Protein, lipid, water and caloric contents of immature rainbow trout, Salmo gairdneri Richardson, growing at different rates

Protein, lipid, water and caloric contents of immature rainbow trout, relative to size of whole fish, growing at different rates were examined by use of allometric analysis (y = ax^b, Huxley, 1932). Fish grew at different rates as a result of differences in ration size (satiation, or 4–5% of dry body weight), temperature (7 and 12°C) and bGH (bovine growth hormone) administration. In fingerlings, protein, lipid and caloric contents tended to increase (v. body weight) as a percentage of body composition, whereas above fingerling size, protein decreased while lipid and caloric contents still increased. These trends occurred regardless of growth rate differences. The correlations between protein, lipid, caloric contents and body weight were high so reliable estimates of body components can be made from body weight for all experimental treatments. At satiation rations (7 and 12°C), there were no significant differences in protein content, but lipid and caloric contents were significantly higher in control fish. At low rations, protein and caloric contents were lower than those at satiation rations and lipid was lower than in the control group. On a dry weight basis, in uninjected fish at 12°C, ration size did not influence the percentage composition (protein and lipid) but the low ration group had lower energy values per unit of body dry weight. At low temperature (7°C satiation), fish had lower lipid and higher protein content (dry weight basis) than their controls (12°C satiation), thereby resembling bGH injected fish at satiation rations. Values of caloric content estimated from protein and lipid values by use of standard conversion factors differed sufficiently from caloric values directly determined by bomb calorimetry to suggest that caloric values of lipids may change during growth. The increase in body caloric content during growth apparently results mainly from an increase in the percentage of lipid. The similarity of body composition between different growth rate groups implied a tendency for conservation of relative proportions of components such that body composition can be approximated from body weight. Equations were also given to describe the relationship between water content and body constituents, and the relationship between condition factor (K) and both the body dry weight and lipid content.     

Specific growth rate and proximate body composition of Atlantic cod (Gadus morhua L.)

The effect of growth rate and maturation on the proximate composition and energy content of Atlantic cod, Gadus morhua L., was investigated over 10 months for each of two consecutive years, 1978–1980 at 5 and 8 °C. Relative energy and lipid content of whole cod increased with specific growth rate for all three sampling periods (November, January, March), each at 5 and 8 °C. Relative water content decreased with specific growth rate and temperature, and was lower in March than in January and November. Relative protein content was positively correlated with specific growth rate, but to a lesser degree than with temperature and age. Relative ash content was negatively correlated with specific growth rate. The effect of season and temperature on the proximate content of gonad, liver, muscle, and carcass was also determined. The major energy and lipid source in cod was the liver. Energy, lipid, and water were highly correlated to each other, and regressions are provided to allow for their prediction, given one of the components. Energy budgets for cod at 5 and 8 °C are calculated and the effect of increased ration size on the budget is estimated. The prediction of short-term specific growth rates of cod from the proximate composition is proposed. The proximate composition of cod is affected by growth rate and thus feeding level, and in turn directly affects behaviour. The relative proximate content of maturing and immature 3-yr-old cod was not found to be significantly different. Keywords: specific growth rate; proximate; bioenergetics; Atlantic cod; energy budget; temperature; season     

Energy allocation in juvenile sablefish: effects of temperature, ration and body size

The lipid deposition of juvenile sablefish Anoplopoma fimbria was examined, in particular, the changes in allocation over time. Growth rates of early juveniles (initial size 36–50 mm total length, L_T) were manipulated using two temperatures (10 and 20° C) and two rations (ad libitum and 3–4% body mass day^?1). Fish L_T, mass and lipid content were measured every 3 weeks for 15 weeks. Irrespective of treatment, the relationship of total lipid content with body size was clearly hyperallometric; small juveniles allocated relatively more energy to growth and less to lipid storage than large juveniles. After adjusting for the influence of body size, temperature and ration significantly influenced body composition but these effects varied over the course of the experiment. In the first 3 week time period, fish on the high ration, high temperature treatment had reduced lipid storage relative to other treatments, but in all subsequent time periods their lipid concentrations were similar to or higher than those of fish on other treatments. In contrast, fish held at low rations and low temperatures initially had average levels of lipid concentration, but after 6 weeks their levels were lower than other treatments. Estimation of allocation to lipid storage over time (proportion of dry mass increase comprised of lipid) suggested that fish in all of the treatments were approaching an asymptotic level of lipid concentration (c. 50–60% of dry mass) but with different rates of lipid increase. Within a treatment, it was predicted that individual differences in allocation would result in trade‐offs between somatic growth and storage. This trade‐off was evident only for fish held on low rations at low temperatures. In contrast, fish held on high rations at high temperatures exhibited the opposite pattern of a positive correlation between somatic growth and storage. These results suggest that lipostatic regulation of appetite is unlikely in juvenile sablefish. When resources are unlimited, this species appears to adopt a maximizing strategy for both somatic growth and lipid accumulation.     

Changes in body composition and morphology of young-of-the-year gulf menhaden, Brevoortia patronus Goode, in Fourleague Bay, Louisiana

Body composition (lipid, water, protein, energy content) and metamorphosis of young-of-the-year gulf menhaden, Brevoortia patronus Goode, were examined over a yearly cycle in Fourleague Bay, Louisiana. Metamorphosis from larvae to juveniles, as indicated by the body depth to length ratio, was completed by 30 mm S.L. Analysis of the length/weight relationship indicated separate equations for larvae/postlarvae (< 30 mm S.L.) and juvenile/subadults (30–100 mm S.L.). Major body composition changes from larvae to subadult included an increase in lipid content, a corresponding increase in energy content, and a decrease in nitrogen content. Young-of-the-year gulf menhaden did not exhibit the seasonal pattern of condition factors typical of adult clupeids. There may be a shift in energy allocation away from protein growth to lipid storage associated with attainment of a larger size. Initiation of emigration may be related to lipid storage and temperature change, with higher lipid content fish migrating first. End-of-year variations in fish size and lipid storage may be related to environmental conditions, such as water temperature and salinity.     

Feeding and growth of juvenile sea bass: the effect of ration and temperature on growth rate and efficiency

The effect of ration on the growth of pairs of juvenile sea bass Dicentrarchus labrax fed squid mantle was recorded at four temperatures: 6, 10, 14 and 18) C, covering the range typical of Welsh coastal waters. Initial weight of the fish ranged from 2.8 to 15.9 g. A predictive model for the maximum meal size (M_max) at temperatures between 10 and 18) C, accounted for 95% of the variance in lnM_max. Even when offered excess food, bass at 6) C had a low rate of food consumption [0.19% body weight (BW) day^?1] and lost weight (G=?0.04% day^?1). Predictive regression models for specific growth rate (G) accounted for 86% of the variance at reduced rations and 70% at maximum meals. The relationship between G (calculated for total biomass per tank) and ration was a decelerating curve. G at maximum meals increased with temperature, at lower rations G decreased with temperature. For a pair of bass with a combined weight of 15 g, predicted maintenance ration ranged between 0.7 and 2.3% BW day^?1 and increased with temperature. Maximum meal size was more sensitive to temperature than maintenance ration. At 18) C optimum ration was 7.4% BW day^?1. At lower temperatures, the optimum ration was the maximum meal. The maximum gross growth efficiency was 17.4% at 18) C. Mean absorption efficiency was 94.8%. Ration level had no significant effect on absorption efficiency, which was lowest at 6) C. Condition indices (Fulton condition factor, wet and dry liver—somatic indices and body depth index) increased with meal size at all temperatures except 6) C. An increase in temperature between 10 and 18) C generally resulted in a decrease in condition indices at a given ration. When comparisons were made at a given standard length, gut and carcass weight increased with ration. Visceral fat and gut weight decreased with increased temperature.     

Temporal changes in the relationship between condition indices and proximate composition of juvenile Coregonus artedi

The Fulton‐condition factor ( K ) and per cent whole‐body water content were examined to determine whether these indices can estimate the proximate composition of juvenile lake herring Coregonus artedi exposed to a simulated Lake Superior winter over a 225 day laboratory experiment. The K was positively correlated to whole‐body crude lipid, crude protein, and gross energy content and negatively correlated to whole‐body water content for each sampling period of the experiment (days 75, 150 and 225). In contrast, there was only a weak positive correlation between K and whole‐body ash content. While per cent water content was negatively correlated with crude lipid, crude protein and gross energy content for each of the three sampling periods, the correlation between this predictor and ash content was only weakly negative. The indices can be used to accurately estimate temporal changes in proximate composition of juvenile lake herring during winter periods.     

Relationship between Fulton's condition factor and proximate body composition in three freshwater fish species

Morphometric‐based condition indices are widely used to assess proximate body composition and, collaterally, feeding and living conditions of fish. However, the exact relationship between condition indices and proximate body composition of fish and its relatedness to life history traits and seasonality has yet to be fully explored. Therefore, the aim of this study was to examine how the Fulton's condition factor (K‐factor) is related to the chemical composition (i.e. lipid, protein, water and carbon content, and molar carbon : nitrogen ratio), length and gonadal development of fish, and how these relationships are influenced by gender and seasonality in three freshwater fish species: Amur sleeper (Perccottus glenii), pumpkinseed (Lepomis gibbosus) and rudd (Scardinius erythrophthalmus). It was found that the strength and direction of association between the K‐factor and proximate body composition can vary markedly among fish species. The K‐factor correlated positively with gonadal development in pumpkinseed and Amur sleeper, while no such relationship existed in rudd. Condition factor can be a reliable measure of lipid content; however, the relationship was stronger in species with higher and more variable lipid contents. Moreover, a striking and consistently negative linkage was found between the K‐factor and water content of the fish body, which corresponds to the findings of several other studies. In turn, any relationship between the K‐factor and the protein content of fish was not detected. Gender seemed to exert a negligible effect on the relationship between the K‐factor and proximate body composition, while seasonal variance was obvious in most relationships.     

Estimation in vivo of the body composition of the ewe by measuring the diffusion space of heavy water

Relationships for estimating body lipids, energy and proteins from the deuterium-oxide (D2O) diffusion space have been determined using 38 dry, pregnant or lactating ewes. The animals were fed as usual and had free access to water up to slaughter. They were injected with deuterium-oxide in the morning, half an hour to 3 hours after being fed the whole daily ration or half of it. Blood samples were collected between 1 and 24 h after injection. The deuterium-oxide content of blood water was determined by infra-red spectrophotometry. The ewes were slaughtered 24 h after injection and their body water, protein, ash and energy were chemically determined. Different blood samples were used to measure the deuterium-oxide diffusion space (theoretical initial concentration Co method). The closest relationship between measured body water (ECM) and D2O diffusion space (ECD2O) was found when the blood sample collected 8 h after injection was discarded. Ewe body composition was estimated using a linear relationship in which ewe body weight and D2O diffusion space were two independent variable (relationship number 5). The best ewe body weight to use was the weight measured at injection time. The estimation of body lipid was significantly affected by physiological status when slaughter weight and true body water, measured at slaughter, were used. The effect of this physiological status decreased when the water content of the digestive tract was taken into account and it was no longer significant when the water content of the uterus and conceptus was considered. Body composition could always be predicted by estimating the water content; the standard error was nearly 1.4 kg. However, the method was not accurate enough to obtain a significant effect of physiological status on lipid estimation. Thus, at present we have proposed identical equations for dry, pregnant or lactating ewes.     

Variation in body condition, fat content and growth rate of Icelandic summer-spawning herring Clupea harengus L.

Interannual variation in Fulton's condition factor ( K ), total body-fat content and growth rate were investigated for the Icelandic summer-spawning herring Clupea harengus over the period from 1963 to 1999. The results show a close relationship between K and the body-fat content in the autumn, both of which correlated to the growth rate in the same calendar year. A relationship was also found between K during the prespawning season (February to June) and K in the autumn (September to December) in the same year. The different measures of body condition were not found to be related to sea temperature or zooplankton indices, while the spawning stock biomass had negative effects on the fish condition. The results imply that the body condition in the autumn is representative of the feeding success over the whole foregoing year and is therefore related to the growth rate that year. The growth rate appears to be relatively uniform throughout the year.     

Do random fluctuations in daily ration affect the growth rate of juvenile three-spined sticklebacks?

The effect of random fluctuations in daily ration on the growth performance of individual juvenile three-spined sticklebacks Gasterosteus aculeatus was studied in experiments lasting 21 days at 14°C and a photoperiod of 10L:14D. Two mean ration levels were used: a maintenance ration of 2%, and a high ration of 6% of initial body weight per day. For experimental fish, the daily ration varied randomly about the required mean value with a coefficient of variation of 33%. The controls received a constant daily ration. The experiment was replicated in winter (Dec.-Jan.) and spring (Mar.-Apr.). At a given ration, there was no significant difference between the specific growth rates of fish receiving constant or varying ration. Neither the final dry weight, final body water content nor final lipid content (% dry weight) differed significantly. As expected, ration had a significant effect on growth rate, final dry weight and body lipid and water content. There was a significant difference in mean growth rate between the winter and spring replicates. The growth rates observed in these experiments were compared with those predicted from a previously described empirical growth model developed for sticklebacks fed constant rations. The model underestimated mean growth rates.     

Effects of diet and temperature on condition, proximate composition and three major macro elements, Ca, P and Mg, in barbel Barbus barbus juveniles

Six feeding groups of 60 early juveniles (0.6?g) of a rheophilic cyprinid barbel Barbus barbus were reared in triplicate in 18?glass aquaria. Fish fed a commercial formulated dry diet Aller Futura were compared with those on natural food??commercially available frozen Chironomidae larvae at 17, 21 and 25°C. Daily food rations were adjusted according to fish biomass, differences in hydration between the two diets, and rearing temperature. No mortality occurred during experiment. Temperature, and especially diet, both affected whole body proximate and mineral composition of B. barbus juveniles. Under conditions of this experiment the formulated diet Aller Futura seems to satisfy the demands for calcium and magnesium, while an elevated lipid content in this diet was found and marginally deficient phosphorus content can be concluded. These resulted in a depressed content of phosphorus and total minerals in the body, and in elevated lipid content and condition coefficient in the Futura-fed fish, but no visible body deformities occurred. Advantages and limitations of the condition coefficient K were discussed. This is a non-destructive, express indicator that can be used as a supplementary tool for estimation of changes in body proximate composition in fish juveniles of similar size within a population.     

Influence of dietary protein and lipid levels on growth, body composition and utilization efficiencies of Arctic charr, Salvelinus alpinus L.

Growth, body composition and feed utilization were evaluated for Arctic charr fed a control diet and nine practical diets formulated to contain three protein levels (34, 44 or 54%) at each of three lipid levels (10, 15 or 20%). Fish were raised for 24 weeks at 12° C. Increasing either dietary protein and/or lipid resulted in improved weight gain, feed efficiency and energy retained. Protein efficiency ratio and protein retained were directly related to dietary lipid and inversely related to dietary protein. A sparing effect of lipid on protein was evident where, at each protein level, Arctic charr utilized protein more efficiently with each increase in dietary lipid. Levels of 34% protein and/or 10% lipid were well below the requirements for Arctic charr (initial weight, 4.6 g). Within the range tested, the dietary combination of 54% protein with 20% lipid maximized weight gain and feed efficiency and resulted in fish with low liver and body lipid contents. However, if lowest feed cost per kg gain is an important factor, the 44% protein—20% lipid diet would be less expensive to use, although it results in a minor reduction in growth potential.     

Effects of Ration on Growth and Reproductive Investment from Larva to Sexual Maturity in the Female Cyprinid Minnow,Tanichthys albonubes

Somatic growth and reproductive investment in female Tanichthys albonubes (Cyprinidae) individually cultured at the laboratory from larva to sexual maturity were examined under low, medium, and satiation food rations. All the 72‐day post‐hatch fish reached sexual maturity under all rations. The standard length, wet body mass, dry ovarian mass, dry liver mass, condition factor, energy content, and number of vitellogenic oocytes were all increased with ration levels. However, food conversion efficiency decreased with ration. There were no significant differences in total number of oocytes per female between rations. The species is a continuous batch‐spawner. The size‐frequency distributions of oocyte diameters showed a continuous pattern, ranging from 0.03 to 0.70 mm, at different rations. The proportion of energy intake allocated to growth decreased with ration levels. Only 3.29–4.60% of energy intake was stored in the ovary. These biological and energetic characteristics allow this fish to reach first maturity with low food intake, although it produced fewer vitellogenic oocytes at lower rations. This property enables T. albonubes survive in its native habitats where food is not only scarce, but also variable temporally and spatially. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Effect of food on the growth of carcase, liver and ovary in female Gasterosteus aculeatus L.

The effect of ration size on the growth of the carcase, liver and ovary in female three-spined sticklebacks, Gasterosteus aculeatus L., was studied experimentally at monthly intervals throughout the annual reproductive cycle. Temperature and photoperiod during each experiment corresponded to the average, natural conditions for that month.
Growth of the carcase was sensitive to ration in all months. At a given ration, the growth rate of the carcase was relatively constant. The water content of the carcase was inversely correlated with ration size. Growth of the liver was also sensitive to ration, but the sensitivity varied in different phases of the reproductive cycle. High rates of weight loss occurred in the liver under starvation, but at maximum rations the liver showed high rates of weight gain particularly in the post-spawning period. Growth of the ovary was relatively insensitive to ration but directly reflected the changes occurring naturally in the reproductive cycle.
It is suggested that the liver acts as a buffer between the carcase and the ovary and this helps to insulate the ovary from the immediate effects of fluctuations in food supply.
The growth rates of the carcase and liver may provide useful indices of the rate of food consumption in natural populations. Although, if the liver is used, its growth has to be interpreted in the context of the annual reproductive cycle.     

Protein, lipid and caloric contents of bluntnose minnow, Pimephales notatus Rafinesque, during growth at different temperatures

Changes in the relative proportions of protein, lipid, water and caloric contents of bluntnose minnow growing at various temperatures (15, 25, 30° C) were investigated by application of the allometry equation, y=ax^b . Fish grew significantly faster at 25° C (closest to optimum), more slowly at 30° C and most slowly at 15° C. Protein, as a percentage of body wet weight, tended to increase with fish size at all temperatures ( b > 1.000), whereas in juveniles (<0.7 g) it decreased ( b < 1.000). However, with the exception of the 15° C group, protein as a percentage of body dry weight, decreased in all groups ( b < 1.000). Temperature appeared to modify the body composition of bluntnose minnows, e.g. decreasing temperature led to significantly enhanced protein content during growth. Lipid (%) and caloric content (cal g⁻¹) increased with increasing fish weight ( b > 1). The slower growing fish (15°, 30° C) deposited significantly more lipid (and had higher caloric contents) than those growing most rapidly (at 25° C). Water content (%) decreased with increasing body weight in all groups. Despite intergroup growth rate differences, all groups showed evidence of a tendency to follow similar trends in b values for body constituents and caloric content (except for protein v. body dry weight for the 15° C group). This suggests a general conservativeness of body composition in bluntnose minnow. The correlations between body constituents, caloric content and body weight were high ( r ²>0.9) so that estimates of body composition can be obtained from body weight for all temperature groups.     

The bioenergetics of the southern catfish (Silurus meridionalis Chen): growth rate as a function of ration level, body weight, and temperature

A feeding-growth experiment was conducted in the laboratory on 114 young southern catfish ( Silurus meridionalis Chen) with initial weights of 8.71–127.9g at 15, 20, 25 and 30°C. The experiment consisted of eight weight-temperature groups, with five ration levels ranging from starvation to satiation in each group. A multiple regression equation fitted to the experimental data was developed to describe the relation between specific growth rate (SGR) and the three factors, ration level (RL), body weight ( W ) and temperature ( T ): SGR = 0.471 + 0.172ln W −0.0443 T +0.0682 T ln(RL + l). This predicts that with increasing temperature the specific growth rate decreases at lower ration levels and increases at higher ration levels. The equation, SGR = a + b ln(RL + l), may be considered as the basic growth model where a is the maintenance metabolism exponent and b is the conversion exponent of the net energy; body weight and temperature influence the two parameters. With this relationship the two antagonistic effects of temperature on growth can be understood, increasing temperature imposes a negative effect on growth due to increment in energy cost for maintenance metabolism, and a positive effect due to higher efficiency of transforming food energy into net energy; the positive effect will increase at higher ration levels. This could also explain why at a restricted ration level relationships between growth and temperature are different in different species.     

Influence of body size and food ration on the energy budget of rainbow trout Salmo gairdneri Richardson

Three size classes (3 g, 200 g, 1–3 kg) of rainbow trout, Salmo gairdneri Richardson, were each fed five rations ranging from zero to a 'maximum' ration and the relationships among body size, food ration, metabolism and growth examined. Water temperature was constant at 11–4°C. The gross growth efficiency in wet weight, K , increased asymptotically with increasing ration but decreased with increasing body size. Changes in fat and moisture content of the body, however, were such that Kin terms of energy utilization was independent of body size when the fish were feeding at the same level. This result was not apparent when wet weight values only were considered and use of a constant to convert wet weight offish to calorific values can lead to errors in interpretation of results. Assimilation efficiency averaged 71 -7 % over all feeding groups and was independent of body size and food ration. Total metabolic expenditure, however, increased with increasing body size and feeding level, but at any given feeding level a similar proportion of the energy of the assimilated ration was used for metabolism. At a single feeding level, therefore, the percentage utilization of energy in each category of the energy budget remained similar over all body sizes. Such a result can only be expected under laboratory conditions but it provides a base-line for comparison with the performance of other more complicated systems.     

Bioenergetics of growth of a cyprinid, Phoxinus phoxinus (L.): the effect of ration and temperature on growth rate and efficiency

Growth of minnows, Phoxinus phoxinus , weighing 1-5.5 g was studied experimentally at five ration levels from starvation to ad libitum and four temperatures ranging from 5 to 15°C. The relationship between specific growth rate (SGR) and ration was a decelerating curve. SGR at maximum rations increased with increased temperature, but at restricted rations it decreased with increased temperature. Predictive models for the specific growth rates were developed using multiple regression. Maintenance rations and optimum rations both increased with increased temperature. Maintenance rations were less sensitive to temperature than optimum rations and mostly lay between 1 and 2% of body weight per day. Conversion efficiencies increased with increased ration from zero value at the maintenance ration to a peak at the optimum ration, then decreased with further increases in ration. At a given restricted ration level, conversion efficiencies generally decreased with increased temperature. At maximum rations, conversion efficiencies were relatively insensitive to temperature. Growth in wet weight, dry weight and energy content showed similar responses to ration, temperature and body weight.     

The effect of ration and temperature on the growth of the three-spined stickleback, Gasterosteus aculeatus L.

The effect of ration on the growth of the three-spined stickleback, Gasterosteus aculeatus , was studied over the temperature range 3·19°C, which corresponded to the range in their natural habitat. With an ad lib ration, the specific growth rate increased with temperature. The relationship between specific growth rate and ration at a given temperature was curvilinear. Regression models were used to predict the maximum, optimum and maintenance rations at each temperature. Whereas maximum ration increased rapidly with temperature, the optimum and maintenance rations were relatively insensitive to temperature. A regression model was used to describe the overall relationship between specific growth rate and ration, temperature and body weight. This model predicted that at low rations, growth rate decreased with temperature, whereas at high rations, growth rate increased slightly with temperature. The gross growth efficiency increased rapidly with rations above maintenance ration up to a maximum at the optimum ration and declined at higher rations. The maximum gross growth efficiencies were at high temperatures and rations about half of the maximum ration. The maximum net growth efficiencies occurred at relatively low temperature and at rations just above maintenance.     

Condition factor and hepatosomatic index as estimates of energy status in male three-spined stickleback

Seasonal variation in direct and indirect measures of energy status was examined using estimates of glycogen, lipid and protein levels in a single cohort of male three-spined sticklebacks from an annual population collected each month over one complete year. Condition factor, somatic condition factor and hepatosomatic index (HSI) were calculated as indirect indices of energy status and the accuracy of these indirect measures as predictors of energy status was investigated. Results indicate that both condition factors were significant predictors of energy reserves (lipid, protein, glycogen and total energy), but that the proportion of variance accounted for was small. Both condition factors perform better as predictors of energy content per unit body weight. The HSI was a significant, but a weak predictor of total glycogen levels over the whole year. On a seasonal basis the relationship between HSI and energy reserves was highly variable. These indices are therefore poor predictors of energy reserves in male three-spined sticklebacks.