Fuel, fasting, fear: routine metabolic rate and food deprivation exert synergistic effects on risk-taking in individual juvenile European sea bass

1. Individuals of the same species often exhibit consistent differences in metabolic rate, but the effects of such differences on ecologically important behaviours remain largely unknown. In particular, it is unclear whether there is a cause-and-effect relationship between metabolic rate and the tendency to take risks while foraging. Individuals with higher metabolic rates may need to take greater risks while foraging to obtain the additional food required to satisfy their energy requirements. Such a relationship could be exacerbated by food deprivation if a higher metabolic demand also causes greater mass loss and hunger. 2. We investigated relationships among metabolic rate, risk-taking and tolerance of food deprivation in juvenile European sea bass. Individual fish were tested for risk-taking behaviours following a simulated predator attack, both before and after a 7-day period of food deprivation. The results were then related to their routine metabolic rate (RMR), which was measured throughout the period of food deprivation. 3. The amount of risk displayed by individual fish before food deprivation showed no relationship with RMR. After food deprivation, however, the amount of risk among individuals was positively correlated with RMR. In general, most fish showed an increase in risk-taking after food deprivation, and the magnitude of the increase in risk-taking was correlated with the rate of individual mass loss during food deprivation, which was itself strongly correlated with RMR. 4. The observation that RMR was related to risk-taking behaviour after food deprivation, but not before, suggests that although RMR can influence risk-taking, the strength of the relationship is flexible and context dependent. The effects of RMR on risk-taking may be subtle or non-existent in regularly feeding animals, but may lead to variability in risk-taking among individuals when food is scarce or supply is unpredictable. This synergistic relationship between RMR and food deprivation could lead to an increased likelihood of being predated for individuals with a relatively high intrinsic energy demand during times when food is scarce.     

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.     

Refuge use by fish as a function of body weight changes

Refuge use provides a good model for the study of trade-offs between the benefits of predator avoidance and the costs of lost feeding opportunities. We manipulated the latter costs by subjecting similar-sized three-spine sticklebacks to 2 days of food deprivation followed by a 2-day re-feeding period and recorded associated changes in body weight and refuge use. Food deprivation resulted in a decrease and re-feeding in an increase in the duration of refuge use by fish. Emergence times of fish from the refuge were extremely variable (with a ratio of 1:127 between the shortest and the longest ones) but individual ranks were highly consistent between different days of testing, suggesting that emergence times were individually characteristic. Percentage weight change of fish in response to the experimental treatments also showed a high level of inter-individual variation ranging from 0–17%. A significant positive correlation was found between the percentage weight lost and the percentage decrease in emergence time from a refuge after food deprivation and similarly between the percentage weight gained and the percentage increase in refuge use after re-feeding. The relationship between energy turnover and behavioural strategies is discussed. Received: 13 December 1998 / Received in revised form: 2 May 1999 / Accepted: 7 June 1999     

Reduction of predation risk under the cover of darkness: Avoidance responses of mayfly larvae to a benthic fish

Summary Mayfly larvae of Paraleptophlebia heteronea (McDunnough) had two antipredator responses to a nocturnal fish predator (Rhinichthys cataractae (Valenciennes)): flight into the drift and retreat into interstitial crevices. Drift rates of Paraleptophlebia abruptly increased by 30 fold when fish were actively foraging in the laboratory streams but, even before fish were removed, drift began returning to control levels because larvae settled to the substrate and moved to areas of low risk beneath stones. This drifting response was used as an immediate escape behavior which likely decreases risk of capture from predators which forage actively at night. Surprisingly, drift most often occurred before contact between predator and prey, and we suggest that in darkness this mayfly may use hydrodynamic pressure waves for predator detection, rather than chemical cues, since fish forage in an upstream direction. Although drifting may represent a cost to mayfly larvae in terms of relocation to a new foraging area with unknown food resources, the immediate mortality risk probably out-weighs the importance of staying within a profitable food patch because larvae can survive starvation for at least 2 d. In addition to drifting, mayflies retreated from upper, exposed substrate surfaces to concealed interstitial crevices immediately after a predator encounter, or subsequent to resettlement on the substrate after predator-induced drift. A latency period was associated with this response and mayflies remained in these concealed locations for at least 3 h after dace foraging ceased. Because this mayfly feeds at night and food levels are significantly lower in field refugia under stones, relative to exposed stone surfaces, predator avoidance activity may limit foraging time and, ultimately, reduce the food intake of this stream mayfly.     

Effect of food deprivation on oxidative stress biomarkers in fish (Sparus aurata)

Oxidative stress in fish (Sparus aurata) as a consequence of food restriction and fasting, has been studied. Four groups of fish were maintained for 46 days under different conditions of food supplementation: a control group with no food restriction (ratio of food/fish of 2% w/w), two groups of animals with restricted food supplement (1 and 0.5%) and a fasting group (no meal addition). Finally, all the fish were provided with food at the same ratio as the control group for the last 7 days. Sampling and weighing of fish were carried out every week and their livers were used for the analysis of known biomarkers of oxidative stress. Malondialdehyde and oxidized glutathione levels increased at the third week in fish with partial or total food deprivation, but these levels returned to normal values when the fish readapted to the control conditions. Antioxidant enzymes were also analyzed and significant increases in superoxide dismutase (SOD), glutathione reductase and glutathione peroxidase activities were found in parallel with food restriction; however catalase activity decreased in fasting fish. New SOD isoforms were detected by isoelectrofocusing in fish under food restriction at the second week, which disappeared when starved fish returned to the control conditions. These new SOD isoforms were detected before the appearance of other usual oxidative stress biomarkers.     

Bombesin stimulates insulin secretion and reduces food intake in the baboon

Baboons received a 5-minute intravenous infusion of either saline or bombesin (BBS; 1-4 micrograms/kg) following 3 1/2 or 16 1/2 hours of food deprivation and were then allowed to eat for 30 minutes. Plasma insulin was significantly elevated following five minutes of BBS infusion, but there was no change of plasma glucose over the same interval. Bombesin infusion resulted in a dose-dependent decrease of food intake that was independent of deprivation time. Plasma insulin levels following the 30-minute meal were significantly depressed after BBS infusions such that there was essentially no change of plasma insulin over the duration of the meal, even though the baboons did not totally suppress their food intake. Following 3 1/2 hours food deprivation, BBS suppressed the post-prandial rise of plasma glucose in a dose-dependent manner. The results provide further evidence that BBS and/or structurally-related peptides are involved in the regulation of feeding and metabolism.     

Effects of cholecystokinin octapeptide (CCK-8) on food intake in adult and aged rats under different feeding conditions

The effects of CCK on food intake were investigated under fixed feeding conditions in comparison to a test meal taken after 16 h of food deprivation. The experiments were performed on young adult rats (8 weeks old) as well on aged rats (23 months old). Intraperitoneal CCK-8 (8 and 40 μg/kg) significantly reduced the size of a test meal following 16-h food deprivation. This effect was independent of the age of the rats. However, under fixed feeding conditions neither of the doses used in this study reduced food intake in the young adult rats, whereas the highest dose of 40 μg/kg did so in the aged rats. These results suggest that the hypophagic effect of exogenous CCK-8 depends on experimental conditions, food intake being reduced after a period of food deprivation but not under a fixed feeding regimen in adult animals. Furthermore, the data suggest that age is a factor contributing to the complex behavioral actions of CCK, because only old animals were more susceptible to an anorectic action of CCK under the fixed feeding schedule. An explanation may lie in an interaction of other known behavioral effects of CCK (e.g., anxiogenic, mnemonic action) with its effects under the different feeding schedules.     

捕食风险对高原鼠兔食物大小选择的影响

文章报道了捕食风险条件下高原鼠兔对食物大小选择的格局。在实验箱中放置艾虎以改变捕食风险水平, 食物按体积大小分为4种食物项目, 并测定各项目摄入率和取食单个食物项目的进食时间, 结果表明, 摄入率与进食时间依食物项目体积的增大而增加。将大食物与小食物项目配对并供高原鼠兔选择时, 食物项目的利用率视环境状况而不同。捕食风险处理中, 小食物利用率依其进食时间的减少而增加, 其程度与所配对的食物项目的摄入率和进食时间有关。在捕食风险的作用下, 高原鼠兔的食物选择格局反映了能量摄取与风险避免间的权衡。     

Refuge use by fish as a function of body length-related metabolic expenditure and predation risks

Theory predicts that staying in a refuge has benefits in terms of predator avoidance and costs in terms of lost feeding opportunities. In this study, we investigated how the relative importance of these costs and benefits changes with increasing body length. This is of particular interest in animals such as fish, which show continuous growth throughout their lives. Our results suggest that larger fish are subject to lower predation risks and are less affected by food deprivation than small fish, with fish decreasing their responses to food-deprivation treatments more strongly with increasing body length than to predation treatments. This may explain our observation that large fish emerged later from a refuge than small ones and spent shorter times outside the refuge. The key role of differential responses to food deprivation was further illustrated by the finding that the relative weight loss of individual fish was strongly correlated with a reduction in hiding time even in the absence of body length differences. The importance of inter-individual differences in metabolic rates for the decision-making behaviour of animals is discussed.     

Feeding while evading predators by a lotic mayfly: linking short-term foraging behaviours to long-term fitness consequences

Laboratory experiments were performed to quantify the combined effects of food abundance (low, high) and predation threat, imposed using a model fish (safe, risky) on fitness correlates (i.e. growth, time to emergence, adult body mass, fecundity, egg size) of the mayfly Baetis tricaudatus. These effects were determined by rearing larvae under different combinations of food abundance and predator threat. Fitness correlates were significantly affected by food abundance, predation threat or the interaction of these factors. High food abundance and the absence of predation threat significantly increased larval growth rates, adult body mass, fecundity, egg size, and decreased time to emergence. Long-term effects of predator threat and food abundance on fitness correlates of B. tricaudatus were compared to previously conducted short-term patch choice trials to test for concordance between short-term measures of patch choice and their potential longer-term fitness consequences. This comparison indicated that patches that were utilized the most were those that yielded the highest fitness benefits. These results suggest that behavioural costs of balancing mortality risk due to predation against food acquisition by B. tricaudatus can have strong fitness consequences.     

The effect of feeding hierarchy on individual variability in daily feeding of rainbow trout, Oncorhynchus mykiss (Walbaum)

Three groups of rainbow trout, Oncorhynchus mykiss , (initial weight 40 g) were fed one of three rations (low, medium or high) for 73 days. Consumption by individual fish within the three ration groups was measured on four occasions (days 27, 55, 64 and 72) using radiography. Food intake by individual fish varied between days and this variability was expressed using the coefficient of variation (CV). Dominant fish within each ration group were defined as individuals with the greater share of the group meal and these fish had low CVs for food intake indicating relatively little variation in daily consumption. By contrast, the fish which had consumed a low mean proportion of the group meal displayed high CVs for food intake, indicating considerable variability in the sizes of individual meals consumed by these fish. As group ration increased, the range of meal sizes and individual CVs in daily feeding decreased, suggesting that the strength of the feeding hierarchy and the variability in individual consumption decreased as food availability increased. It is suggested that radiography can be used to assess social relationships within groups of fish allowing the assessment of feeding hierarchies in larger groups of fish than would be possible by observational techniques.     

Compensatory Growth in Juveniles of Freshwater Redclaw Crayfish Cherax quadricarinatus Reared at Three Different Temperatures: Hyperphagia and Food Efficiency as Primary Mechanisms

Feeding restriction, as a trigger for compensatory growth, might be considered an alternative viable strategy for minimizing waste as well as production costs. The study assessed whether juvenile redclaw crayfish Cherax quadricarinatus (initial weight 0.99 ±0.03 g) was able to compensate for feeding restriction at different temperatures (23±1, 27±1 and 31±1°C). Hyperphagia, food utilization efficiency, energetic reserves, and hepatopancreas structure were analyzed. Three temperatures and two feeding regimes (DF-daily fed throughout the experiment and CF- 4 days food deprivation followed by 4 days of feeding, intermittently) were tested. The restriction period was from day 1 to 45, and the recovery period was from day 45 to 90. The previously restricted crayfish held at 23, 27, and 31 ± 1°C displayed complete body weight catch-up through compensatory growth following the restriction period with depressed growth. The mechanisms that might explain this response were higher feed intake (hyperphagia), and increased food utilization efficiency. Hepatopancreatic lipids were used as a metabolic fuel and hepatosomatic index was reduced in the previously restricted crayfish, but recovery at the same level of unrestricted crayfish occurred after the shift to daily feeding. The highest temperature affected adversely growth, feed intake, food efficiency, and metabolism of crayfish, whereas the lowest temperature and feeding restriction induced a more efficient growth of the crayfish.     

The effects of vitamin B6 deprivation with 4-deoxypyridoxine in meal-fed rats

Weanling male rats were trained to consume a control diet in one 4-hr meal a day. Rats were then fed one of three experimental diets for 7 days: (a) control diet minus B6 (-B6); (b) control diet (B6); or (c) control diet restricted to food intake of -B6 (B6R). Xanthurenic acid excretion was greater before and after a tryptophan load in -B6 than in B6R. Body weight gain, food intake, and food efficiency were not different between -B6 and B6. However, in B6R body weight gain, food intake and food efficiency were lower than that in both -B6 and B6. Serum glucose (12-16 hr after meal) and percentage body fat were similar in all three groups. No differences were found in organ size between the -B6 and both control groups indicating that short-term deprivation and deoxypyridoxine did not affect organ size. No significant differences were observed for serum growth hormone (GH), pituitary GH, serum prolactin (PRL), or pituitary PRL among the three groups. When meal fed, differences were minimized between B6 deprived and unrestricted control (B6) rats in body weight gain, food intake, food efficiency, and body composition.     

Mercury Concentration in Breast Milk and Infant Exposure Assessment During the First 90 Days of Lactation in a Midwestern Region of Brazil

Breast milk samples collected from 18 nursing mothers between the 15th and 90th day of lactation were digested in nitric acid in a microwave, and total mercury (THg) levels were quantified by atomic fluorescence spectrometry. Participants responded to a 24-h dietary recall questionnaire on the 74th and 76th day of lactation and to a Food Frequency Questionnaire querying the frequency of fish intake over the last 90 days. Usual intake was estimated using the PC-SIDE software package. A meal of fish was offered on the 75th day of lactation. Mothers’ individual mean THg levels ranged from <0.76 to 22.7 ng/mL during the period, and the mean level for all samples (n?=?142) was 6.47?±6.04 ng/mL. The multilevel mixed linear model used showed high heterogeneity of the mercury levels among the mothers, and THg levels did not change significantly over the period under study. However, a significant increase in THg levels was observed after the intervention with the fish meal. Exposure increased for most infants on the 90th day of lactation, with intakes exceeding the THg provisional tolerable weekly intake (PTWI) at least once during the period for 77.8 % of samples. Mothers consumed mostly food from the fat and grain groups, and a significant correlation was detected between consumption of food of these groups and breast milk THg levels (p?=?0.006 and 0.007). A significant correlation was also found between vegetable consumption and carbohydrate intake and THg levels in the samples (p?=?0.015 and 0.045, respectively). No correlation was found between mothers’ daily fish consumption frequency and THg levels. Although this study showed that mercury intake by infants during lactation may exceed the toxicologically safe exposure level (PTWI), we nevertheless believe that the benefits of lactation for both the mother and the infant outweigh the eventual risks that this exposure may represent.     

Reduced feeding during water deprivation depends on hydration of the gut

Removal of drinking water at the start of the dark period reduced food intake in freely feeding rats within 45 min. Both first and later meals were smaller during 7.5 h of water deprivation, but meal frequency did not change. Ingestion of a normal-sized meal (3 g) rapidly increased plasma tonicity when drinking water was withheld, but intravenous infusions of hypertonic NaCl causing similar increases in plasma tonicity did not reduce feeding. Feeding during 6 h of water deprivation was restored by slowly infusing the volume of water normally drunk into the stomach, jejunum, or cecum, but not in the vena cava or hepatic portal vein. The infusions did not alter water or electrolyte excretion or affect food intake in rats allowed to drink. We conclude that the inhibition of feeding seen during water deprivation is mediated by a sensor that is located in the gastrointestinal tract or perhaps in the mesenteric veins draining the gut, but not the hepatic portal vein or the liver. In the absence of drinking water, signals from this sensor provoke the early termination of a meal.     

An analysis of feeding patterns in the adult pig

Feeding behavior has been analyzed in Large-White and Chinese pig breeds. Maximally automatized apparatuses, especially adapted to animal size, were used. Besides the usual parameters measured such as food intake, meal size, meal duration, meal frequency, intermeal intervals and postprandial correlation, these apparatuses also permitted us to measure the rate of intake during the meal. This rate was constant throughout the meal in ad libitum conditions and after 24-hour deprivation. A positive correlation between meal size and length of the postmeal interval was found in only half of the animals. During this work, we began to study the mechanisms initiating food intake.     

Development and validation of methods for measuring multiple biochemical indices of condition in juvenile fishes

The investigation established and validated methods to measure multiple biochemical indices of condition (whole body total lipids, whole body triglycerides, muscle RNA : DNA ratio and muscle protein) simultaneously in the same individual juvenile fish. It also provided examples of their application using a species comparison (salmonids and cyprinids) of the degree of change in these indices after food deprivation. The results showed that juvenile rainbow trout Oncorhynchus mykiss were much more susceptible to a decline in all biochemical indices of condition than juvenile fathead minnows Pimephales promelas upon food deprivation. The combination of these techniques can be used to accurately assess condition and growth potential of individuals from wild fish populations.     

豆粕和土豆蛋白替代饲料中鱼粉对异育银鲫生长及能量收支的影响

利用全能量收支同步测定呼吸仪系统测定了三种不同蛋白源(鱼粉、豆粕、土豆蛋白)饲料对银鲫生长及能量收支各组分的影响。对照饲料全部以鱼粉为蛋白源,另两种饲料中30％的蛋白质分别来自豆粕和土豆蛋白。试验结果表明,银鲫对三种蛋白源的饲料的摄食率,食物转化效率均有显差异,以鱼粉组最高,豆粕其次,土豆蛋白最低。能量收支各组分也均有显差异,能量收支式如下：鱼粉88．1C=13．77F 2．95U 24．1G 41．8R 豆粕 102．3C=17．1F 2．8U 16．4G 55．4R土豆蛋白105．5C=14．9F 1．3U 14．4G 67．4R式中C、F、U、G、R分别代表摄食能、排粪能、排泄能、生长能、代谢能。     

Influence of the subfornical organ on meal-associated drinking in rats

A lesion of the subfornical organ (SFO) may disrupt drinking after a meal of dry chow as it does drinking after intragastric administration of hypertonic saline. Food and water intakes of SFO-lesioned (SFOX) and sham-lesioned rats were measured during 90-min tests following various lengths of food deprivation. During the tests, all rats began eating before they began drinking. After 20-24 h of food deprivation, latency to begin drinking after eating had started was longer for SFOX than for sham-lesioned rats. Plasma osmolality was elevated by 2-3% in both lesion groups at 12 min, the latency for sham-lesioned rats to drink, but SFOX rats nevertheless continued eating and delayed drinking. Eating after shorter 4-h food deprivations and ad libitum feeding produced more variable drinking latencies and less consistent effects of SFO lesion. During 24 h of water deprivation, SFO lesion had no effect on the suppression of food intake and did not affect food or water intakes during the first 2 h of subsequent rehydration. These findings indicate that the SFO is involved in initiating water intake during eating and in determining drinking patterns and the amount of water ingested during a meal.     

Patch use behaviour in benthic fish depends on their long-term growth prospects

Animals foraging in a heterogeneous environment may combine prior information on patch qualities and patch sample information to maximize intake rate. Prior information dictates the long-term expectations, whereas prior information in combination with patch sample information determines when to leave an individual food patch. We examined patch use behaviour of benthic feeding fish in their natural environment at different spatial scales to test if they could determine patch quality and if patch use behaviour was correlated with environmental quality. In seven lakes along a gradient of environmental quality (measured as maximum benthivore size), we made repeated measurements of giving-up density (GUD) in artificial food patches of different qualities. At the largest spatial scale, between lakes, we tested if giving-up densities revealed the long-term growth expectation of benthic fish. At the local scale of patches and micro patches we tested for the ability of benthic fish to assess patch quality, and how this ability depended on the patch exploitation levels between the different lakes. We found that GUD was positively related to maximum size of bream, suggesting that short-term behavioural decisions reflected long-term growth expectations. Benthic fish discriminated between nearby rich and poor patches, but not between rich and poor micropatches within a food patch. This suggests that the foraging scale of benthic fish lies between the patch and micro patch scale in our experiments. We conclude that patch use behaviour of benthic fish can provide a powerful measure of habitat quality that reveals how benthic fish perceive their environment.