Risk Sensitivity in Foraging Jack Dempsey Cichlids (Cichlasoma octofasciatum)

Studies have documented that animals with positive energy budgets tend to prefer feeding sites offering constant amounts of food over those offering variable amounts of food. The present study tested whether Jack Dempsey cichlids (Cichlasoma octofasciatum) in a two-patch environment preferred the patch offering a constant amount of food over the one offering a variable amount of food. The study also examined (1) whether fish displayed shifts in risk sensitivity when they were allowed only one choice per trial (discrete-choice treatment), as opposed to when they could freely sample both alternatives (free-choice treatment), and (2) whether or not fish displayed shifts in risk sensitivity between the first trial of the day, when they were hungrier, and the final trial of the day, when they were more satiated. Two of six fish significantly preferred the constant alternative and one showed a numerical preference for the constant alternative that approached significance. The mean proportion of first choices to the constant side did not differ significantly between the discrete-choice treatment and the free-choice treatment. In the discrete-choice treatment, however, fish were significantly more risk averse in the first trial of daily sessions than in the final trial — a shift in risk sensitivity opposite to that seen in a number of other studies. This suggests that animals may pass through more than one threshold in risk sensitivity on different points along an energy budget continuum.     

Risk sensitivity in starlings: variability in food amount and food delay

Starlings' preferences for constant versus variable food sourceswere studied in the laboratory. The constant alternative gavea fixed amount of food after a fixed delay. The variable alternativeoffered either a varying amount of food after a fixed delay(treatment A) or a fixed amount of food after a variable delay(treatment B). In both treatments the ratio of amount of foodover trial length (the sum of intertrial interval plus delayand handling times) of the constant alternative equaled theaverage of the two ratios of the variable alternative. The variableratios were 30% higher and 30% smaller than the fixed ratio.In free-choice trials (both options available in each trial),the subjects were risk-averse or indifferent in treatment Aand indifferent or riskprone in treatment B. In no-choice trials(only one source available per trial), the latency to respondwas longer in the variable than in the constant source in treatmentA and the opposite in treatment B. The greater preference forvariability in time than for variability in reward amount isnot consistent with either maximizing the ratio of expectedenergy over expected time or the expected ratio of energy overtime for individual trials. There was a negative correlationbetween individual intake rate and degree of risk pronenessfor both kinds of variability. We present a model of choicebased on an information-processing theory for temporal memorythat accounts for the different effects of variability in delayand in amount but cannot explain the effects of intake rate.[Behav Ecol 1991;2:301–308]     

Influence of budget and reinforcement location on risk-sensitive preference

Little is known about the effect that procedural variables have on risk-sensitive preference. This study assessed the effect of procedural variables on pigeons' choice between a fixed and variable amount of reinforcement (amount risk) and, in a separate condition, between a fixed and variable delay until reinforcement (delay risk). Experiment 1 investigated the impact of water reinforcement and risk dimension when pigeons were in a restrictive budget, where access to water was less than that necessary to maintain current body weight, and a condition where the pigeons had ample access to water. Pigeons exhibited a greater tendency to prefer the variable alternative for delay risk than for amount risk in both restrictive and ample budgets. Varying water budget had no effect on risk preference. Experiment 2 investigated the influence of water reinforcer location while in a restrictive budget, in which reinforcers were delivered to a single location, two distinct locations, or a randomly selected location. With amount risk, pigeons were risk averse when reinforcers were delivered in separate or random locations and were indifferent to risk when delivered to a single location. With delay risk, pigeons were generally risk prone with no effect from reinforcement location. The finding that pigeons were risk averse when reinforcers were delivered to separate locations and were indifferent to risk when delivered to a single location offers a methodological explanation to the inconsistent findings in the literature with amount risk.     

Modeling Global Macroclimatic Constraints on Ectotherm Energy Budgets

We describe a mechanistic individual-based model of how globalmacroclimatic constraints affect the energy budgets of ectothermicanimals. The model uses macroclimatic and biophysical charactersof the habitat and organism and tenets of heat transfer theoryto calculate hourly temperature availabilities over a year.Data on the temperature dependence of activity rate, metabolism,food consumption and food processing capacity are used to estimatethe net rate of resource assimilation which is then integratedover time. We detail and explore the significance of assumptionsused in these calculations. We present a new test of this model in which we show that thepredicted energy budget sizes for 11 populations of the lizardSceloporus undulalus are in close agreement with observed resultsfrom previous field studies. This demonstrates that model testsare feasible and the results are reasonable. Further, sincethe model represents an upper bound to the size of the energybudget, observed residual deviations form explicit predictionsabout the effects of environmental constraints on the bioenergeticsof the study lizards within each site that may be tested byfuture field and laboratory studies. Three major new improvements to our modeling are discussed.We present a means to estimate microclimate thermal heterogeneitymore realistically and include its effects on field rates ofindividual activity and food consumption. Second, we describean improved model of digestive function involving batch processingof consumed food. Third, we show how optimality methods (specificallythe methods of stochastic dynamic programming) may be includedto model the fitness consequences of energy allocation decisionssubject to food consumption and processing constraints whichare predicted from the microclimate and physiological modeling Individual-based models that incorporate macroclimatic constraintson individual resource acquisition, assimilation and allocationcan provide insights into theoretical investigations about theevolution of life histories in variable environments as wellas provide explicit predictions about individual, populationand community level responses to global climate change.     

Energy budget versus temporal discounting as determinants of preference in risky choice

Four pigeons and three ringneck doves responded on an operant simulation of natural foraging. After satisfying a schedule of reinforcement associated with search time, subjects could "accept" or "reject" another schedule of reinforcement associated with handling time. Two schedules of reinforcement were available, a variable interval, and a fixed interval with the same mean value. Food available in the session (a variable related to the energy budget) was manipulated in the different conditions either by increases of the value of the search state schedule of reinforcement, or by increases in the mean value of the handling state schedules. The results indicate that the amount of food available in the session did not affect the preference for variable schedules of reinforcement, as would be predicted by an influential theory of risk sensitive foraging. Instead, the preference for variability depended on the relationship between the time spent in the search and the handling states, as is predicted by a family of models of choice that are based on the temporal proximity to the reinforcer.     

Cattle use visual cues to track food locations

We tested the hypothesis that cattle aided by visual cues would be more efficient than uncued animals in locating and consuming foods placed in either fixed or variable locations within a 0.64-ha experimental pasture. Eight yearling steers were randomly selected and trained to associate traffic barricades and traffic cones with high- (oat-barley mixture) and low- (straw) quality foods, respectively. Initially steers were randomly assigned to 1 of 4 food location/visual cue treatments: fixed locations/with cues (F/C), variable locations/with cues (V/C), fixed locations/no cues (F/NC), or variable locations/no cues (V/NC). High- and low-quality foods and their respective cue (or no cue) were placed in the experimental pasture. Individual animals were allowed to explore the pasture for 10 min twice per day every other day for 1 week. Minutes until feeding, first feed type consumed (i.e., high-quality, low-quality, or no food consumed), animal location and activity (i.e., feeding, standing, or moving), and total intake of high- and low-quality feed were recorded during each 10-min trial. At the end of each week, location/visual cue treatments were randomly assigned to another 2 steers, which permitted an independent test of each animal in each treatment over a 4-week period. Animals in the F/C and V/C treatments took about 2 min to initially locate and consume a food, compared to F/NC and V/NC animals who took nearly 4 and 6 min, respectively. The high-quality food was the first food located and consumed by F/C, V/C, F/NC, and V/NC animals during 79, 77, 67, and 54% of sampling occasions, respectively. Cued animals typically spent more time feeding (P=0.0004) and less time standing (P=0.005) than uncued animals. Cued animals had a higher mean intake than uncued animals of high- (P=0.001) and low- (P=0.04) quality food. Mean high-quality intake for F/C, V/C, F/NC, and V/NC treatments was 266, 245, 214, and 126 (+/-22) g, respectively; mean low-quality intake for the same treatments was 36, 32, 12, and 10 (+/-10) g. Cued animals also consumed more food per distance traveled than uncued animals (P=0.005). Animals located food quicker (P=0.03) and consumed more high-quality food (P=0.02) when food locations were constant than when they were variable. Our data strongly indicate that cattle can learn to associate visual cues with disparate food qualities and use this information to forage more efficiently in both fixed and variable foraging environments.     

Energy requirements of a Great white pelican (Pelecanus onocrotalus) population and its impact on fish stocks

The Great white pelican Pelecanus onocrotalus is a major predator of some freshwater fish stocks in much of Africa. Detailed population time-activity budgets allowed us to estimate that the south-western Cape Province population consumed 184 tonnes offish annually. Of this, 69% was required for adult existence, 17% for chicks and 14% for flight. Per capita consumption averaged 840 g per day, or 8–8% of body weight; very much less than the figure of 17–6% used in some energy budget calculations. Our integrative bioenergetics approach lends some support for the use of 10% of body weight as the amount of food consumed per day by large piscivorous birds for initial estimates of their impact on stocks.
Predation by Great white pelicans may account for some 10–25% offish production, a shortage of reliable data on fish production in African freshwater ecosystems makes a more precise estimate impossible. Food supplies and shoreline length may both be important factors for pelican populations.     

建鲤和异育银鲫摄食不同质量饲料时的氮收支和能量收支比较

实验探讨了建鲤和异育银鲫摄食低质和高质饲料时氮和能量的收支情况.低质饲料以豆粕为主要蛋白源,饲料蛋白含量为33.91%,高质饲料以鱼粉为主要蛋白源,饲料蛋白含量为45.59%.55d的生长结果显示,氮收支和能量收支受到饲料质量和鱼类种类的显著影响:摄食低质饲料时,建鲤的生长氮和生长能比例显著低于异育银鲫,排泄氮、排泄能和代谢能比例显著高于异育银鲫;摄食高质饲料时,两种鱼的氮收支和能量收支无显著差异;建鲤的氮收支和能量收支受饲料质量的显著影响,摄食低质饲料时,其生长氮和生长能比例均显著低于摄食高质饲料时,而排泄氮、粪能和代谢能比例均显著高于摄食高质饲料时;异育银鲫的氮收支、生长能和代谢能比例不受饲料质量的显著影响.结果表明,在低质饲料条件下,建鲤利用氮和能量的能力弱于异育银鲫,在高质饲料条件下,两种鱼没有显著差异.与异育银鲫相比,建鲤利用氮和能量的能力受饲料质量的影响更为显著.       

食物质量差异对树麻雀能量预算和消化道形态特征的影响

为比较小型鸟类对不同食物的能量预算策略,本文对3组树麻雀(Passer monanus)进行2周的食物质量差异性驯化(Acclimation)。结果表明,相同和较低的生存压力下,树麻雀体重、体温、体脂含量和水分含量表现出显著的组间差异。高能食物组摄入能和可消化能增加极显著,消化率和同化率增加不显著,体内能量储备增加;低能食物组摄入能、可消化能、消化率和同化率降低极显著,体内能量储备消耗极显著。小肠和总消化道长度以及肝脏重量出现极显著的表型可塑性响应。体内能量储备增减的预算,能量摄入和能量转化器官的功能能力与自身能耗之间的预算,以及摄食低能食物时的节能预算都是树麻雀能量预算策略的重要组成部分。总之,树麻雀对不同含能食物采用不同的能量预算策略,器官水平的表型可塑性(phenotypic plasticity)响应是个体能值出现显著变化的基础,也是个体能量预算策略成功的关键。对不同食物采用不同的能量预算策略是树麻雀重要的生存对策。     

The effects of reward quality on risk-sensitivity in Rattus norvegicus

Risk-sensitive foraging theory (RSFT) was developed to explain a choice between a variable (risk-prone) or constant (risk-averse) option. In the RSFT literature, qualitative shifts in risk-sensitivity have been explained by fluctuations in daily caloric energy budget (DEB). The DEB rule describes foragers’ choices as being based on fitness and rate of gain. If the DEB rule is correct, rewards that differ in caloric returns should cause differences in foragers’ sensitivity to risk. However, few studies have explored the influence of reward quality on risk-sensitivity in mammals. The present study was designed to examine the effects of reward quality on risk-sensitivity when reward magnitude, delay to reward, body mass, and response effort were controlled. Results from the current study demonstrated that subjects rewarded with a high calorie reward (i.e., sugar) made significantly fewer choices for a variable option than subjects rewarded with a lower calorie reward (i.e., grain). These results are consistent with the predictions of the DEB rule, and add to the RSFT literature where reward quality was manipulated by describing difference in risk-sensitivity in mammals. Suggestions for future research include an examination of risk-sensitivity where flavor and caloric return are manipulated.     

Carpenter Bee (Xylocopa micans) Risk Indifference and a Review of Nectarivore Risk-sensitivity Studies

This paper presents new results of risk-sensitive foraging studiesof the carpenter bee, Xylocopa micans, and reviews the workto date on risk sensitivity in nectarivores. In the field, nectarivoreschoose among alternative food sources (flowers) that differin the variabilities of their nectar rewards. In the lab, theforaging situation for carpenter bees was experimentally simplifiedby offering the bees a choice between either "low variance"or "high variance" artificial flowers. The two flower typesdiffered in their variabilities but offered the same expectedshortterm rates of net energy gain to test the predictions ofthe short-term rate maximization mechanism. Foragers were testedunder two energy budget conditions, hungry and well-fed, totest the predictions of the z-score model. Individual carpenterbees were indifferent to variability in both nectar volume andnectar sugar concentrations, and their risk-indifference wasunaffected by energy budget. These findings of risk indifferencesupport neither the variance discounting nor the z-score modelof risk sensitivity. Since the low and high variance flowertypes are equivalent for carpenter bees in short-term rate ofenergy gain, there can be no selection on carpenter bees tobe sensitive to variability based on differences in rate ofgain. Studies of risk sensitivity in honey bees and bumble beesusing variance in nectar concentration support this contention.These findings are compared with other nectarivore risk sensitivitystudies in order to highlight the most likely mechanisms underlyingaversion to variation in nectar rewards (short-term rate maximizing,the Weber-Fechner law of perception and learning non-empty flowers)and to suggest future research in the interplay of these threemechanisms.     

Effects of temporal clumping and payoff accumulation on impulsiveness and cooperation

Animals show impulsiveness when they prefer a smaller more immediate option, even though a larger more delayed option produces a higher intake rate. This impulsive behavior has implications for several behavioral problems including social cooperation. This paper presents two experiments using captive blue jays (Cyanocitta cristata) that consider the effects of payoff accumulation and temporal clumping on impulsiveness and cooperation. Payoff accumulation refers to a situation where the benefits gained from each choice trial accumulate from one trial to the next, and only become available to the animal after it has completed a fixed number of trials. We hypothesized that this would reduce impulsiveness because it removes the advantage of quickly realizing food gains. Clumping refers to situation in which the animal experiences several choice trials in quick succession followed by a long pause before the next clump. We hypothesized that if payoffs accumulated over a clump of trials this would enhance the effect of accumulation. We tested the effects of accumulation and clumping on impulsiveness in a self-control situation. We found a significant interaction between clumping and accumulation. Payoff accumulation reduced impulsiveness, but only when trials were clumped. Post hoc analyses suggest that clumping alone increases impulsiveness. A second experiment applied these results to cooperation. This experiment reveals an interaction between payoff accumulation and trial's position within the clump. Jays were more likely to cooperate on the first trial of a clump, but the likelihood of cooperation dropped after the first trial. However, this drop was larger when payoffs did not accumulate. This observation suggests that the difference between accumulated and un-accumulated treatments that we reported previously may be largely due to differences in how animals behave in the first trial of a clump.     

Optimal risky choice in humans: effects of amount of variability

The present study investigated whether the predictions of an optimal risk-sensitive foraging model (the energy-budget rule) would extend to humans’ choices between high- and low-variance monetary response options. In Experiment 1, participants were presented with repeated choices between two options that had the same mean values but different variances across positive- and negative-budget conditions. In Experiment 2, a within-subject comparison was conducted to investigate choice under positive- and negative-budget conditions when the options were either (a) a fixed option and a high-variance option, or (b) a low-variance option and a high-variance option. Session-wide choices were analyzed in relation to the predictions of the energy-budget rule and sequential choices were analyzed with dynamic optimization modeling. When both options were variable choice was generally consistent with predictions of the energy-budget rule and was more risk prone under negative-budget than positive-budget conditions. Sequential choices were sensitive to local budget conditions, but choices were less consistent with optimality when both options were variable, possibly because of the greater similarity in expected earnings for optimal and nonoptimal choices in these conditions. Overall, the results provide further evidence that the energy-budget rule may have broad applicability and that it can extend to human risky choice between multiple variable response options.     

A study of the food and feeding of perch, Perca fluviatilis L., in Windermere

SUMMARY. Perch were sampled for their stomach contents at regular intervals throughout 24 h from June until October in 1973 and from February 1975 until January 1976. They were found to feed on benthic organisms from November until April, on benthos and plankton during May and June and on perch fry and zooplankton from July until October. Perch over a wide size range feed on similar prey. Fish showed great variability in the weight of their stomach contents. A method based on a points system was developed to estimate the weight of food in the stomach for a given weight of fish at a known time. A diel feeding pattern which varied with the season was apparent from these data. Rates of gastric evacuation were assumed to be exponential and were calculated from the drop in night-time stomach content weights when food intake was assumed to be zero. The rates ranged from 0.18 mg h⁻¹ at a mean water temperature of 11°C in May to 0.35 mg h⁻¹ at a mean water temperature of 17°C in July. Assuming that food consumption followed a linear rate of intake, the standard Bajkov method was considered an adequate model to calculate daily food consumption. Daily food consumption (mg dry weight) was calculated for 150 g perch for all months of the year (November to April and September and October being combined). A second series of values was calculated making corrections for the time spent in the fishing gears when food intake was zero but gastric evacuation continued. Daily food consumption figures for 150g male perch based on Winberg's hypotheses (1956) and growth data showed no significant differences from this second series of values, when both were expressed in energy terms. When certain assumptions have been tested, growth data and Winberg's equations together may be a suitable method for calculating an energy budget for the Windermere perch population.     

Food restriction and refeeding in growing rats.

1. The interrelationship between food intake, body weight and oxygen consumption was analysed at 25 degrees C in growing rats. 3. The experiment was divided into two phases lasting four weeks each. During the first phase the animals were subjected to energy restriction and during the second phase they were allowed ad lib energy intake. Four groups of rats were studied: Control with ad lib food intake and three restricted groups R4, R5, and R6 which received during the first phase 4, 5, and 6 g per day of stock diet in a single meal. 3. The results showed a decrease in body weight and oxygen consumption during the restriction period and a recovering of the latter during the refeeding period, without body weight recovering. 4. It is concluded that an energy conservation mechanism is present during food restriction and for some time after refeeding.     

Risk-sensitive foraging in rats: the effects of response-effort and reward-amount manipulations on choice behavior

The literature on risk-sensitive foraging theory provides several accounts of species that fluctuate between risk-averse and risk-prone strategies. The daily energy budget rule suggests that shifts in foraging strategy are precipitated by changes in the forager's energy budget. Researchers have attempted to alter the organism's energy budget using a variety of techniques such as food deprivation, manipulation of ambient temperatures, and delays to food reward; however, response-effort manipulations have been relatively neglected. A choice preparation using a wheel-running response and rats examined risk-sensitive preferences when both response effort and reward amounts were manipulated. Concurrently available reinforcement schedules (FI/60 and VI/60) yielded equivalent food amounts per unit time in all treatments. Two levels of response effort (20 or 120 g tangential resistance) and two levels of reward amount (three or nine pellets) were combined to form four distinct response-effort/reward-amount pairings. Increasing reward amounts significantly shifted choice toward the FI schedule in both response-effort conditions. The incidence of choice preference and the magnitude of shifts in choice were greater for the high response-effort conditions than for the low response-effort conditions. Implications of the significant interaction between response effort and reward amount are discussed in terms of a general energy-budget model.     

随机限食和重喂食小鼠能量收支和生长发育的可塑性

为阐明能量收支和生长发育的可塑性对动物适应食物资源变化的作用和意义,将断乳后的雄性KM小鼠40只随机限食4周,再重喂食4周。采用封闭式流体压力呼吸计测定基础代谢率(BMR)。限食使摄食量显著增加,BMR和活动行为降低,胴体和生殖腺重量显著降低。重喂食后上述指标均恢复到对照组水平,表现出显著的可塑性变化。结果表明,KM小鼠能通过摄食量、BMR、活动行为和身体组成的可塑性调节以适应难以预测的食物资源变化。     

温度对台湾红罗非鱼能量收支的影响

于１９９６年７－８月在盐度为１４的条件下,测定了台湾红罗非鱼（体重２５．３３４－２６．２２５ｇ）在水温为２２、２８和３４℃时的能量收支,结果表明,温度对台湾红罗非鱼的特定生长率,转化效率和最大摄食率均有显著影响,２８℃时上述各值均达到最大,分别为１．７９,６０．０１％和２．５９％Ｂ．Ｗ．ｄ＾－１;对吸收率无显著影响,对于物质的吸收率的平均值为６４．５８％、温度对食物能分配于能量支出各部分的比例有显     

Partitioning and utilization of energy during the larval development of the xanthid crab, Rhithropanopeus harrisii (Gould)

An energy budget is constructed for the larval development of the crab Rhithropanopeus harrisii (Gould) fed nauplii of the brine shrimp Artemia salina (L.). Between the first zoeal instar and the megalopa, there is a 5.4-fold increase in caloric consumption and a 13.2-fold increase in dry weight. Weight specific energy content increases through the zoeal stages and drops at the megalopa. Rate of oxygen consumption increases steadily throughout development. Assimilation, gross growth, and net growth efficiencies increase steadily through zoeal development and drop at the megalopa. Calories put into tissue production exceed those expended via respiration in zoeal stages II–IV; the reverse is true in zoeal stage I and the megalopa.

A total energy budget has been calculated and the partitioning of energy is discussed in relation to other physiological studies on this species.     

Rhinopithecus bieti at Tacheng,Yunnan: Diet and Daytime Activities

We collected data on diet and activity budget in a group of Rhinopithecus bieti at Tacheng (99°18E, 27° 36'N, between 2,700–3,700 m asl), Yunnan, from March 1999 to December 2000. We mainly recorded species-parts eaten with feeding scores from scanning state behaviors of one-male units in tree-crowns. We also conducted microscopic analysis of feces collected monthly. The subjects consumed 59 plant species, belonging to 42 genera in 28 families, of which 90 species-parts were distributed as follows: 21 in Winter, 38 in spring, 39 in Summer, 47 in autumn. Conversely, the group annually spent, on average, 35% of daytime feeding, 33% resting, 15% moving, and 13% in social activities. Seasonal changes are apparent in daytime budget and food item-related feeding time in tree-crowns, food remains in feces, and the number of species-parts eaten. Correlations within and between food items and time budget clearly indicate maximization of foraging effectiveness and minimization of energy expenditure. In consideration of reports from northern and southern groups, that which underlay the specific adaptation to the habitat appeared to be similar to those of other colobines. Thus, the ultimate factors for survival of the species are more hopeful than expected.