Energy Intake Rate Research Articles

Limits to sustained energy intake. I. Lactation in the laboratory mouse Mus musculus.

Laboratory mice (strain MF1) were used to determine whether sustainable rates of energy intake are limited during lactation. Mice raising natural-sized litters (N=71) reached an asymptote in their daily food intake between days 13 and 16 of lactation at 23.1 g x day(-1) and also between litter sizes of 9 and 15 pups (22.8 g x day(-1)). A second group of 37 females had their litter sizes manipulated at birth to raise more or fewer offspring than they gave birth to. When the litter size was increased, females did not increase their food intake to match their new litter size. However, when litter size was decreased, females decreased their asymptotic daily food intake during late lactation in relation to the extent of reduction in litter size. Therefore, it appeared that females were limited during late lactation and with large litter sizes. The milk energy exported amounted to 44% of the gross energy intake, and the estimated daily energy expenditure was therefore considerably lower than the sustained energy intake [8.0xRMR(gross), 6.6xRMR(assimilated)], and averaged 3.1xRMR, where RMR is resting metabolic rate. It was not possible to determine whether the apparent limit on sustained energy intake was acting centrally or peripherally because of the asymptotes in both food intake and milk energy output with increasing litter size.

Spatial Variation in Tuber Depletion by Swans Explained by Differences in Net Intake Rates

We tested whether the spatial variation in resource depletion by Tundra Swans (Cygnus columbianus) foraging on belowground tubers of sago pondweed (Potamogeton pectinatus) was caused by differences in net energy intake rates. The variation in giving-up densities within the confines of one lake was nearly eightfold, the giving-up density being positively related to water depth and, to a lesser extent, the silt content of the sediment. The swans' preference (measured as cumulative foraging pressure) was negatively related to these variables. We adjusted a model developed for diving birds to predict changes in the time allocation of foraging swans with changes in power requirements and harvest rate. First, we compared the behavior of free-living swans foraging in shallow and deep water, where they feed by head-dipping and up-ending, respectively. Up-ending swans had 1.3–2.1 times longer feeding times than head-dipping swans. This was contrary to our expectation, since the model predicted a decrease in feeding time with an increase in feeding power. However, up-ending swans also had 1.9 times longer trampling times than head-dipping swans. The model predicted a strong positive correlation between trampling time and feeding time, and the longer trampling times may thus have masked any effect of an increase in feeding power. Heart rate measurements showed that trampling was the most energetically costly part of foraging. However, because the feeding time and trampling time changed concurrently, the rate of energy expenditure was only slightly higher in deep water (1.03–1.06 times). This is a conservative estimate since it does not take into account that the feeding costs of up-ending are possibly higher than that of head-dipping. Second, we compared captive swans foraging on sandy and clayey sediments. We found that the harvest rate on clayey sediment was only 0.6 times that on sandy sediment and that the power requirements for foraging were 1.2–1.4 times greater. Our results are in qualitative agreement with the hypothesis that the large spatial variation in giving-up densities was caused by differences in net rates of energy intake. This potentially has important implications for the prey dynamics, because plant regrowth has been shown to be related to the same habitat factors (water depth and sediment type).

Under-reporting in dietary surveys – implications for development of food-based dietary guidelines

To illustrate issues related to under-reporting in dietary surveys relevant for development of food-based dietary guidelines. Food and nutrient intakes in two national dietary surveys are analysed in relation to under-reporting using the concept of cut-off values for the ratio between energy intake and basal metabolic rate as well as biochemical markers of nutritional status. Subjects with an implausibly low EI/BMR ratio had dietary patterns more close to recommended levels, e.g. lower fat intake and higher fibre intake, than those with acceptable energy intakes. Intakes of micronutrients were lower. This was due to both general and selective under-reporting of foods. Under-reporting was due to less frequent consumption, larger proportion of non-consumers and smaller portion sizes. Low EI/BMR ratios did not influence the characterisation of protein and micronutrient intake of low and high consumers of fruit and vegetable intake. Under-reporting can influence the usefulness of dietary data as a tool for developing food-based dietary guidelines, and the extent of under-reporting should be assessed using appropriate methods.

Examining feeding strategies and position choice of drift-feeding salmonids using an individual-based, mechanistic foraging model

We demonstrated the ability of a mechanistic habitat selection model to predict habitat selection of brown trout (Salmo trutta) and mountain whitefish (Prosopium williamsoni) during summer and winter conditions in the Blacksmith Fork River, Utah. By subtracting energy costs and losses from the gross energy intake rate (GEI) obtained through simulation of prey capture, the model calculates the potential net energy intake rate (NEI) of a given stream position, which is essentially the rate of energy intake available for growth and reproduction. The prey capture model incorporates the size, swimming speed, and reaction distance of the fish; the velocity, depth, temperature, and turbidity of the water; and the density and size composition of the drifting invertebrates. The results suggest that during both summer and winter, the brown trout and mountain whitefish in our study reach avoided locations providing low NEI and preferred locations providing a high ratio of NEI to the swimming cost (SC) at the focal position of the fish (NEI/SC). This supports the idea that the drift-feeding fish in this study selected stream positions that provided adequate NEI for the least amount of swimming effort.

Foraging behaviour of brent geese, Branta b. bernicla, on grasslands: effects of sward length and nitrogen content.

Feeding behaviour and preferences of brent geese, Branta b. bernicla were observed on pastures of different sward lengths and nitrogen contents. On swards of 2.0-6.0cm captive geese took larger bites and had a higher intake rate when feeding on 6-cm swards than when feeding on shorter ones. In the field wild geese chose 6-cm swards in preference to both shorter and longer ones. For unfertilized pastures there was a strong negative relationship between nitrogen content and sward height: 11-cm swards contained 2.8% nitrogen, 3.5-cm swards 4.2% nitrogen. Application of 75kgNha-1 organically based fertilizer at the end of October eliminated this relationship between nitrogen content and sward height, swards of all heights then having a mean content of 4.1% N. On fertilized plots the geese preferred swards longer than 6cm with no indication of a decrease in preference up to the maximum height investigated, 16cm. Breaking strain of grass laminae was measured to give an indication of the proportion of strengthening tissues in the leaves and hence their digestibility. Apical laminae from longer swards had a higher breaking strain than those from shorter swards. Unfertilized swards had a higher breaking strain than fertilized swards but the difference in breaking strain between long and short swards was the same on fertilized and control treatments. These results are discussed in relation to the forage maturation hypothesis and are interpreted as indicating that the primary constraint on maximizing energy intake rates is not the reduced energy digestibility of older foliage but the reduced nitrogen content in the higher-biomass swards. We conclude that it is the balance between maximizing energy intake and nitrogen absorption rates which results in the observed preference for intermediate-height swards.

EVOLUTION OF INTRINSIC GROWTH AND ENERGY ACQUISITION RATES. II. TRADE-OFFS WITH VULNERABILITY TO PREDATION IN MENIDIA MENIDIA

The Atlantic silverside (Menidia menidia) exhibits countergradient latitudinal variation in somatic growth rate along the East Coast of North America. Larvae and juveniles from high-latitude populations display higher intrinsic rates of energy consumption and growth than genotypes from low-latitude populations. The existence of submaximal growth in some environments suggests that trade-offs must counter the oft-cited theoretical benefits of energy and growth maximization (e.g., “bigger is better,” “faster is better”) in the immature life stages. We hypothesized that energy and growth maximization trades off against investment in defense from predators. We conducted laboratory selection experiments to compare vulnerability to predation of silversides from: (1) fast-growing northern (Nova Scotia, NS) versus slow-growing southern (South Carolina, SC) source populations; (2) phenotypically manipulated fast-growing versus moderately-growing NS fish; and (3) recently fed versus unfed NS and SC fish. Tests involved fish drawn from common-garden environments and were conducted by subjecting mixed-treatment schools of size-matched silversides to natural, common piscine predators. NS silversides suffered significantly higher predation mortality than SC silversides. Parallel results were found in phenotypic manipulation of growth: NS silversides reared on a fast-growth trajectory (∼1.0 mm/day) were significantly more vulnerable to predation than those growing at a moderate rate (∼0.5 mm/day). Food consumption also affected vulnerability to predators: Silversides with large meals in their stomachs suffered significantly higher predation mortality than unfed silversides. Differences in predation vulnerability were likely due to swimming performance, not attractiveness to predators. Our findings demonstrate that maximization of energy intake and growth rate engenders fitness costs in the form of increased vulnerability to predation.Corresponding Editor: D. Roff

BIASES IN THE FIELD ESTIMATION OF SHOREBIRD PREY SIZES

Several studies have attempted to calculate the energy intake rates of shorebirds. One method of doing this involves estimating the size of prey items in relation to the bird's bill length, and then using an average bill length to convert this value to actual prey length in order to estimate prey mass/energy content. Although this technique is used regularly, no study has assessed under what conditions the method is reliable. Observer distance from the bird, prey type, size and orientation in the bill, bill length and shape, and light conditions may all affect the accuracy of the estimates. This study sought to establish the conditions under which the method is or is not reliable, using three different prey types (“thin” and “fat” polychaetes, and crabs) and a long- and short-billed bird. It was found that if observers were calibrated, and their results were corrected using regression analysis, errors in estimates of food intake were much reduced. The time allowed to view the prey significantly affected the error in the result, with shorter viewing time causing greater errors, and observations made into the light were less accurate than those made with the light behind the observer. Large prey types, or even larger size classes of the same species, are estimated less accurately than small prey, and accuracy decreases as prey size increases as multiples of bill length. If separate calibration regressions are generated for large and small prey, food intake estimates are more accurate than if a single regression is used incorporating the full range of prey sizes. Sexual dimorphism in bill length can also increase error in the results.

State dependent behavior and the Marginal Value Theorem

The Marginal Value Theorem (MVT) is the dominant paradigm in predicting patch use and numerous tests support its qualitative predictions. Quantitative tests under complex foraging situations could be expected to be more variable in their support because the MVT assumes behavior maximizes only net energy-intake rate. However across a survey of 26 studies, foragers rather consistently ‘‘erred’’ in staying too long in patches. Such a consistent direction to the errors suggests that the simplifying assumptions of the MVT introduce a systematic bias rather than just imprecision. Therefore, I simulated patch use as a statedependent response to physiological state, travel cost, predation risk, prey densities, and fitness currencies other than net-rate maximization (e.g., maximizing survival, reproductive investment, or mating opportunities). State-dependent behavior consistently results in longer patch residence times than predicted by the MVT or another foraging model, the minimize /g rule, and these rules fail to closely approximate the best behavioral strategy over a wide range of conditions. Because patch residence times increase with state-dependent behavior, this also predicts mass regulation below maximum energy capacities without direct mass-specific costs. Finally, qualitative behavioral predictions from the MVT about giving-up densities in patches and the effects of travel costs are often inconsistent with state-dependent behavior. Thus in order to accurately predict patch exploitation patterns, the model highlights the need to: (1) consider predator behavior (sit-and-wait versus actively foraging); (2) identify activities that can occur simultaneously to foraging (i.e., mate search or parental care); and (3) specify the range of nutritional states likely in foraging animals. Future predictive models of patch use should explicitly consider these parameters.

Vulnerability to severe weather and regulation of body mass of Icelandic and British Redshank Tringa totanus

During severe weather, Redshanks suffer the heaviest mortality amongst all the shorebird species wintering around the North Sea coasts of the British Isles. An earlier study had suggested that this resulted from a failure to accumulate sufficient body fat reserves before mid‐winter. Detailed field studies in northeast England between 1993 and 1995 of seasonal changes in body mass, and in estimated lean and fat masses, of two races of Redshank, both of which winter in the same estuary, were accompanied by similar studies of small numbers held in captivity with unlimited food. After differences in body size were allowed for, there were no differences in body composition and its seasonal pattern of change in birds of the Icelandic and British races. Body mass changes in wild birds paralleled those in captives between November and March, and mid‐winter levels were not limited by food supply; indeed they were slightly higher in a winter with lower prey densities. It is concluded that Redshanks regulate body mass and, indirectly, fat reserves at levels set by a trade‐off between the risks of predation and starvation. Unlike most other shorebird species, they take very small prey in relation to their body size and hence must feed for long periods during each tidal cycle to achieve their daily energy intake needs. Thus they have little scope to extend their feeding time during severe weather, which also forces them to feed on ice‐free exposed coastal habitats where wind chill cannot be avoided. Both factors lead to more rapid depletion of fat reserves than in other species which have higher energy intake rates or lower total daily requirements.

Seed selection in the Java Sparrow (Padda oryzivora): preference and mechanical constraint

Very few studies address the effect of hardness on seed selection in granivorous birds. As a defense against predators, plant species may produce seeds of various hardnesses, some of which are too hard for a bird to crack. Unsuccessful cracking attempts lead to loss of time and, thus, lowers energy-intake rate. Birds may prefer seeds with a short handling time and a large chance of being cracked. However, without knowing the maximal cracking force of the bird, it is difficult to distinguish between seed selection as a result of mechanical constraints and selection as a result of preference. Our experiments aimed to discriminate between these two effects. During two series of experiments, Java Sparrows (Padda oryzivora) were offered safflower seeds. Size characters and hardness of the seeds that remained after feeding were compared with a control group. Without prior experience, the birds showed selection as a result of mechanical constraints. Seeds were chosen randomly and only seeds with a hardness less than the maximal crushing force were eaten, with the rest being rejected. After some experience, birds started to actively select for seed size (e.g., depth) and preferred to eat the smallest seeds. Although the correlation between size and hardness is low, the birds successfully used size characteristics as a predictor for hardness.

The energy budget of captive Siberian hamsters, Phodopus sungorus, exposed to photoperiod changes: mass loss is caused by a voluntary decrease in food intake.

Unlike most nonhibernating small mammals, Siberian hamsters (Phodopus sungorus) undergo a pronounced mass loss, almost exclusively as white adipose tissues, in response to a switch from long- to short-photoperiod exposure. This mass loss can be caused by an increase in the rate of energy expenditure, a decrease in the rate of energy intake, or a decrease in assimilation efficiency. In order to determine how they relate to photoperiod-induced mass loss, we measured these energy budget components every 2 wk on 12 captive Siberian hamsters exposed to 8 wk of long photoperiod followed by 12 wk of short photoperiod. Body mass decreased shortly after short-photoperiod exposure. This was accompanied by an immediate decrease in the rate of energy intake and, after a 2-wk delay, by a decrease in the rate of energy expenditure. The overall cumulative decrease in energy intake (376 kJ) could account for the mass loss (249 kJ) observed during short-photoperiod exposure. Regression analyses indicated that only the rate of energy intake was significantly related to the rate of mass change. Therefore, we conclude that photoperiod-induced mass loss in Siberian hamsters is caused by a decrease in the rate of food intake, rather than by changes in energy expenditure or in assimilation efficiency.

Ingestion patterns and daily energy intake on a sugary diet: the Red Lory Eos bornea and the Malachite Sunbird Nectarinia famosa

This study examines whether nectarivorous birds regulate daily energy intake as proposed by Lloyd in 1991. Two Old World nectarivorous species, a large non‐passerine, the Red Lory Eos bornea, and a small passerine, the Malachite Sunbird Nectarinia famosa, were fed 0.25 mol/1 sucrose (9%), 0.73 mol/1 sucrose [24%] or 0.73 mol/1 glucose in separate laboratory trials to determine hourly and overall daily rate of sugar intake and consumption. Overall daily energy intake rates of the Malachite Sunbird and the Red Lory support the hypothesis of regulated energy intake for nectarivorous Old World birds. However, the species differ in their ingestion patterns through the day. The Red Lory ingests large volumes initially, regardless of sugar type. This is possibly a consequence of their large size and having a crop in which food can be stored. The Malachite Sunbird showed more regular hourly consumption through the day, and no change in mass during the day. Intake rates of both species were greater on lower sugar concentrations.

Reproducibility of energy and macronutrient intake and related substrate oxidation rates in a buffet-type meal

The aim of this present study was to determine the reliability of a buffet-type meal as a measure of spontaneous energy and macronutrient intake. In addition, we evaluated the short-term effects of diet on the composition of the substrate mix oxidized postprandially. Fourteen male subjects had ad libitum access to a variety of foods from a buffet-type meal offered in the laboratory during two identical sessions. The foods comprising the test meal had varying amounts of protein, lipid and carbohydrate. The results showed that there were significant intraclass correlations (ri) for energy (ri 0.97, P = 0.0001), lipid (ri 0.97, P = 0.0001), carbohydrate (ri 0.92, P = 0.0003) and protein (ri 0.82, P = 0.0072) intake between the two meal sessions. Hunger and fullness levels measured immediately before and during 4 h after the meal were identical under the two conditions. In addition, there was no significant difference between the two sessions for RQ and resting energy expenditure, which showed significant reproducibility for measurements obtained immediately before, immediately after, as well as 30 min after, the buffet. This present study demonstrates the high reproducibility of energy and macronutrient intake and oxidation rate values obtained with a buffet-type meal in healthy male subjects and suggests that the use of this test is a reliable method for assessment of macronutrient preferences in the laboratory.

Lipid content and energy density of forage fishes from the northern Gulf of Alaska.

Piscivorous predators can experience multi-fold differences in energy intake rates based solely on the types of fishes consumed. We estimated energy density of 1151 fish from 39 species by proximate analysis of lipid, water, ash-free lean dry matter, and ash contents and evaluated factors contributing to variation in composition. Lipid content was the primary determinant of energy density, ranging from 2 to 61% dry mass and resulting in a five-fold difference in energy density of individuals (2.0-10.8 kJg(-1) wet mass). Energy density varied widely within and between species. Schooling pelagic fishes had relatively high or low values, whereas nearshore demersal fishes were intermediate. Pelagic species maturing at a smaller size had higher and more variable energy density than pelagic or nearshore species maturing larger. High-lipid fishes had less water and more protein than low-lipid fishes. In some forage fishes, size, month, reproductive status, or location contributed significantly to intraspecific variation in energy density. Differences in quality are sufficient to potentially affect diet selection of breeding seabirds, especially when transporting food for their young to the nest site.

Foraging behaviour of Blue Tits Parus caeruleus in a patchy environment under contrasting levels of natural food supply

We analysed the foraging behaviour of free-ranging Blue Tits Parus caeruleus in open holm oak Quercus ilex woodlands of western Spain during winter. Such woodlands are patchy for foraging tits because of the scattered distribution of trees and the patterns of abundance of canopy arthropods within and among trees. Results were compared with those obtained in spring of the same year, when we found that the foraging behaviour and spatial distribution of Blue Tits were largely unaffected by food availability (Pulido and Díaz 1997). Patch (tree) residence time was highly variable both within and among individual birds, and it was uncorrelated with either previous travel time or patch quality. Contrary to a priori expectations, the behaviour of tits did not conform to short-term energy maximizing rules in winter, in spite of a 2.5-fold decrease in food supply from spring to winter and a likely 2-fold increase in bird requirements. Instead, birds tended to fly towards patches that were further away than locally available. Overall, we conclude that energy intake rate was not the fitness-related currency that birds were trying to maximize while foraging.

Foraging ecology of the South Australian glossy black‐cockatoo (Calyptorhynchus lathami halmaturinus)

AbstractThe endangered South Australian glossy black‐cockatoo (Calyptorhynchus lathami halmaturinus Mathews 1912) feeds almost exclusively on the seeds of the drooping sheoak (Allocasuarina verticillata), and shows marked preferences for individual trees. This field study investigated foraging ecology and tree selection through observations of foraging birds and measurements of trees and seed cones. The cockatoos spent the vast majority of their foraging time (94%) handling seed cones, and handling behaviour was highly stereotyped. Handling time per cone was correlated primarily with cone size, while seed intake rate was correlated primarily with seed mass per cone. The cockatoos fed mostly in trees with signs of previous feeding. They tended initially to sample trees with large seeds, and to stay for long feeding bouts in trees with high ratios of seed‐to‐cone mass. As a result of these biases, feeding was concentrated in trees with high seed mass per cone. Preferred trees were also larger, with higher ratios of seed‐to‐cone mass and larger seeds containing more lipid and protein. By feeding from selected trees the cockatoos increased both their seed intake rate and the nutritional quality of the seeds ingested, thereby increasing their energy intake rate by an estimated 28%. They did not discriminate against trees that had re‐grown from basal shoots after fires. Insect larvae were present in some seed cones but the cockatoos did not appear to actively seek them. Males foraged 19% more efficiently than females, resulting in greater daily food intake. The characteristics of individual A. verticillata trees that determined the cockatoos’ feeding rates were also correlated with their distribution on a regional scale. This suggests that the distribution of this endangered cockatoo depends not only on the presence of food trees, but also on their regionally varying feeding profitability.

Maximum daily energy intake: it takes time to lift the metabolic ceiling.

Conventionally, maximum capacities for energy assimilation are presented as daily averages. However, maximum daily energy intake is determined by the maximum metabolizable energy intake rate and the time available for assimilation of food energy. Thrush nightingales (Luscinia luscinia) in migratory disposition were given limited food rations for 3 d to reduce their energy stores. Subsequently, groups of birds were fed ad lib. during fixed time periods varying between 7 and 23 h per day. Metabolizable energy intake rate, averaged over the available feeding time, was 1.9 W and showed no difference between groups on the first day of refueling. Total daily metabolizable energy intake increased linearly with available feeding time, and for the 23-h group, it was well above suggested maximum levels for animals. We conclude that both intake rate and available feeding time must be taken into account when interpreting potential constraints acting on animals' energy budgets. In the 7-h group, energy intake rates increased from 1.9 W on the first day to 3.1 W on the seventh day. This supports the idea that small birds can adaptively increase their energy intake rates on a short timescale.

Seasonal energy utilization in bamboo by the red panda (Ailurus fulgens)

Seasonal energy intake and digestion of bamboo by red pandas were measured in energy balance trials at the field station at an altitude of 3,100 m in Yele Natural Reserve, Mianning County of Sichuan Province, Peoples Republic of China. Two-week trials were conducted at monthly intervals by feeding animals with only natural bamboo browse, Bashania spanostachya, from May to November 1995. Results demonstrated that energy intake, digestible energy, and digestibility differed significantly among seasons and diets. Energy digestibilities were high in summer-autumn, intermediate in spring, and low in winter. These variations correlated positively with the nutrient contents of bamboo. The daily metabolic energy requirement measured varied from 2,603.3 kJ in the spring to 3,139.8 kJ in the summer-autumn to 2,740.8 kJ in the winter. The results also showed that shoots were more easily digested than leaves and exhibited the highest digestibility. Restricted by the capacity of digestive tracts, red pandas poorly utilized the energy provided by bamboo as indicated by the lower energy digestibility (averaging 26.47% for leaves and 44.08% for shoots among the seasons). To fulfill their daily energy requirements, red pandas ingested a large amount of energy from bamboo leaves, which varied from 10,145.8 kJ in the spring, 12,045.1 kJ in the summer-autumn, and 12,276.9 kJ in the winter to maximize the rate of energy intake. Zoo Biol 19:27–33, 2000. © 2000 Wiley-Liss, Inc.

Validation of a self-administered 3-day estimated dietary record for use in the elderly.

was to assess the validity of a 3-day estimated dietary record, especially developed for the longitudinal study on nutrition and health status in an aging population (aged > or = 60 years) in Giessen, Germany. The dietary record comprises 146 food items. Portion sizes of the foods consumed were estimated by means of household measures and corresponding dimensions. The validity of the dietary record was evaluated by means of nitrogen excretion in the 24 h urine in 14 women and 9 men, and by comparing energy intake and basal metabolic rate determined by indirect calorimetry in 238 women and 105 men. Neither in women nor in men were any differences found between calculated nitrogen intake (13.4 +/- 2.2 g and 15.0 +/- 2.0 g, respectively) and nitrogen excretion (12.9 +/- 2.1 g and 15.7 +/- 3.0 g, respectively). The ratio between reported energy intake and basal metabolic rate was 1.62 +/- 0.46 in women and 1.53 +/- 0.46 in men. Regarding energy intake, 7.6% of the females and 16.2% of the males were classified as under-reporters. They show a significantly higher body weight and BMI than the adequate reporters. The results have proved the new 3-day estimated dietary record as suitable to determine energy and nutrient supply of the elderly.

Dynamic changes in prey choice by stickleback during simultaneous encounter with large prey

When a three-spined stickleback Gasterosteus aculeatus encountered prey simultaneously the probability of hanging and the median pursuit time were greater than when prey were encountered sequentially. During simultaneous prey encounter fish did not choose to attack the more profitable prey but instead the nearer prey was handled first except when the difference between the two prey sizes was large. No difference was found in the level of total energetic intake by the fish regardless of prey size pairing. Fish that handled and ate the first prey of a pair in <5 s attacked the second prey with a high probability of success, demonstrating an opportunistic feeding strategy. Importantly however, the fish did not choose to maximize long term energy intake rate by eating both prey, but rather short-term considerations over the course of feeding took precedence. With an empty stomach, the probability of a fish eating (Peat) the first prey handled was high regardless of prey size. As stomach fullness increased, the Peat the first prey handled decreased if it was the larger prey. Hence, the fish were unselective when the stomach was empty but thereafter there was a shift in preference towards the smaller prey. The decision of which prey to attack and eat appeared to be based on short-term energy considerations and the level of stomach fullness. This study demonstrates that feeding on a short-term scale is a crucial factor to take account of when analysing fish feeding during simultaneous prey encounter.