .9 0.724637 383 77.32 82.01 6.065700 12.1 12 -0.82644 521 80.05 61.8 -22.7982 16.1 17 .1 6.211180 46.347 43.893 -4.66841 13.77 14. 01 1.651055 15, 1988 I n i t i a l Sample % I n i t i a l Sample % 210 30.61 28.41 -7.18719 13.4 13 .2 -1.49253 237 39.68 37.38 -5.79637 14 14 .2 1.428571 261 30.44 29.42 -3.35085 13.1 13 .3 1.526717 281 19.06 20.52 7.660020 11.4 11 .6 1.754385 285 24.85 22.88 -7.92756 12.5 12 .6 0.8 289 37.75 34.55 -8.47682 14.9 14 .8 -0.67114 307 33.75 52.38 55.2 13 15 .1 16.15384 308 24.43 22.99 -5.89439 11.7 11 .7 0 309 40.55 65.75 62.14549 13.4 15 .7 17.16417 383 23.62 22.08 -6.51989 12.1 11 .9 -1.65289 30.474 33.636 7.985242 12.95 13. 41 3.501112 82 Group IC, I n i t i a l Values Dec. 10/ 1987 Tag Weight Length New Tag 205 39 14.5 461 222 50.35 14.6 226 24.18 12.2 231 22.21 11.7 257 20.39 11.2 264 15.13 10.5 315 18.13 11 317 28.65 13.1 319 29.74 12.9 320 29.52 12.8 322 45.34 14.4 323 28 12.6 325 28.85 11.8 327 64.12 15.8 329 22.77 12.3 330 21.52 11.3 332 29.44 12.9 333 18.98 11.7 350 75.1 15.6 351 38.82 14.4 352 23.25 11.2 353 26.75 12.5 358 59.2 15.5 361 30.87 13.3 363 13.45 10.1 364 27.4 13.1 365 41.95 14 366 27.5 11.4 368 24.3 12 370 28.91 12.9 371 78.62 16.2 372 35.25 14 373 20.45 11.6 375 26.37 12.3 376 35.67 13.7 377 32 13.4 378 11.98 9.5 379 23.34 12.6 380 21.6 11.5 381 27.82 13.2 382 24.55 12.6 count 41 avg 31.49926 12.77804 sum 1291.47 feed 64.5735 g/day 83 Group IC, Samples Dec. 14, 1987 Tag I n i t i a l Sample % I n i t i a l Sample % 222 50.53 45.33 -10.2909 14.6 14.9 2.054794 322 45.34 45.33 -0.02205 14.4 14.5 0.694444 329 22.77 22.28 -2.15195 12.3 12.3 0 332 29.44 28.84 -2.03804 12.9 12.9 0 351 38.82 39.85 2.653271 14.4 14.5 0.694444 353 26.75 26.83 0.299065 12.5 12.5 0 358 59.2 54.97 -7.14527 15.5 15.8 1.935483 371 78.62 75.37 -4.13380 16.2 16.5 1.851851 372 35.25 35.01 -0.68085 14 14 0 376 35.67 35.25 -1.17746 13.7 13.8 0.729927 42.239 40.906 -2.46880 14.05 14.17 0.796094 Dec. 17, 1987 Tag I n i t i a l Sample % I n i t i a l Sample % 319 29.74 29.772 0.107599 12.9 12.9 0 329 22.77 21.92 -3.73298 12.3 12.3 0 332 29.44 27.86 -5.36684 12.9 12.5 -3.10077 353 26.75 26.01 -2.76635 12.5 12.6 0.8 358 59.2 52.57 -11.1993 15.5 15.8 1.935483 365 41.95 38.86 -7.36591 14 14.2 1.428571 371 78.62 71.88 -8.57288 16.2 16.6 2.469135 376 35.67 34.78 -2.49509 13.7 13.8 0.729927 379 23.34 23.21 -0.55698 12.6 12.5 -0.79365 381 27.82 27.44 -1.36592 13.2 13.1 -0.75757 382 24.55 23.72-3.38085 12.6 12.6 0 36.35 34.36563 -4.24505 13.49090 13.53636 0.246465 Dec. 21, 1987 Tag I n i t i a l Sample % chg I n i t i a l Sample % chg 222 50.35 53.58 6.415094 14.6 14.9 2.054794 257 20.39 19.71 -3.33496 11.2 11.6 3.571428 320 29.52 28.87 -2.20189 12.8 12.5 -2.34375 329 22.77 21.7 -4.69916 12.3 12.2 -0.81300 353 26.75 26.11 -2.39252 12.5 12.7 1.6 358 59.2 50.02 -15.5067 15.5 15.8 1.935483 371 78.62 69.41 -11.7145 16.2 16.4 1.234567 372 35.25 33.77 -4.19858 14 13.9 -0.71428 381 27.82 27.29 -1.90510 13.2 13 -1.51515 382 24.55 24.01 -2.19959 12.6 12.6 0 37.522 35.447 -4.17380 13.49 13.56 0.501007 84 Dec. 24, 1987 Tag I n i t i a l Sample % chg I n i t i a l Sample % chg 257 20.39 19.1 -6.32663 11.2 11.7 4.464285 319 29.74 28.84 -3.02622 12.9 12.8 -0.77519 329 22.77 20.98 -7.86122 12.3 12 -2.43902 332 29.44 27.53 -6.48777 12.9 12.8 -0.77519 353 26.75 25.6 -4.29906 12.5 12.6 0.8 365 41.95 36.69 -12.5387 14 14.3 2.142857 372 35.25 33.31 -5.50354 14 14 0 379 23.34 22.58 -3.25621 12.6 12.6 0 381 27.82 26.59 -4.42127 13.2 13.1 -0.75757 382 24.55 23.47 -4.39918 12.6 12.6 0 28.2 26.469 -5.81198 12.82 12.85 0.266015 Dec. 28, 1987 Tag I n i t i a l Sample % chg I n i t i a l Sample % chg 320 29.52 28.89 -2.13414 12.8 12.7 -0.78125 322 45.34 49.91 10.07940 14.4 14.8 2.777777 329 22.77 21.25 -6.67545 12.3 12.1 -1.62601 332 29.44 27.08 -8.01630 12.9 12.7 -1.55038 350 75.1 74.59 -0.67909 15.6 16.1 3.205128 353 26.75 25.62 -4.22429 12.5 12.5 0 358 59.2 58.69 -0.86148 15.5 16.1 3.870967 372 35.25 34.02 -3.48936 14 14 0 379 23.34 23.38 0.171379 12.6 12.5 -0.79365 381 27.82 26.53 -4.63695 13.2 13.1 -0.75757 37.453 36.996 -2.04663 13.58 13.66 0.434499 Dec. 31, 1987 Tag I n i t i a l Sample % chg I n i t i a l Sample % chg 222 50.35 69.63 38.29195 14.6 15.5 6.164383 319 29.74 28.74 -3.36247 12.9 13 0.775193 332 29.44 26.95 -8.45788 12.9 12.8 -0.77519 351 38.82 39.15 0.850077 14.4 14.6 1.388888 365 41.95 49.46 17.90226 14 14.6 4.285714 372 35.25 36.35 3.1205.67 14 14.1 0.714285 376 35.67 34.47 -3.36417 13.7 13.8 0.729927 379 23.34 23.6 1.113967 12.6 12.5 -0.79365 381 27.82 26.71 -3.98993 13.2 13.2 0 382 24.55 23.58 -3.95112 12.6 12.6 0 33.693 35.864 3.815325 13.49 13.67 1.248954 85 Jan. 4, 1988 Tag I n i t i a l Sample % chg I n i t i a l Sample % chg 205 39 37.46 -3.94871 14.5 14.5 0 222 50.35 67.17 33.40615 14.6 15.8 8.219178 319 29.74 28.79 -3.19435 12.9 13 0.775193 320 29.52 30.4 2.981029 12.8 12.8 0 351 38.82 40.99 5.589902 14.4 14.8 2.777777 353 26.75 25.28 -5.49532 12.5 12.7 1.6 358 59.2 67.21 13.53040 15.5 16.6 7.096774 364 27.4 28.85 5.291970 13.1 13 -0.76335 365 41.95 54.11 28.98688 14 14.9 6.428571 372 35.25 37.73 7.035460 14 14.2 1.428571 376 35.67 34.52 -3.22399 13.7 13.9 1.459854 381 27.82 26.85 -3.48670 13.2 13.1 -0.75757 382 24.55 23.19 -5.53971 12.6 12.7 0.793650 35.84769 38.65769 5.533308 13.67692 14 2.235279 Jan. 7, 1988 Tag I n i t i a l Sample % chg. I n i t i a l Sample % chg. 222 50.35 67.14 33.34657 14.6 15.9 8.904109 320 29.52 30.5 3.319783 12.8 12.8 0 322 45.34 53.52 18.04146 14.4 15.6 8.333333 353 26.75 25.31 -5.38317 12.5 12.6 0.8 358 59.2 64.59 9.104729 15.5 16.7 7.741935 365 41.95 50.81 21.12038 14 15.1 7.857142 372 35.25 38.92 10.41134 14 14.3 2.142857 376 35.67 34.48 -3.33613 13.7 13.9 1.459854 381 27.82 26.51 -4.70884 13.2 13.1 -0.75757 382 24.55 23.12 -5.82484 12.6 12.6 0 37.64 41.49 7.609127 13.73 14.26 3.648165 Jan. 14, 1988 Tag I n i t i a l Sample % Chg I n i t i a l Sample % Chg 226 24.18 23.28 -3.72208 12.2 12.2 0 257 20.39 18.86 -7.50367 11.2 11.7 4.464285 319 29.74 28.57 -3.93409 12.9 13 0.775193 320 29.52 29.18 -1.15176 12.8 12.8 0 322 45.34 51.12 12.74812 14.4 15.6 8.333333 329 22.77 21.84 -4.08432 12.3 12.2 -0.81300 332 29.44 27.09 -7.98233 12.9 12.8 -0.77519 365 41.95 46.05 9.773539 14 15.3 9.285714 379 23.34 23.68 1.456726 12.6 12.6 0 381 27.82 26.31 -5.42774 13.2 13.1 -0.75757 29.449 29.598 -0.98276 12.85 13.13 2.051274 86 Jan. 21, 1988 I n i t i a l Sample % Chg I n i t i a l Sample % Chg 222 50.35 62.62 24.36941 14.6 16 9.589041 257 20.39 18.3 -10.2501 11.2 11.6 3.571428 319 29.74 28.35 -4.67383 12.9 12.8 -0.77519 320 29.52 28.89 -2.13414 12.8 12.7 -0.78125 329 22.77 21.62 -5.05050 12.3 12.3 0 332 29.44 26.78 -9.03532 12.9 12.7 -1.55038 353 26.75 24.92 -6.84112 12.5 12.6 0.8 364 27.4 27.8 1.459854 13.1 13.1 0 372 35.25 36.14 2.524822 14 14.1 0.714285 461 39 38.52 -1.23076 14.5 14.6 0.689655 31.061 31.394 -1.08617 13.08 13.25 1.225757 87 Group ID, I n i t i a l Values Dec. 15, 1987 Tag Weight Length New Tag 255 28.9 12.2 447 60.47 15.5 558 55.65 13.8 559 65.95 15.7 560 60.63 14.8 561 25.25 12.4 562 28.62 12.6 563 97.72 17.5 565 74.23 15.9 566 36.6 13.8 567 40.21 14.8 568 47.4 14.1 569 74.91 16.2 570 24.4 12.4 564 571 39.7 13.3 572 76.69 16.2 573 29.01 12.2 574 69.03 16.3 575 28.32 12.3 576 77.96 16.4 577 23.58 10.9 579 79.24 16.3 582 30.9 13.6 583 69.11 16.5 534 584 50.49 14.3 585 14.28 10.5 586 29.65 13.2 587 23.78 12.3 588 24.18 12.3 589 109.14 18.4 590 38.14 14.3 591 38.8 13.1 592 25.48 12.9 593 26.18 12.5 594 20.82 11.2 595 62.22 15.1 596 31.61 13.7 597 22.9 11.5 598 26.72 13.4 599 38.3 12.6 501 500 27.18 13.3 503 27.18 11.9 504 36.65 13.3 509 53.98 14.8 523 33.08 13.3 528 27.69 12.2 count 4 6 avg. 44.19413 13.82173 sum 2032.93 feed 60.9879 g/day 88 Group ID, Samples Sample 1, Dec. 25, 1987. Tag i n i t i a l sample % d i f f . I n i t i a l Sample % chg. 501 38.3 34.73 -9.32114 12.6 12 -4.76190 523 33.08 33.97 2.690447 13.3 13.2 -0.75187 565 74.23 65.29 -12.0436 15.9 16.3 2.515723 583 69.11 60.81 -12.0098 16.5 16.7 1.212121 588 24.18 24.03 -0.62034 12.3 12.2 -0.81300 589 109.14 99.86 -8.50284 18.4 18.3 -0.54347 591 38.8 34.68 -10.6185 13.1 13.6 3.816793 593 26.18 25.88 -1.14591 12.5 12.5 0 597 22.9 20.93 -8.60262 11.5 12 4.347826 598 26.72 26.45 -1.01047 13.4 13-2.98507 46.264 42.663 -6.11849 13.95 13.98 0.203711 Sample 2, Jan. 1, 1988. Tag i n i t i a l sample % d i f f . I n i t i a l Sample % chg. 501 38.3 33.82 -11.6971 12.6 12.3 -2.38095 523 33.08 33.5 1.269649 13.3 13.4 0.751879 558 55.65 47.2 -15.1841 13.8 13.2 -4.34782 560 60.63 51.9 -14.3988 14.8 15.4 4.054054 566 36.6 35.99 -1.66666 13.8 13.8 0 574 69.03 61.35 -11.1255 16.3 16.9 3.680981 576 77.96 67.73 -13.1221 16.4 16.9 3.048780 588 24.18 23.55 -2.60545 12.3 12.2 -0.81300 594 20.82 20.15 -3.21805 11.2 11.2 0 596 31.61 31.15 -1.45523 13.7 13.8 0.729927 44.786 40.634 -7.32036 13.82 13.91 0.472383 Sample 3, Jan. 8, 1988. Tag i n i t i a l sample % d i f f . I n i t i a l Sample % chg. 255 28.9 24.16 -16.4013 12.2 12.2 0 504 36.65 33.41 -8.84038 13.3 13.5 1.503759 509 53.98 49.4 -8.48462 14.8 14.9 0.675675 528 27.69 25.29 -8.66738 12.2 12.4 1.639344 558 55.65 45.81 -17.6819 13.8 13.5 -2.17391 577 23.58 18.78 -20.3562 10.9 10.9 0 586 29.65 28.5 -3.87858 13.2 13.2 0 589 109.14 95.83 -12.1953 18.4 18.6 1.086956 597 22.9 20.25 -11.5720 11.5 11.9 3.478260 598 26.72 26.12 -2.24550 13.4 12.9 -3.73134 41.486 36.755 -11.0323 13.37 13.4 0.247874 89 Sample 4, Jan. 15, 1988 Tag i n i t i a l sample % d i f f . I n i t i a l Sample % chg. 528 27.69 28.91 4.405922 12.2 12.6 3.278688 534 69.11 75.38 9.072493 16.5 16.7 1.212121 564 24.4 24.45 0.204918 12.4 12.3 -0.80645 574 69.03 65.35 -5.33101 16.3 17.2 5.521472 575 28.32 27.58 -2.61299 12.3 12.4 0.813008 576 77.96 67.72 -13.1349 16.4 17.1 4.268292 586 29.65 28.6 -3.54131 13.2 13.2 0 589 109.14 95.39 -12.5984 18.4 18.8 2.173913 594 20.82 19.82 -4.80307 11.2 11.1 -0.89285 598 26.72 25.61 -4.15419 13.4 13 -2.98507 48.284 45.881 -3.24926 14.23 14.44 1.258311 Sample 5, Jan. 22, 1988. Tag i n i t i a l sample % d i f f . I n i t i a l Sample % chg, 500 27.18 27.36 0.662251 13 .3 13.3 0 528 27.69 36.58 32.10545 12 .2 13 6. 557377 559 65.95 73.84 11.96360 15 .7 16.7 6. 369426 570 24.4 23.78 -2.54098 12 .4 12.4 0 574 69.03 76.16 10.32884 16 .3 17.5 7. 361963 576 77.96 77.28 -0.87224 16 .4 17.2 4. 878048 579 79.24 87.15 9.982332 16 .3 17.5 7. 361963 587 23.78 22.53 -5.25651 12 .3 12.3 0 588 24.18 22.75 -5.91397 12 .3 12.1 -1 .62601 592 25.48 25.48 0 12 .9 12.7 -1 .55038 44.489 47.291 5.045876 14. 01 14.47 2. 935237 Sample 6, Jan. 29, 1988 Tag i n i t i a l sample % d i f f . I n i t i a l Sample % chg. 255 28.9 37.72 30 .51903 12.2 13 6. 557377 503 25.02 34.65 38 .48920 11.9 12.6 5. 882352 504 36.65 55.69 51 .95088 13.3 14.6 9. 774436 509 53.98 80.27 48 .70322 14.8 16.5 11 .48648 534 69.11 90.03 30 .27058 16.5 17.7 7. 272727 558 55.65 60.12 8. 032345 13.8 14 1. 449275 565 74.23 93.46 25 .90596 15. 9 17.7 11 .32075 576 77.96 87.67 12 .45510 16.4 17.7 7. 926829 584 50.49 66.91 32 .52129 14.3 16 11 .88811 587 23.78 22.25 -6 .43397 12.3 12.3 0 49.577 62.877 27 .24136 14.14 15.21 7. 355835 90 Group IE, I n i t i a l Values Dec. 6, 1987 Tag Weight Length New Tag 219 19.91 11.1 220 26.8 12.9 508 221 19.41 11 229 11.38 10.1 252 30.08 12.6 253 44.89 14.3 256 24.65 12.1 265 44.4 14.9 502,524 266 35.25 14.2 269 30.38 13.5 270 63.7 15.5 271 36.12 14.5 272 73.8 15.8 273 26.98 12.9 274 66.68 16.4 275 61.92 16 276 29.23 12.8 277 23.09 12.4 278 48.54 15 279 39.81 13.6 280 56.38 15.1 283 28.22 12.8 300 43.1 14.5 302 18.64 11.1 303 24.92 13.1 449 304 35.24 13.4 305 20.9 11.8 306 40.08 14.1 406 310 21.4 11.8 311 31.29 13.9 313 28.55 12.4 314 20.18 11.8 316 41.41 14.6 326 36.52 13.6 355 75.4 16.8 356 25.98 12.9 357 23.59 12.4 359 44.75 14.3 360 64.02 16 421 362 40.95 13.6 count 40 avg 36.9635 13.54 sum 1437.59 feed 71.8795 g/day 91 Group IE, Samples Weight Dec. 25, 1987: Sample 1 Tag Length Tag Tag I n i t i a l Sample % I n i t i a l Sample % chg. 271 36.12 34.02 -5.81395 14.5 14.3 -1.37931 277 23.09 22.25 -3.63793 12.4 12.3 -0.80645 283 28.22 27.95 -0.95676 12.8 13.1 2.34375 300 43.1 42.48 -1.43851 14.5 14.4 -0.68965 310 21.4 20.18 -5.70093 11.8 11.6 -1.69491 316 41.41 40.38 -2.48732 14.6 14.5 -0.68493 326 36.52 39.44 7.995618 13.6 14.3 5.147058 355 75.4 89.05 18.10344 16.8 18.2 8.333333 502 44.4 44.85 1.013513 14.9 14.7 -1.34228 508 26.8 24.29 -9.36567 12.9 12.8 -0.77519 37.646 38.489 -0.22885 13.88 14.02 0.845140 1/ : 1988: Sample 2 I n i t i a l Sample % I n i t i a l Sample % chg. 253 44.89 50.71 12.96502 14.3 15.5 8.391608 256 24.65 26.22 6.369168 12.1 12.8 5.785123 272 73.8 79.95 8.333333 15.8 17.1 8.227848 273 26.98 25.32 -6.15270 12.9 12.8 -0.77519 275 61.92 68.82 11.14341 16 16.7 4.375 283 28.22 26.94 -4.53579 12.8 13.1 2.34375 304 35.24 37.59 6.668558 13.4 14.1 5.223880 316 41.41 39.93 -3.57401 14.6 14.4 -1.36986 357 23.59 22.38 -5.12929 12.4 12 -3.22580 406 40.08 41.46 3.443113 14.1 14.3 1.418439 40.078 41.932 2.953080 13.84 14.28 3.039478 8, : 1988: Sample 3 I n i t i a l Sample % I n i t i a l Sample % chg. 256 24.65 26.09 5.841784 12.1 12.7 4.958677 270 63.7 68.71 7.864992 15.5 16.7 7.741935 278 48.54 55.64 14.62711 15 16 6.666666 304 35.24 37 4.994324 13.4 14.3 6.716417 311 31.29 30.3 -3.16395 13.9 13.4 -3.59712 355 75.4 84.6 12.20159 16.8 18.3 8.928571 357 23.59 22.36 -5.21407 12.4 12.2 -1.61290 362 40.95 47.52 16.04395 13.6 14.7 8.088235 406 40.08 41.41 3.318363 14.1 14.4 2.127659 421 64.02 62.59 -2.23367 16 16.2 1.25 44.746 47.622 5.428042 14.28 14.89 4.126813 92 Jan. 15, 1988: Sample 4 Tag I n i t i a l Sample % I n i t i a l Sample % chg. 269 30.38 29.3 -3.55497 13.5 13.4 -0.74074 270 63.7 75.32 18.24175 15.5 17.1 10.32258 274 66.68 83.28 24.89502 16.4 17.8 8.536585 277 23.09 23.22 0.563014 12.4 12.3 -0.80645 283 28.22 30 6.307583 12.8 13.2 3.125 302 18.64 22.07 18.40128 11.1 11.4 2.702702 304 35.24 40.88 16.00454 13.4 14.3 6.716417 311 31.29 30.11 -3.77117 13.6 13.3 -2.20588 326 36.52 41.89 14.70427 13.6 14.7 8.088235 406 40.08 41.89 4.515968 14.1 14.5 2.836879 37.384 41.796 9.630730 13.64 14.2 3.857532 Jan. 22, 1988: Sample 5 Tag I n i t i a l Sample % I n i t i a l Sample % chg. 271 36.12 35.81 -0.85825 14.5 14.6 0.689655 272 73.8 102.09 38.33333 15.8 17.9 13.29113 274 66.68 87.22 30.80383 16.4 18.3 18.3 283 28.22 33.86 19.98582 12.8 13.4 13.4 302 18.64 26.42 41.73819 11.1 11.7 5.405405 304 35.24 47.18 33.88195 13.4 14.9 11.19402 311 31.29 30.31 -3.13199 13.9 13.5 -2.87769 359 44.75 66.08 47.66480 14.3 16 11.88811 449 24.92 33.98 36.35634 13.1 14.3 9.160305 467 11.38 11.14 -2.10896 10.1 10 -0.99009 37.104 47.409 24.26650 13.54 14.46 7.946085 Jan. 29, 1988: Sample 6 Tag I n i t i a l Sample % I n i t i a l Sample % chg. 269 30.38 28.92 -4.80579 13.5 13.5 0 271 36.12 38.01 5.232558 14.5 14.5 0 274 66.68 98.08 47.09058 16.4 18.5 12.80487 277 23.09 28.53 23.55998 12.4 12.6 1.612903 283 28.22 37.12 31.53791 12.8 13.5 5.46875 311 31.29 30.37 -2.94023 13.9 13.5 -2.87769 316 41.41 41.61 0.482975 14.6 14.6 0 357 23.59 23.88 1.229334 12.4 12.4 0 467 11.38 11.09 -2.54833 10.1 10.2 0.990099 502 44.4 67.31 51.59909 14.9 15.8 6.040268 33.656 40.492 15.04380 13.55 13.91 2.403920 93 Group IF, I n i t i a l Values Dec. 17, 1987 Tag Weight Length New Tag 505 39 13.9 507 29.75 13 509 16.45 10.9 510 33.52 13.5 511 38.65 13.7 512 51.24 15.5 513 44.58 14.2 514 29.361 14.1 516 39.02 14.6 517 83.6 17.3 518 40.64 14.3 519 40.12 13.3 525 520 64.66 15.7 526 29.31 12.8 527 27.5 12.5 529 48.35 15 530 106.54 18.9 531 19.55 11.8 413 532 58.65 15.8 424 533 39.92 14.2 536 88.28 17.9 537 26.42 12.3 417 538 16.08 10.3 539 34.48 13.4 540 31.82 13.3 541 43.95 14.6 542 34.46 12.6 543 20.61 11.6 544 58.98 15.5 545 46.7 14.1 546 18.3 11.3 547 54.59 15.8 548 22.08 12 549 67.71 16.5 550 56.75 14.7 551 110.28 19.1 552 21.8 11.9 553 42.29 14.8 554 51.38 14 555 35.69 13.7 556 28 12.4 557 31.32 12.7 578 71.68 16.3 580 35.3 13.7 581 17.6 10.3 94 Group IF, Samples Weight Dec. 25, 1987: Sample 1 Tag Length Tag Tag I n i t i a l Sample % change I n i t i a l Sample % chg. 424 58.65 54.01 -7.91133 11 15 .9 44.54545 507 29.75 29.5 -0.84033 13 12 .9 -0.76923 509 16.45 16.8 2.12.7659 10.9 10 .9 0 513 44.58 38.42 -13.8178 14.2 14 .2 0 516 39.02 38.92 -0.25627 14.6 14 .4 -1.36986 519 40.12 34.68 -13.5593 13.3 13 .6 2.255639 526 29.31 28.01 -4.43534 12.8 12 .9 0.78125 546 18.3 18.06 -1.31147 11.3 11 .1 -1.76991 551 110.28 98.15 -10.9992 19.1 18 .5 -3.14136 557 31.32 28.06 -10.4086 12.7 12 .8 0.787401 41.778 38.461 -6.14122 13.29 13. 72 4.131937 1/ : 1988: Sample 2 I n i t i a l Sample % I n i t i a l Sample % chg. 417 26.42 25.09 -5.03406 12.3 12 .3 0 507 29.75 29 -2.52100 13 13 0 509 16.45 15.65 -4.86322 10.9 10 .8 -0.91743 511 38.65 29.8 -22.8978 13.7 13 .5 -1.45985 512 51.24 45.55 -11.1046 15.5 15 .3 -1.29032 527 27.5 24.42 -11.2 12.5 12 .4 -0.8 542 34.46 29.4 -14.6836 12.6 12 .3 -2.38095 546 18.3 17.85 -2.45901 11.3 11 .2 -0.88495 552 21.8 20.55 -5.73394 11.9 11 .8 -0.84033 555 35. 69 33.81 -5.26758 13.7 13 .4 -2.18978 30.026 27.112 -8.57649 12.74 12 .6 -1.07636 8, 1988: Sample 3 I n i t i a l Sample % I n i t i a l Sample % chg. 413 19.55 18.92 -3.22250 11.8 11 .6 -1.69491 424 58.65 52.88 -9.83802 15.8 15 .9 0.632911 513 44.58 37.09 -16.8012 14.2 14 .2 0 516 39.02 39.03 0.025627 14.6 14 .5 -0.68493 518 40.64 39.38 -3.10039 14.3 14 .3 0 520 64.66 54.9 -15.0943 15.7 15 .6 -0.63694 527 27.5 24.51 -10.8727 12.5 12 .5 0 541 43.95 38.49 -12.4232 14.6 14 .7 0.684931 546 18.3 17.6 -3.82513 11.3 11 .1 -1.76991 553 42.29 40.82 -3.47599 14.8 14 .6 -1.35135 39.914 36.362 -7.86279 13.96 13 .9 -0.48202 95 Jan. 15, 1988: Sample 4 Tag I n i t i a l Sample % I n i t i a l Sample % chg. 413 19.55 18.46 -5.57544 11.8 11.6 -1.69491 505 39 37.78 -3.12820 13.9 13.9 0 507 29.75 28.62 -3.79831 13 13.9 6.923076 516 39.02 39.5 1.230138 14.6 14.7 0.684931 517 83.6 84.12 0.622009 17.3 17.4 0.578034 520 64.66 55.69 -13.8725 15.7 16 1.910828 527 27.5 24.89 -9.49090 12.5 12.6 0.8 530 106.54 97.28 -8.69157 18.9 19.3 2.116402 544 58.98 59.66 1.152933 15.5 15.2 -1.93548 546 18.36 17.35 -5.50108 11.3 11.2 -0.88495 48.696 46.335 -4.70530 14.45 14.58 0.849791 Jan. 22, 1988: Sample 5 Tag I n i t i a l Sample % I n i t i a l Sample % chg. 507 29.75 28.02 -5.81512 13 12.9 -0.76923 509 16.45 15.15 -7.90273 10.9 10.7 -1.83486 513 44.58 48.61 9.039928 14.2 14.7 3.521126 516 39.02 38.95 -0.17939 14.6 14.7 0.684931 542 34.46 31.08 -9.80847 12.6 12.5 -0.79365 546 18.3 17.88 -2.29508 11.3 11.2 -0.88495 548 22.08 22.28 0.905797 12 11.9 -0.83333 551 110.28 109.28 -0.90678 19.1 19.2 0.523560 553 42.29 40.58 -4.04350 13.7 13.9 1.459854 555 35.69 37.55 5.211543 12.7 13.3 4.724409 557 31.32 37.71 20.40229 39.29 38.938 -1.57938 13.41 13.5 0.579784 Jan. 2 9, 1988: Sample 6 Tag I n i t i a l Sample % I n i t i a l Sample % chg. 413 19.55 20.43 4.501278 11.8 11.8 0 507 29.75 27.58 -7.29411 13 13 0 514 29.61 62.98 112.6984 14.1 15.5 18.3 520 64.66 70.39 8.861738 15.7 16.5 13.4 525 40.12 45.9 14.40677 13.3 13.9 4.511278 527 27.5 29.89 8.690909 12.5 12.8 2.4 533 39.92 58.81 47.31963 14.2 15.1 6.338028 543 20.61 21.72 5.385735 11.6 11.9 2.586206 554 51.38 69.11 34.50759 14 15.5 10.71428 555 35.69 42.82 19.97758 13.7 14 2.189781 35.879 44.963 24.90555 13.39 14 6.043957 96 Group 2A: I n i t i a l Values July 19, 1988 Tag Weight Length 650 123.68 19.4 651 136.81 20.1 610 66.91 15.7 652 151.62 20.2 653 48.12 14.3 654 97.59 17.5 655 100.01 18.8 656 163.8 20.7 657 134.52 20.8 658 109.22 18.5 659 126.22 19.6 660 56.92 15 661 97.63 18.2 662 147.68 20.7 664 181.35 22.5 665 184.72 22.2 666 115.5 19.4 667 179.18 21.4 668 189.11 21.7 669 71.59 15.7 670 120.05 19.2 671 128.78 20.2 672 108.75 18.2 673 130.6 20.1 674 107.57 18.5 675 81.95 16.9 676 76.47 16.5 678 68.77 15.4 679 142.6 20.3 680 128.98 19.7 681 72.65 17 682 132.24 19.6 683 104.82 18.7 684 135.05 19.7 685 139.38 19.9 686 49.72 14.5 687 115.3 18.8 688 93.98 17.1 689 169.92 21.5 690 143.57 20.3 691 76.61 16.6 692 112.18 19.4 693 173.22 20.9 694 120.68 19.8 695 106.15 18.2 696 113.47 19.2 97 Tag Weight Length 697 69.72 16.5 698 100.11 18.2 699 61.12 15.6 700 130.5 19.4 701 127.78 19.8 702 165.88 21.2 avg: 116.6052 18.796 sum: 5830.26 feed (5%) 291.513 98 Group 2B: I n i t i a l Values July 19, 1988 Tag Weight 600 115. 23 602 159. 11 603 119. 26 604 118 i . l 605 75. 33 606 120. 17 607 165 .6 608 78. 51 609 183. 65 611 50. 24 612 95. 42 613 123. 52 614 117. 75 615 49 1.2 617 125. 35 618 146. 45 619 137 .5 620 112. 35 621 174. 09 622 144. 05 623 115. 68 625 155. 77 626 120. 21 627 125. 45 628 131. 05 629 126. 83 630 110. 63 631 171. 16 632 42 634 143. 95 635 106. 73 636 106 .1 637 107. 25 638 123. 21 639 97. 17 640 93. 02 641 100. 52 642 133. 25 643 109. 64 644 113. 31 645 103. 23 646 183. 01 647 130. 62 648 112. 72 649 168. 18 Length 18.7 21.4 19.3 19.3 16.7 19.1 21.5 16.5 22.5 14.9 17.5 19.7 19.1 13.4 19.9 20.4 20.5 17.6 21.3 20.1 18.6 20.2 19.1 18.8 19.8 18.9 18.5 21.8 13.3 20.6 18.2 18.8 18.6 19.3 17.6 17.6 17.8 20 18 19.8 18 22.5 19.3 18.7 21.2 99 Tag Weight Length 750 124.81 19.3 751 162.59 21.1 752 157.83 21.6 753 160.98 21.4 754 143.28 19.5 avg: 123.8212 19.146 t o t a l : 6191.06 feed (5%) 309.553 100 Experiment 2: Sample 1 August 10, 1988 Group 2B Weight l e n g t h Tag # I n i t i a l Sample % change i n i t i a l sample % d i f f 604 118.1 108.48 -8.14563 19.3 19.2 -0.51813 605 75.33 59.78 -20.6425 16.7 16.8 0.598802 606 120.17 109.58 -8.81251 19.1 19.3 1.047120 608 78.51 57.31 -27.0029 16.5 17 3.030303 613 123.52 111.08 -10.0712 19.7 19.5 -1.01522 614 117.75 107.35 -8.83227 19.1 19.2 0.523560 618 146.45 132.15 -9.76442 20.4 20.2 -0.98039 621 174.09 151.91 -12.7405 21.3 21.1 -0.93896 636 106.1 98.58 -7.08765 18.8 18.8 0 637 107.25 97.61 -8.98834 18.6 18.7 0.537634 639 97.17 88.29 -9.13862 17.6 17.6 0 640 93.02 85.42 -8.17028 17.6 17.8 1.136363 646 183.01 160.05 -12.5457 22.5 22.4 -0.44444 668 189.11 171.65 -9.23272 21.7 21.8 0.460829 751 162.59 150.7 -7.31287 21.1 21 -0.47393 avg 126.1446 112.6626 -11.2325 19.33333 19.36 0.197567 Group 2A weight l e n g t h Tag # I n i t i a l Sample % change i n i t i a l sample % d i f f 655 100.01 92.83 -7.17928 18.8 18.5 -1.59574 657 134.52 124.25 -7.63455 20.8 20.6 -0.96153 662 147.68 244.53 65.58098 20.7 22.8 10.14492 666 115.5 117.65 1.861471 19.4 19.7 1.546391 670 120.05 104.77 -12.7280 19.2 18.9 -1.5625 674 107.57 165.2 53.57441 18.5 20 8.108108 676 76.47 93.46 22.21786 16.5 17 3.030303 678 68.77 60.08 -12.6363 15.4 15.5 0.649350 685 139.38 199.24 42.94733 19.9 21.1 6.030150 689 169.92 242.13 42.49646 21.5 22.8 6.046511 690 143.57 200.54 39.68099 20.3 21.8 7.389162 692 112.18 101.91 -9.15492 19.4 19.4 0 693 173.22 155.42 -10.2759 20.9 20.7 -0.95693 694 120.68 213.61 77.00530 19.8 21.6 9.090909 695 106.15 95.25 -10.2684 18.2 18.5 1.648351 701 127.78 117.81 -7.80247 19.8 19.6 -1.01010 avg 122.7156 145.5425 16.73030 19.31875 19.90625 2.974834 101 Group 2A: sample 2 August 31, 1988 tag i n i t . wt sample wt% d i f i n i t l n . sample l n % d i f f 652 151. 62 305. 77 101.6686 20 .2 24 .4 20.79207 653 48. 12 57. 85 20.22028 14 .3 14 .9 4.195804 656 163 .8 216. 82 32.36874 20 .7 22 .7 9.661835 657 134. 52 89. 51 -33.4597 20 .8 17 .2 -17.3076 660 109. 22 69. 48 -36.3852 18 .5 15 .8 -14.5945 661 97. 63 99. 62 2.038307 18 .2 17 .7 -2.74725 664 181. 35 355 .7 96.14006 22 .5 26 .7 18.66666 666 115 .5 175. 82 52.22510 19 .4 21 .2 9.278350 670 120. 05 106. 32 -11.4369 19 .2 18 .9 -1.5625 671 128. 78 219. 31 70.29818 20 .2 22 .9 13.36633 672 108. 75 229 '.8 111.3103 18 .2 22 20.87912 675 81. 95 89. 51 9.225137 16 .9 17 .2 1.775147 676 76. 47 116. 09 51.81116 16 .5 18 9.090909 677 130. 32 149. 55 14.75598 20 .4 21 .1 3.431372 678 68. 77 78. 82 14.61393 15 .4 16 .8 9.090909 680 128. 98 117. 33 -9.03240 19 .7 19 .6 -0.50761 681 72. 65 165. 58 127.9146 17 21 .5 26.47058 682 132. 24 128. 28 -2.99455 19 .6 19 .7 0.510204 683 104. 82 162. 91 55.41881 18 .7 20 .3 8.556149 685 139. 38 257. 28 84.58889 19 .9 23 .2 16.58291 686 49. 72 48. 44 -2.57441 14 .5 14 .5 0 687 115 .3 109. 51 -5.02168 18 18 .8 4.444444 688 93. 98 110. 62 17.70589 17 .1 18 .1 5.847953 689 169. 92 330. 38 94.43267 21 .5 24 .9 15.81395 691 76. 61 99. 62 30.03524 16 .6 17 .7 6.626506 692 112. 18 112. 48 0.267427 19 .4 19 .5 0.515463 694 120. 68 304. 68 152.4693 19 .8 24 .3 22.72727 697 69. 72 83. 28 19.44922 16 .5 17 3.030303 698 100. 11 187. 58 87.37388 18 .2 21 .2 16.48351 110.4531 157. 86 39.49748 18.54827 19.92413 7.279936 102 Group 2B: sample 2 August 31, 1988 tag i n i t . wt sample wt% d i f i n i t l n . sample l n % d i f f 600 115. 23 172. 11 49.36214 18 .7 20 .8 11.22994 602 159. 11 192. 88 21.22431 21 .4 22 .6 5.607476 603 119. 26 165. 58 38.83951 19 .3 21 .1 9.326424 606 120. 17 142. 09 18.24082 19 .1 20 .2 5.759162 607 165 .6 247. 35 49.36594 21 .5 24 11.62790 609 183. 65 160 .2 -12.7688 22 .5 22 .2 -1.33333 614 117. 75 158 .3 34.43736 19 .1 20 .5 7.329842 615 49 .2 56. 16 14.14634 13 .4 14 .3 6.716417 617 125. 35 176. 35 40.68607 19 .9 21 .9 10.05025 620 112. 35 148. 38 32.06942 17 .6 19 .1 8.522727 622 144. 05 125. 05 -13.1898 21 .1 19 .9 -5.68720 623 115. 68 108. 08 -6.56984 18 .6 18 .6 0 626 120. 21 169.06 40.63721 19 .1 20 .8 8.900523 627 125. 45 184. 98 47.45316 18 .8 20 .9 11.17021 629 126. 83 186. 77 47.26011 18 .9 21 .1 11.64021 631 171. 16 180. 21 5.287450 21 .8 21 .9 0.458715 634 143. 95 129. 78 -9.84369 20 .6 20 .5 -0.48543 635 106. 73 133. 58 25.15693 18 .2 19 .3 6.043956 637 107. 25 142 .4 32.77389 18 .6 20 .2 8.602150 638 123. 21 156. 35 26.89716 19 .3 21 .4 10.88082 640 93. 02 150. 28 61.55665 17 .6 19 .8 12.5 641 100. 52 104. 21 3.670911 17 .8 18 .1 1.685393 644 113. 31 163. 87 44.62095 19 .8 21 .4 8.080808 645 103. 23 154. 08 49.25893 18 19 .3 7.222222 646 183. 01 244 .3 33.48997 22 .5 24 .4 8.444444 648 112. 78 114. 08 1.152686 18 .7 18 .7 0 628 131. 05 164. 72 25.69248 19 .8 21 .4 8.080808 750 124. 81 157. 31 26.03958 19 .3 20 .5 6.217616 751 162. 59 201. 35 23.83910 21 .1 22 .2 5.213270 752 157. 83 227. 66 44.24380 21 .6 23 .5 8.796296 753 160. 98 273. 85 70.11429 21 .4 23 .9 11.68224 Avg 128.8812 164.2377 27.90790 19.51935 20.79032 6.589802 i n i t i a l % change sample t o t a l r a t i o n r a t i o n (5%) 6191.06 27.9079 7918.854 395.9427 1 0 3 Experiment 2: Sample 3 Sept. 21, 1988 Group 2A Weight Length Tag # i n i t i a l sample % change i n i t i a l sample % change 654 97.59 246.87 152.9664 17.5 21.7 24 661 97.63 227.02 132.5309 18.2 23 26.37362 666 115.5 252.63 118.7272 19.4 23.2 19.58762 676 76.47 145.72 90.55838 16.5 19.2 16.36363 679 142.6 244 71.10799 20.3 22.7 11.82266 681 72. 65 135.81 86.93737 17 19.6 15.29411 685 139.38 336.49 141.4191 19.9 24.4 22.61306 691 76.61 118.79 55.05808 16.6 18.3 10.24096 694 120.68 405.71 236.1866 19.8 26.4 33.33333 696 113.47 136.25 20.07579 19.2 19.6 2.083333 699 61.12 108.14 76.93062 15.6 17.2 10.25641 700 130.5 444.98 240.9808 19.4 26.1 34.53608 Avg. 103.6833 233.5341 118.6233 18.28333 21.78333 18.87540 Group 2B Weight Length Tag # i n i t i a l sample % change i n i t i a l sample % change 617 125.35 159.37 27.14000 19.9 21.9 10.05025 622 144.05 126.25 -12.3568 20.1 20.1 0 625 155.77 215.8 38.53758 20.2 22.7 12.37623 627 125.45 158.78 26.56835 18.8 20.6 9.574468 628 131.05 162.91 24.31133 19.8 21.2 7.070707 629 126.83 155.57 22.66025 18.9 20.7 9.523809 630 110.63 125.58 13.51351 18.5 19.6 5.945945 635 106.73 117.08 9.697367 18.2 19.4 6.593406 641 100.52 99.78 -0.73617 17.8 18.3 2.808988 751 162.59 164 0.867212 21.1 22.3 5.687203 752 157.83 217.81 38.00291 21.6 23.6 9.259259 757 136.31 19.1 Avg. 131.5272 154.8118 17.10959 19.53636 20.94545 7.171843 Feeding Amounts i n i t i a l t o t a l feed/day (g) Tank 5 (5%) 5830.26 12746.30 637.3153 Tank 7 (5%) 6191.06 7250.324 362.5162 104 Experiment 2: Sample 4 Oct. 12, 1988 Group 2A Weight Length # i n i t i a l sample % change i n i t i a l sample % < change 654 97.59 314 .88 222.6560 17 .5 24. 1 37 .71428 655 100.01 151 .16 51.14488 18 .8 21 11 .70212 661 97.63 327 .15 235.0916 18 .2 25. 3 39 .01098 662 147.68 483 .52 227.4106 20 .7 28. 4 37 .19806 666 115.5 313 .58 171.4978 19 .4 25. 3 30 .41237 669 71.59 157 .33 119.7653 15 .7 19. 8 26 .11464 670 120.05 210 .61 75.43523 19 .2 22. 1 15 .10416 680 128.98 234 .91 82.12901 19 .7 22. 6 14 .72081 681 72.65 220 .65 203.7164 17 22. 3 31 .17647 683 104.82 338 .38 222.8200 18 .7 25. 7 37 .43315 687 115.3 167 .47 45.24718 18 .8 21. 1 12 .23404 688 93.98 204 .05 117.1206 17 .1 20. 6 20 .46783 692 112.18 271 .69 142.1911 19 .4 24. 2 24 .74226 697 69.72 186 .54 167.5559 16 .5 20. 9 26 .66666 698 100.11 316 .65 216.3020 18 .2 24. 8 36 .26373 700 130.5 578 .28 343.1264 19 .4 28. 9 48 .96907 Average 104.8931 279.8031 165.2006 18.39375 23.56875 28.12066 S.D. 21.25444 114.2811 78.43601 1.286209 2.646157 10.73070 105 Group 2B Weight Length Tag # i n i t i a l sample % change i n i t i a l sample % change 612 95.42 201.65 111.3288 17.5 21.3 21.71428 614 117.75 229.78 95.14225 19.1 22.3 16.75392 625 155.77 361.22 131.8931 20.2 25.1 24.25742 629 126.83 255.6 101.5296 18.9 22.8 20.63492 630 110.63 201.63 82.25616 18.5 21.6 16.75675 631 171.16 318.61 86.14746 21.8 25 14.67889 635 106.73 201.45 88.74730 18.2 21.2 16.48351 640 93.02 253.05 172.0382 17.6 23 30.68181 641 100.52 165.73 64.87266 17.8 20.3 14.04494 642 133.25 259.98 95.10694 20.1 23.1 14.92537 643 109.64 209.65 91.21670 18.1 21.2 17.12707 644 113.31 237.98 110.0255 19.8 23.6 19.19191 750 124.81 239.03 91.51510 19.3 22.7 17.61658 751 162.59 280.91 72.77200 21.1 23.9 13.27014 752 157.83 379.05 140.1634 21.6 26.1 20.83333 757 219.35 20.8 Average 125.284 250.9168 102.3170 19.30666 22.75 18.59806 S.D. 24.63195 57.11309 26.80272 1.374756 1.625576 4.394011 Feeding i n i t i a l % change T o t a l feed/day (g) Tank 5 5830.26 165.2 15461.84 773.0924 Tank 7 S t a r v a t i o n P e r i o d 106 Experiment 2: Sample 5 November 2, 1988 Group 2A Weight Length Tag # i n i t i a l sample % change i n i t i a l sample % change 655 100. 01 173 .37 73. 35266 18 .8 22. 4 19 .14893 659 126. 22 253 .05 100 .4832 19 .6 24. 4 24 .48979 661 97. 63 405 .77 315 .6201 18 .2 27. 6 51 .64835 666 115 .5 383 .55 232 .0779 19 .4 27. 2 40 .20618 667 179. 18 586 .28 227 .2016 21 .4 29. 8 39 .25233 671 128. 78 463 .44 259 .8695 20 .2 29. 4 45 .54455 679 142 .6 428 .88 200 .7573 20 .3 27. 7 36 .45320 680 128. 98 291 .21 125 .7791 19 .7 25. 1 27 .41116 681 72. 65 286 .31 294 .0949 17 25. 3 48 .82352 687 115 .3 183 .91 59. 50563 18 .8 22. 2 18 .08510 688 93. 98 258 .87 175 .4522 17 .1 22. 3 30 .40935 689 169. 92 573 .35 237 .4234 21 .5 31. 1 44 .65116 694 120. 68 669 .85 455 .0629 19 .8 31. 4 58 .58585 696 113. 47 260 .15 129 .2676 19 .2 23. 5 22 .39583 697 69. 72 229 .02 228 .4853 16 .5 23. 1 40 Average 118.308 363.134 207.6289 19.16666 26.16666 36.47369 S.D. 29.54218 149.5474 100.2556 1.437435 3.133191 11.98323 107 Group 2B Weight Length Tag # i n i t i a l sample % change i n i t i a l sample % change 605 75. 33 135 .14 79. 39731 16. 37 20 .4 24 .61820 614 117. 75 191 .06 62. 25902 19 .1 22 .8 19 .37172 619 137 .5 235 .51 71.28 20 .5 24 .6 20 627 125. 45 225 .68 79. 89637 18 .8 23 .1 22 .87234 629 126. 83 213 .21 68. 10691 18 .9 23 21 .69312 630 110. 63 176 .08 59. 16116 18 .5 21 .9 18 .37837 631 171. 16 272 .55 59. 23697 21 .8 25 .4 16 .51376 632 42 59 .25 41. 07142 13 .3 15 .4 15 .78947 639 97. 17 137 .62 41. 62807 17 .6 20 .3 15 .34090 642 133. 25 210 .87 58. 25140 20 .1 23 .4 16 .41791 644 113. 31 203 .38 79. 48989 19 .8 23 .9 20 .70707 645 103. 23 207 .38 100 .8912 18 .1 22 .6 24 .86187 648 112. 72 153 .81 36. 45315 18 .7 21 12 .29946 751 162. 59 225 .81 38. 88308 21 .1 24 .2 14 .69194 752 157. 83 308 .31 95. 34309 21 .6 26 .4 22 .22222 757 182 .27 20 .9 Average 119.1166 196.1206 64.75660 18.95133 22.45625 19.05189 S.D. 32.12609 56.29581 19.37769 2.092070 2.489972 3.678371 Feeding i n i t i a l % change T o t a l feed/day (g) Tank 5 5830.26 207.6 17933.87 896.6939 Tank 7 6191.06 64.8 10202.86 510.1433 108 Experiment 2: Sample 6 November 23, 1988 Group A Weight Length • # i n i t i a l sample % change i n i t i a l sample % * change 656 163.8 402.52 145 .7387 20.7 27.6 33 .33333 662 147.68 534.92 262 .2156 20.7 31 49 .75845 666 115.5 415.19 259 .4718 19.4 28.5 46 .90721 670 120.05 329.11 174 .1441 19.2 25.4 32 .29166 671 128.71 504.98 292 .3393 20.2 30.3 50 676 76.47 313.45 309 .8993 16.5 24.6 49 .09090 679 142.6 471.91 230 .9326 20.3 28.3 39 .40886 680 128.98 314.38 143 .7432 19.7 25.3 28 .42639 681 72.65 344.91 374 .7556 17 26.4 55 .29411 685 139.38 586.31 320 .6557 19.9 29.2 46 .73366 687 115.3 202.32 75. 47267 18.38 22.5 22 .41566 688 93.98 290.31 208 .9061 17.1 22.8 33 .33333 689 169.92 606.79 257 .1033 21.5 31.6 46 .97674 694 120.68 710.39 488 .6559 19.8 32.1 62 .12121 697 69.72 261.39 274 .9139 16.5 24.4 47 .87878 age 120.3613 419.2586 254 .5965 19.12533 27.33333 42 .93135 1. 30.11348 140.4963 97. 95022 1.582377 3.030438 10 .54467 109 Group B Weight Length Tag # i n i t i a l sample % change i n i t i a l sample % change 605 75. 33 182. 31 142.0151 16.7 21 .7 29. 94011 614 117. 75 272. 58 131.4904 19.1 24 25. 65445 617 125. 35 318. 79 154.3199 19.9 25 .7 29. 14572 619 137 .5 319. 68 132.4945 20.5 25 .6 24. 87804 627 125. 45 301. 11 140.0239 18.8 23 .9 27. 12765 628 131. 05 282. 11 115.2689 19.8 23 .8 20. 20202 629 126. 83 318. 51 151.1314 18.9 24 .4 29. 10052 631 171. 16 330. 67 93.19350 21.8 26 .9 23. 39449 635 106. 73 248. 68 132.9991 18.2 22 .9 25. 82417 636 106 ; . i 174. 68 64.63713 18.8 21 .5 14. 36170 639 97. 17 187. 55 93.01224 17.6 21 .3 21. 02272 641 100. 52 195. 18 94.17031 17.8 21 .5 20. 78651 642 133. 25 263. 91 98.05628 20 24 .5 22.5 644 113. 31 273. 28 141.1790 19.8 25 .3 27. 77777 648 112. 72 216. 29 91.88254 18.7 22 .1 18. 18181 750 124. 81 263. 78 111.3452 19.3 24 .1 24. 87046 752 157. 83 428. 11 171.2475 21.6 27 .5 27. 31481 753 160. 98 455. 13 182.7245 21 28 .8 37. 14285 757 112. 35 235. 11 109.2656 17.6 21 .9 24. 43181 Avg 122.9573 277.2347 123.7082 19.25789 24.07368 24. 92935 S.D. 22.61857 74.01842 29.82537 1.342548 2.121529 4.866129 Feeding i n i t i a l % change T o t a l feed/day (g) Tank 5 5830.26 254.6 20674.10 1033.705 Tank 7 6191.06 123.7 13849.40 692.4700 (Star v a t i o n ) 110 

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