State-dependent feed acquisition among two strains of hatchery-reared Arctic charr

The effects of genetic and environmental factors on aggression and feeding hierarchies were studied using X-radiography to measure food intake by hatchery-reared individuals of two strains (Hammerfest and Svalbard) of Arctic charr Salvelinus alpinus . A reduction in food rations and/or water current speed increased intraspecific aggression, and both factors led to increased interindividual variability in food intake, increasing the coefficient of variation (CV). Following a return to pre-manipulation conditions, CVs decreased to their original level. In control groups, CVs and share of group meals were stable throughout the experiment. The increase in CVs following manipulation was the result of a small number of dominant individuals obtaining a high share of the meal. Restriction in food ration affected share of meals, specific growth rates and the frequencies of non-feeding fish, while reductions in water current speed affected only share of meals. Feeding hierarchies were size-dependent in the control groups. In contrast, no relationships between body weight and feeding rank were evident in groups in which food ration or water current speed were reduced. A small, but consistent, difference was revealed in feeding hierarchy responses between the two strains.     

Energetic costs at different levels of feeding in juvenile cod, Gadus morhua L.

The energetic costs associated with feeding by juvenile cod were determined by means of an open-circuit respirometer. Fish acclimated to several temperatures (7, 10, 15 and 18°C) were kept at natural lighting levels, and fed inside their individual respirometers. They consumed a diet compounded from natural foods, at five different ration levels, their oxygen consumption being monitored continually over an 11–16 day period.
After each meal the rate of oxygen consumption increased to above the pre-feeding level, reaching a peak 8–10 h later. With each successive meal the oxygen consumption showed a cumulative increase, reaching a maximum usually after the last meal.
The elevation in metabolic rate associated with feeding was dependent upon ration size, increasing linearly as the food intake increased. The effect was also dependent upon temperature; for fish fed to satiation the total energy cost was equivalent to 11.9, 10.9, 16.4 and 17.1% of the ingested energy at 7, 10, 15 and 18°C respectively. For resting satiated fish the rate of oxygen consumption was close to the maximum rate for active fish.     

Interindividual variations in feeding and growth in rainbow trout during restricted feeding and in a subsequent period of compensatory growth

Compensatory growth responses of rainbow trout Oncorhynchus mykiss were studied by examining food intake and growth of individual fish held within groups that were switched between regimes that involved full and restricted feeding. Restricted feeding led to marked interindividual variability in food intake, probably as a result of the establishment of feeding hierarchies. This disparity in food acquisition was reflected in highly heterogeneous growth amongst the fish fed low rations. When fish were transferred from restricted to full rations, they became hyperphagic and displayed high rates of growth. Growth compensation was most marked amongst those fish which had shown the poorest growth during the period of feed restriction. These results suggest that the feeding hierarchies established under feed restriction did not persist, but were rapidly broken down when food became increasingly available, enabling the previously suppressed fish to gain access to food and to display rapid growth.     

Development of a system for individual feeding of baboons maintained in an outdoor group social environment

We developed a system that allows individual feeding of adult baboons, 8-15 years of age, maintained in an outdoor group social environment. The purpose of the system is to allow careful monitoring and control of individual diet. Baboons were housed in two group cages, 16 females and a single male in one and 12 females and a single male in the other. Baboons exited the group cage once daily and passed along a chute and over a scale into individual cages where they received their individual diets. Food intake was monitored during their 2-hour stay in the individual cages. Baboons rapidly learned to use this system. Food intake and weight were stable within 20 days. Food consumed decreased during the period of sexual receptivity. The maintenance of the group social environment allowed observations on the group's dominance structure and the relationship of dominance to food consumption. Speed of food access in the group cage was related to dominance. Dominance was not related to food consumed in individual cages. The system permits study of many variables related to behavior and food intake while still retaining critical social interactions.     

Growth depensation and group behaviour in juvenile hybrid striped bass Morone chrysops×Morone saxatilis: effects of group membership, feeding method, ration size and size disparity

The effects of group membership (three group sizes), feeding method (single site v. scattered), variation in ration size (low v. high) and starting size disparity (range of starting sizes) on growth, growth depensation and group behaviour of juvenile hybrid striped bass (HSB) Morone chrysops female × Morone saxatilis male were tested. Group membership, starting size disparity and ration size affected the growth, growth depensation and behaviour of juvenile HSB. Growth was higher in fish held within large group memberships and high ration treatments. Growth depensation (size variation among individuals of the same age) in group memberships of three and 12 fish per tank was minimal when compared to intermediate memberships (six fish per tank). In addition, small fish held in tanks with fish of intermediate sizes demonstrated higher growth when compared to small fish housed with large individuals. Some behaviours, specifically activity and scramble feeding, were more elevated in large group memberships when compared to groups with fewer members. Activity was also found to increase over time during size disparity testing. Results suggest that HSB are ideally suited for high-density, intensive aquaculture since they appear to thrive in high group memberships of similar starting sizes and when a sufficient amount of feed in administered.     

The effect of ration level and social rank on the development of fin damage in juvenile rainbow trout

The development and severity of fin damage was examined in groups of juvenile rainbow trout Oncorhynchus mykiss of different strength of feeding hierarchies. The development of dorsal and caudal fin damage over time was compared between four groups fed different ration levels (0·25, 0·5, 1·0 and 1·5% body weight day⁻¹) and between individuals of different feeding rank within each group. Dominance hierarchies were assessed from repeated daily measurements of food consumption of individuals using radiography. The feeding and growth data indicated that the strength of the social hierarchy weakened with increasing ration. Caudal fin damage developed with time in all groups whereas dorsal fin damage developed only under limited rations. The severity of both dorsal and caudal fin damage was significantly dependent on the ration size fed to the group, with lower ration groups sustaining more fin damage. The severity of dorsal fin erosion was greater than for the caudal fin. Within the two lower ration groups, subordinate fish suffered the most dorsal fin damage. The results suggested that the severity of dorsal fin damage within groups of juvenile rainbow trout can be used as an indicator of hierarchy Strength.     

The effects of chronic plasma cortisol elevation on the feeding behaviour, growth, competitive ability, and swimming performance of juvenile rainbow trout

Plasma cortisol elevation, a common consequence of stress, occurs in salmonids of subordinate rank; these fish acquire a smaller share of available food and grow more slowly. This study examined the role of cortisol itself in these phenomena. Cortisol implants, with parallel sham and control treatments, were used to create a chronic threefold elevation in plasma cortisol levels in juvenile rainbow trout, and the individual feeding patterns of the fish were evaluated using X-ray radiography. The three treatment groups were (1) held alone and fed to satiation, thereby providing a measure of voluntary appetite, or mixed together in equal proportions and fed to either (2) satiation or (3) half-satiation, thereby allowing assessment of the additional effects of competitive interaction and food limitation. Chronic plasma cortisol elevation had significant negative effects on individual appetite, growth rate, condition factor, and food conversion efficiency, independent of whether the fish were held under unmixed or mixed conditions. Under the latter, mean share of meal was reduced and fin damage increased in cortisol-treated fish; negative growth effects were more severe with food limitation, but the response patterns were otherwise unchanged. Even in the absence of other groups, cortisol-treated fish showed more variable feeding patterns. When compared at the same individual ration levels, cortisol-treated fish had lower growth rates, reflecting a higher "cost of living." Cortisol treatment had no effect on aerobic swimming performance. These results suggest that the structure of the feeding hierarchy may not be determined solely by competitive ability but may also be greatly influenced by differences in the feeding behaviour of unstressed fish versus stressed fish caused by cortisol elevation in the latter.     

Evidence of differences in growth and food intake regulation in different genetic strains of channel catfish

Norris and USDA-103 strains of channel catfish Ictalurus punctatus were compared for growth rate and food conversion ratio under satiation feeding and restricted feeding (1% body weight day⁻¹) regimes. At the start of the experiment Norris fish weighed 2·8 g, USDA-103 fish weighed 14·0 g. Therefore, a regression of the loge of specific growth rate against the log_e of mean body size with an empirically derived fixed slope of -0·37 was used to compare growth rates. Under both feeding regimes the USDA-103 strain had faster specific growth rates and more efficient food conversion. In subsequent studies, voluntary food intake of size matched fish (60 g average) from these two strains was compared using a radiographic method. Fish were acclimatized to tank conditions for 3 weeks prior to voluntary food intake measurement. Half of the groups were deprived of food for 2 days prior to food intake measurement, while the remaining groups were fed 1% body weight day⁻¹. The USDA-103 strain fish ate significantly more food and grew faster than the Norris strain fish. Previously fasted Norris fish subsequently ate more than their fed counterparts, whereas the fed USDA-103 fish consumed more food than the fasted USDA-103 group. When the USDA-103 strain fish were deprived of food for 4 , 2 or 0 days, all groups subsequently consumed between 4·5 and 5·0% of body weight in one meal. The USDA-103 fish, unlike the Norris fish were not stimulated to consume more after short-duration fasting. Taken together, these results suggest that there are genetic differences in growth, food conversion ratio and regulation of food intake between Norris and USDA-103 strains.     

Maximum daily ration and the pattern of food consumption in haddock, Melanogrammus aeglefinus (L.), and dab, Limanda limanda (L.)

Food intake was measured under conditions of continuous food availability for haddock, Melanogrammus aeglefinus (L.), and dab, Limanda limanda (L.). Haddock consumed approximately 75% of all food during the day, with a peak of feeding at dusk and an inferred peak at dawn; peaks of feeding were associated with larger meals. By contrast, dabs were mainly night-time feeders with daily consumption rates that were approximately 40% lower than those shown by haddock. Comparison of haddock maximum daily ration with published estimates of consumption in the wild indicate that wild haddock feed at between 15 and 22% of their maximum capacity. Using published data on gastric evacuation, the relationship between meal size and stomach residuum was investigated. There was no detectable relationship between these two variables.     

Food consumption and growth of larvae and juveniles of three cyprinid species at different food levels

Synopsis The relationships between food availability, consumption and growth were analyzed from the onset of feeding to an age of 90 days in three cyprinid species. Fish were held at 20 ± 0.5° C and given two (three) constant rations of approximately 30, (40) or 100% dry body weight (dbw) ind^-1 day^-1. Food consisted of living zooplankton, the size of which correlated with fish size. At high food densities consumption rates decreased rapidly with fish size in all three species. At reduced rations, fish consumed most of the food offered until they were larger than 10 mg dbw. In all species and at each feeding level daily rations consumed increased allometrically with body size. Respiration rate, expressed as routine metabolic rate differed insignificantly between the three species. At high ration levels, growth rates of small bleak, Alburnus alburnus, were distinctly lower than those of roach, Rutilus rutilus, and blue bream, Abramis ballerus. At low food supply all three species grew at similar rates. Assimilation efficiency at low food conditions was approximately twice that of the well-fed groups. If the caloric equivalents of oxygen consumption as measured in well-fed fish are applied to fish fed at low rations their energy budgets do not balance. This indicates the limitations of fish larvae in the partitioning of energy for growth or activity at such conditions.     

Temperature and state‐dependence of feeding and gastric evacuation in juvenile Pacific halibut

Relationships between nutritional state, behavioural response to prey and gastric evacuation rates were examined in juvenile Pacific halibut Hippoglossus stenolepis feeding on squid. Pacific halibut reared at 2, 6 and 10° C were fasted for 1 or 7 days to generate variation in energetic state. The 7 day fast resulted in measurable declines in condition indices at 10 and 6° C but not at 2° C. At 10° C, all Pacific halibut consumed the first meal offered, but fish previously fasted for 7 days took significantly longer to locate and consume the meal than fish fasted for only 1 day. At 2° C, Pacific halibut fasted for 7 days did not generally consume the first meal offered, but resumed feeding 2·1 days sooner, on average, than fish fasted for only 1 day. The gastric evacuation rate of the squid meal was best described by a power model with near‐exponential curvature ( a  = 1·011). The evacuation rate was strongly temperature‐dependent ( Q ₁₀ = 3·65) but displayed the same degree of variability at each temperature. The evacuation rate in Pacific halibut was not affected by feeding history, body size or energetic state. Furthermore, individual variation in gastric evacuation rate was not correlated with feeding responsiveness at any temperature. These results indicate a general plasticity in the behavioural but not physiological aspects of energy acquisition.     

The effect of feeding regime on growth, locomotor activity pattern and the development of food anticipatory activity in greenback flounder

The mean final weight of juvenile greenback flounder Rhombosolea tapirina fed a single daily meal during the scotophase was lower than that of groups fed continuously or fed twelve, six, three or one meal during the photophase. Final weight did not differ among the latter five groups. Size variation in all groups of fish decreased during the course of the trial over 120 days. Groups assigned to a higher frequency (and hence a longer total daily meal duration) of daily meals were less active during mealtime. A distinct food anticipatory effect was observed in the groups of fish receiving a single daily meal during either photophase or scotophase, and also developed before at least one meal in the group of fish fed three meals per day.     

The influence of ration size and feeding frequency on ammonia excretion by juvenile Atlantic cod, Gadus morhua L.

Small groups of juvenile Atlantic cod, Gadus morhua L., were kept at 14°C in through-flow tanks and were fed known quantities of a compounded diet of natural food. The cod were fed single and multiple meals with ration size in the range 0.5 to 4.1% of total wet fish body weight. Ammonia production in each feeding experiment was monitored continuously.
For single-meal experiments, significant relationships were derived between ration size and (a) total ammonia excreted, (b) total exogenous ammonia excreted above endogenous excretion levels, (c) duration of the elevated phase of ammonia excretion, (d) maximum rate of ammonia excretion, and (e) time delay after feeding to reach maximum rate of ammonia excretion. Relationships between nitrogen loss as ammonia and nitrogen intake were examined and estimates of endogenous excretion rate and maintenance ration made.
Repetitive feeding resulted in cyclical variation in ammonia excretion. At the lowest ration size, ammonia excretion rates had nearly returned to the pre-feeding level within 24 h. At higher feeding levels, the effect of each successive meal tended to be cumulative, resulting in increasingly higher ammonia excretion rates which only stabilized towards the end of the experiments.     

A nutritional analysis of foraging in the Malabar giant squirrel (Ratufa indica)

The effect of nutrients and their availability on the diet of the herbivorous Malabar giant squirrel Ratufa indica (Sciuridae) was investigated at Magod and Bhimashankar in western India. The daily consumption of food items (percent wet diet) and the intake rates of these items and the contained nutrients (wet g s^-1) were determined by continuous observation of focal animals. Water content was a significant positive predictor of relative food item consumption while mineral contents in general had the opposite effect. The intake rates of water and more digestible nutrients such as nonstructural carbohydrates, as well as the intake rate of minerals such as zinc (at Magod), were significant positive predictors of the relative contribution of an item to the daily diet, thus indicating considerations of feeding costs versus benefits. Tannins, some alkaloids, and other secondary metabolites may negatively influence food choice. Protein content, relative to digestibility reducers, influenced food item consumption only at Bhimashankar. There was seasonal variation in daily biomass consumption. At Bhimashankar, daily biomass consumption increased with the proportion of fruit in the diet while this did not occur at Magod. This is perhaps a result of the higher water content and the lower content of some soluble nutrients within fruit at Bhimashankar relative to Magod. There was also intra-month variation between individual squirrels in daily biomass of food consumed. Squirrels consumed ephemeral food items opportunistically and non-ephemeral items such as mature leaves and bark on a regular daily basis. Squirrels probably obtained minerals and nitrogen from both fruit (especially seeds) and non-fruit sources (mature leaves and bark in the case of minerals, young and mature leaves in the case of nitrogen). However, they obtained lipid and non-structural carbohydrates mostly from fruit. Squirrels were selective in their utilization of tree species on a monthly basis, but this selectivity was not evident over a longer time period because individuals were constrained to use different phenological stages of tree species present within their territories. Large body size in this squirrel permits dietary flexibility and enables an overall generalist feeding strategy.     

The influence of environmental manipulations on inter– and intra–individual variation in food acquisition and growth performance of Arctic charr, Salvelinus alpinus

Repeated measurements of food intake made on juvenile Arctic charr, Salvelinus alpinus , held under different rearing conditions enabled examination of the effects of environmental manipulations on both intra– and inter–individual variations in food intake to be made. This permitted the assessment of the influences of differential food acquisition on individual growth rates and biomass gain. When charr were held in isolation individual fish showed relatively little day–to–day variability in food intake and inter–individual differences in intake were small ('base–fine' values). All fish exhibited positive rates of growth and the overall range was narrow. Nevertheless, there was a highly significant positive correlation between food intake and growth, indicating that those individuals that consumed the greatest quantities of food were also those that had the highest rates of weight gain. The rearing of charr in groups led to increases in both intra– and inter–individual variations in food intake to levels considerably above 'base–line'. This increased variability in food intake was reflected in rates of weight gain being more variable amongst the charr reared in groups, with fish that lost weight often being recorded. Manipulation of the rearing environment had marked influences upon intra–individual variability in food intake, inter–individual differences in food acquisition and rates of weight gain. High stocking densities and exposure of the fish to moderate water currents were most effective in reducing levels of variability to approach those observed under 'base–line' conditions.     

Capacity for growth compensation in juvenile three-spined sticklebacks experiencing cycles of food deprivation

The capacity of three‐spined sticklebacks Gasterosteus aculeatus (initial mean mass 0.280 g) to compensate for recurrent periods of food deprivation of 2, 4 or 6 days followed by 2 days of ad libitum feeding on enchytraeid worms over 56 days was assessed by measuring appetite and growth. Control fish were fed daily. The total number of days on which fish were fed ranged from 14 (6‐day cycle) to 56 (controls). Deprived sticklebacks were hyperphagic on the first day of re‐feeding in a cycle and this hyperphagia increased with successive cycles. Mean daily consumption on first day of refeeding was: controls, 62.9 mg; 2 days, 108 mg; 4 days, 98.8 mg; 6 days, 101 mg. The hyperphagia did not increase as the preceding period of deprivation within a cycle increased. Hyperphagia was not maintained on the second day of re‐feeding. The 4 day and 6 day groups initially showed hypophagia on the second day of re‐feeding. Mean daily consumption on second day of re‐feeding was: controls, 62.6 mg; 2 days, 62.2 mg; 4 days, 54.2 mg; 6 days, 50.0 mg. Over the experiment, consumption on the second day of the 4 day and 6‐day groups increased towards the control level, suggesting a developing compensatory response. The highest mean daily consumption per days fed was shown by the 2 day group. The relationship between number of days fed and total food consumption and specific growth rate suggested that the 2 day group almost compensated for the periods of deprivation. Performance declined for the 4 day and 6 day groups, although even at the highest level of deprivation, a positive growth was achieved. Mean specific growth rate in mass (% per day) was: controls, 2.33; 2 days, 1.89; 4 days, 1.21; 6 days, 0.86. Initial mass and total food consumption accounted for most of the variance in specific growth rate. Other indices of performance including lipid concentration, dry matter concentration and the RNA:DNA ratio in white muscle were positively related to quantity of food consumed by each group. Growth efficiency of sticklebacks in terms of wet mass gained and wet mass consumed over the experimental period was 19.4% and did not differ among control and treatment groups. The growth rates of the sticklebacks experiencing cyclical deprivation were comparable to growth rates previously recorded for sticklebacks fed daily, but consuming similar mean daily rations.     

Do random fluctuations in the intervals between feeding affect growth rate in juvenile three-spined sticklebacks?

This study measured the effects of regular and irregular intervals between feeds on the growth performance of juvenile Gasterosteus aculeatus . The experimental period was 21 days at 14) C and photoperiod of 10L: 14D. The fish were housed individually. The control fish received a constant ration every day. Fish on a constant interval were fed on days 1, 5, 9, 13, 17 and 21. Fish on a random regime were fed with the same average interval between feedings, but the interval varied randomly between 1 and 5 days. The rations were calculated so that over the 21-day period, all fish were supplied with the same total quantity of food. The two ration levels were: 2% (maintenance ration) and 6% of the initial body weight per day. At a given ration, the feeding interval had no significant effect on specific growth rate, RNA/DNA ratio and lipid contents. The percentage dry matter was slightly, but significantly lower in treatment groups than in the control group. Groups receiving a mean 2% ratio consumed all the food supplied. At a mean ration of 6%, the control group ate 100%, the regular interval group 95·4% and the irregular interval 98·3% of the total food supplied. For the temporal patterns of feeding used, the fish were able to adjust their food intake, when food become available, to compensate for short periods of food deprivation and maintain their growth performance except for a decrease in dry matter content.     

The effect of feeding frequency on growth performance of juvenile cobia,Rachycentron canadum (Linnaeus, 1766)

The present study was carried out to investigate the number of daily feeding sessions that results in maximum growth of juvenile cobia under laboratory conditions. Groups of eight fish (110 g) were randomly distributed in twenty 500‐L tanks and hand‐fed a commercial diet for 60 days. The same amount of feed was offered daily, divided in 1, 2, 3, four or six meals. None of the parameters associated with growth performance (survival, weight gain, specific growth rate, feed intake, condition factor or size variation) showed any significant differences among treatments. Although under the present conditions feeding frequency had no effect on the growth performance of cobia larger than 110 g, in commercial farming operations where large numbers of fish are kept within a single rearing structure, fish may have aggressive interactions during feeding. Under these conditions, it is difficult to ensure that all cobia are fed to satiation and thus it is usual to provide two or more meals per day. The present results indicate that for an individual cobia the provision of more than one daily meal has no significant effect on growth performance.     

Dairy goats adjust their meal patterns to the fibre content of the diet

Few studies have investigated how meal patterns of ruminants are affected by diet fibre content. Dairy goats (N = 32) in late lactation and early gestation were housed in eight groups of four goats, with all combinations of breed (Alpine and Saanen) and lactation number (1 and 2) represented in each group. Each goat had access to its own individual feed trough placed on a weigh scale with data logged automatically. All goats were fed the same total mixed ration (TMR; 30% concentrate and 44.6% NDF in DM) ad libitum for a control period of 22 days. Using the same feed ingredients, half of the groups were then offered a High fibre diet (20% concentrate; 47.3% NDF), and the other half a Low fibre diet (40% concentrate; 41.5% NDF) for a treatment period of 16 days. Daily meal patterns (meal frequency, duration and size, feeding rate, daily feed intake and daily feeding time) were computed for each animal using a meal criterion of 8 min. The last 10 days for each period (control and treatment) were used to calculate individual period means and individual differences between the two periods. During the control period, the goats ate on average 12.1 ± 0.49 meals/day, consuming 4.2 ± 0.10 kg fresh TMR daily. When the ration changed, all measures of feeding behaviour except meal size changed asymmetrically for the goats on the two diets. Goats fed the High fibre diet reduced their meal frequency by 10%, and the first meal after feed distribution lasted 11% longer, leading to a 9% reduction in feeding rate and no significant changes in daily feed intake and daily feeding time. Goats on the Low fibre diet did not significantly change their meal frequency or meal size, but the combined changes nevertheless led to a 9% increase in daily feed intake. On the Low fibre diet, goats were able to increase their feeding rate by a third, leading to a reduction in meal durations, thus reducing daily feeding time by 13%. Goats adapt their feeding behaviour to the fibre proportion of the offered diet, with more changes when fibre content is lowered, which needs to be taken into account when comparing phenotypes and adaptability of small ruminants to different diets.     

Differences in feeding between 1+ and 2+ hatchery brown trout exposed to low water temperature

Both 1 + and 2+ brown trout fed during the day and at night when held in hatchery tanks at low temperatures (2·7–3° C). Over 60% of the daily ration was consumed during daylight hours, but there were differences in feeding behaviour between fish of the two age groups: the 1 + trout had greater feeding activity at night than the 2+ fish.