Drinking in a patchy environment: the effect of the price of water.

Rats in a laboratory foraging paradigm searched for sequential opportunities to drink in two water patches that differed in the bar-press price of each "sip" (20 licks) of water within a bout of drinking (Experiment 1) or the price and size (10, 20, or 40 licks) of each sip (Experiment 2). Total daily water intake was not affected by these variables. The rats responded faster at the patch where water was more costly. However, they accepted fewer opportunities to drink, and thus had fewer drinking bouts, and drinking bouts were smaller at the more costly patch than at the other patch. This resulted in the rats consuming a smaller proportion of their daily water from the more costly patch. The size of the differences in bout frequency and size between the patches appears to be based on the relative cost of water at the patches. The profitability of each patch was calculated in terms of the return (in milliliters) on either effort (bar presses) or time spent there. Although both measures were correlated with the relative total intake, bout size, and acceptance of opportunities at each patch, the time-based profitability was the better predictor of these intake measures. The rats did not minimize bar-press output; however, their choice between the patches and their bout sizes within patches varied in a way that reduced costs compared to what would have been expended drinking randomly. These data accord well with similar findings for choices among patches of food, suggesting that foraging for water and food occurs on the basis of comparable benefit-cost functions: In each case, the amount consumed is related to the time spent consuming.     

Procurement time as a determinant of meal frequency and meal duration.

Foraging involves the expenditure of both time and effort in the acquisition of food; animals typically modify their meal patterns so as to reduce these expenditures or costs. The contribution of time, as compared with effort, to the overall cost perceived by an animal is not known. We investigated the effect of foraging time as a cost independent of effort by measuring the meal patterns of rats living in a laboratory foraging simulation in which they earned all their daily intake. They pressed a bar once to initiate an interval (procurement interval) leading to the presentation of a large cup of food from which they could eat a meal of any size. As the length of the interval increased from 1 s to 46 hr, meal frequency decreased regularly. Meal size increased in a compensatory fashion, and total daily intake was conserved through an interval of 23 hr. The changes in meal frequency occurred because of changes in the rat's latency to bar press after each meal. The functions relating meal frequency and size to the procurement interval were of the same shape as those seen when cost is the completion of a bar-press requirement, which entails the expenditure of both effort and time. When the bar-press requirement was increased to 10, meal frequency was reduced, but time and effort did not appear to simply add together in the rat's perception of cost. These data reveal that time is preceived to be a cost by rats foraging in this laboratory environment. These results suggest that the time parameters of foraging are different from those of consumption.     

How to measure patch encounter rate: decision-making mechanisms in the parasitic wasp <Emphasis Type="Italic">Asobara tabida</Emphasis>

Parasitic wasps are faced with the decision of where and for how long to search for hosts. Their leaving decisions depend on the rate at which new host-containing patches are encountered: parasitoids increase foraging efficiency by leaving earlier when patch encounter rates become higher. The mechanisms by which these often tiny insects can assess patch encounter rates have not been thoroughly investigated so far. The aim of the present study, where females of the braconid wasp Asobara tabida encountered patches after varying time intervals, was to measure the shape of the travel–time response curve and to analyse how information on inter-patch distances is translated into foraging behaviour. I examined several proxies for travel-time duration, like those of physiological nature as egg content, cues of senescence, amount of energy spent, or muscle fatigue, as well as true cognitive mechanisms, like measurement of distance or interval timing. Constraints in the wasp’s ability to detect patch borders accurately after travelling, e.g. habituation to the patch odour or receptor blocking, are also discussed. From the data presented, most of the above-mentioned mechanisms and constraints can be rejected to work for A. tabida. The effects of inter-patch travel time are strongest when they are short, and even though it cannot be excluded that time measures are processed using an internal clock, I suggest that a Bayesian-like mechanism of timing, the biological basis of which might involve the build-up of neurosecretory material, is the most likely candidate influencing leaving decisions in A. tabida.     

The relationship between feeding rate and patch choice.

Rats in a laboratory foraging simulation searched for sequential opportunities to feed in two patches that differed in the rate at which food pellets were delivered (controlled by fixed-interval schedules) and in the size of the pellets. The profitability of feeding in each patch was calculated in terms of time (grams per minute) and in terms of effort (grams per bar press). These values were the result of the imposed fixed interval, the size of the pellets, and the rate at which the rats pressed the bar in each condition. The rats ate more food and larger meals, but not more frequent meals, at the patch offering the higher rate of food consumption, calculated as grams per minute. The relative intake at any patch was a function of the relative rate of intake during meals at that patch compared to the other patch. Rats respond to explicit manipulations of feeding time in the same manner as they respond to manipulations of feeding effort.     

Time horizons in rats foraging for food in temporally separated patches

An important tenet of optimal foraging theory is that foragers compare prey densities in alternative patches to determine an optimal distribution of foraging behavior over time. A critical question is over what time period (time horizon) this integration of information and behavior occurs. Recent research has indicated that rats do not compare food density in a depleting patch with that in a rich patch delayed by an hour or more (Timberlake, 1984). In the present research we attempted to specify over what time period a future rich patch would affect current foraging. The effect of future food was measured by early entry into the rich patch (anticipation) and by a decrease in food obtained in the depleting patch (suppression). The rats showed anticipation of a rich patch up to an hour distant, but suppressed current feeding only if the rich patch was 16 min distant or less. The suppression effect appeared mediated by competition for expression between anticipatory entries into the rich patch and continued foraging in the depleting patch. These results suggest that optimal foraging is based on a variety of specific mechanisms rather than a general optimizing algorithm with a single time horizon.     

Effects of search cost on foraging and feeding: a three-component chain analysis

An experiment determined whether pigeons minimize number of key pecks per food delivery and maintain their baseline intake of food while key pecking on a three-component chain schedule. Pigeons at either 80% or 100% body weight obtained all their food during baseline and contingency sessions. During baseline sessions, pecks on the left and center keys had no consequences; each peck on the right key activated the feeder. During contingency sessions, pigeons key pecked on a three-component chain schedule simulating components of a foraging chain. In the search component either 3, 9 or 15 key pecks (varied parametrically across blocks of sessions) on the left key produced a stimulus on the middle key, indicating an encounter with either the low-cost prey (3 key pecks) or an equally probable high-cost prey (21 key pecks). In the procurement component the pigeon pecked either: (a) the left key once, thus returning to the search component, or (b) the middle key either 3 or 21 times, which activated the right response key. In the handling component one peck on the right key operated the feeder. The pigeons always procured the low-cost prey and minimized the number of key pecks per hopper by procuring the high-cost prey when the search-cost ratio was high (15 key pecks) but not when it was low (3 key pecks). All pigeons maintained their baselines of eating during contingency sessions by key pecking more frequently and eating more efficiently. The 80% body-weight birds produced higher overall rates of key pecking and eating. These results have implications for ecological theories of optimal foraging and for psychological theories of learned performance.     

Meal patterns of cats encountering variable food procurement cost.

The meal patterns of 2 cats in a laboratory habitat with variable foraging costs were examined in a foraging paradigm in which subjects could initiate meals at any time by completing a predetermined number of bar presses (the procurement price) and then could eat any amount. From meal to meal, the procurement price either was fixed or varied among a geometric series of five prices. As the fixed price or the mean of the variable prices increased, meal frequency decreased and meal size increased; daily intake was unaffected. Within variable-price schedules, meal size was not related to the just-paid procurement price. These results suggest that cats respond to the global rather than to the local cost structure of their habitat. They appear to respond to an average of the prices encountered, initiating meals of a frequency and size appropriate to that average. This was true even when the average price was high, meals were infrequent, and thus price encounters were widely separated in time. Therefore, the time window over which the consequences of behavior can affect behavior is longer than often conceived, at least in economies in which the animal controls its intake and the frequency, size, and distribution of its meals.     

Meal patterns following changes in procurement cost for rats with fornix transection

The purpose of the present study was to explore the effects of a varied procurement cost on the foraging behavior of rats with fornix transection. An operant analog of foraging requirements was used to examine the feeding patterns of the animals under free feeding, low procurement cost (FR5), and high procurement cost (FR80) situations, in an environment with minimal sensory distraction. It was found that animals with fornix transection did not differ from control rats in general consumption. Both groups were also able to adapt their feeding behavior to the varied procurement cost. As the procurement cost increased, the number of meals consumed decreased while the meal duration increased. The meal patterns themselves were different for the fornix transected animals and the control group. Animals with fornix transections ate more meals over the course of a day than did control animals; their meals were of a longer duration, and their intermeal intervals were shorter than those of control animals. During the course of a meal, the rats with fornix transections took a larger number of breaks, during which they drank, explored, or engaged in activities other than eating. These differences in the feeding patterns were seen across all procurement cost levels. The data support the possibility of hippocampal involvement in behavioral organization or sequencing.     

Residence time in concurrent foraging with fixed times to prey arrival

Five pigeons were trained in a concurrent foraging procedure in which reinforcers were occasionally available after fixed times in two discriminated patches. In Part 1 of the experiment, the fixed times summed to 10 s, and were individually varied between 1 and 9 s over five conditions, with the probability of a reinforcer being delivered at the fixed times always .5. In Part 2, both fixed times were 5 s, and the probabilities of food delivery were varied over conditions, always summing to 1.0. In Parts 3 and 4, one fixed time was kept constant (Part 3, 3 s; Part 4, 7 s) while the other fixed time was varied from 1 s to 15 s. Median residence times in both patches increased with increases in the food-arrival times in either patch, but increased considerably more strongly in the patch in which the arrival time was increased. However, when arrival times were very different in the two patches, residence time in the longer arrival-time patch often decreased. Patch residence also increased with increasing probability of reinforcement, but again tended to fall when one probability was much larger than the other. A detailed analysis of residence times showed that these comprised two distributions, one around a shorter mode that remained constant with changes in arrival times, and one around a longer mode that monotonically increased with increasing arrival time. The frequency of shorter residence times appeared to be controlled by the probability of, and arrival time of, reinforcers in the alternative patch. The frequency of longer residence times was controlled directly by the arrival time of reinforcers in a patch, but not by the probability of reinforcers in a patch. The environmental variables that control both staying in a patch and exiting from a patch need to be understood in the study both of timing processes and of foraging.     

An experimental analysis of optimal foraging behaviour in patchy environments

A series of experiments was designed to explore the cognitive mechanisms involved in optimal foraging models by using the behavioural controls of operant methodology. Rats were trained to press one of two levers to obtain reinforcement on a progressive variable-interval schedule, which modelled food patch depletion; the schedule was reset by pressing the other lever. Thus both duration (residence time in a patch) and rate-related (interval before and after the final reward) measures were obtained. Experiment 1, which manipulated environmental stability and quality, and Experiment 2, which varied travel time between patches, found results that supported the marginal value theorem (Charnov, 1976) and suggested that rats adjust capture rate to the environment average by monitoring the length of the interval between rewards. Experiment 3 modelled the clumping of food items and found that capture rate was now adjusted by adoption of a fixed giving-up time. Finally, Experiments 4a and 4b ruled out a time expectancy hypothesis by manipulating the number of food clumps and the series of inter-reinforcement intervals. Overall the experiments demonstrate the value of modelling foraging strategies in operant apparatus, and suggest that rats adopt rate predictive strategies when deciding to switch patches.     

Foraging in a simulated natural environment: There's a rat loose in the lab

Rats were required to earn their food in a large room having nine boxes placed in it, each of which contained food buried in sand. In different phases of the experiment the amount of time allowed for foraging, the amount of food available in each food patch, and the location of the different available amounts were varied. The rats exhaustively sampled all patches each session but seemed to have fairly strong preferences for certain locations over others. If position preferences were for patches containing small amounts of food, the sensitivity to amount available was increased so that when location was compensated for, a pattern of optimal foraging was evident. The importance of environmental constraints in producing optimal behavior and the relation of the observed behavior to laboratory findings are discussed.     

Predictors of children's prosocial lie-telling: Motivation, socialization variables, and moral understanding

Children tell prosocial lies for self- and other-oriented reasons. However, it is unclear how motivational and socialization factors affect their lying. Furthermore, it is unclear whether children’s moral understanding and evaluations of prosocial lie scenarios (including perceptions of vignette characters’ feelings) predict their actual prosocial behaviors. These were explored in two studies. In Study 1, 72 children (36 second graders and 36 fourth graders) participated in a disappointing gift paradigm in either a high-cost condition (lost a good gift for a disappointing one) or a low-cost condition (received a disappointing gift). More children lied in the low-cost condition (94%) than in the high-cost condition (72%), with no age difference. In Study 2, 117 children (42 preschoolers, 41 early elementary school age, and 34 late elementary school age) participated in either a high- or low-cost disappointing gift paradigm and responded to prosocial vignette scenarios. Parents reported on their parenting practices and family emotional expressivity. Again, more children lied in the low-cost condition (68%) than in the high-cost condition (40%); however, there was an age effect among children in the high-cost condition. Preschoolers were less likely than older children to lie when there was a high personal cost. In addition, compared with truth-tellers, prosocial liars had parents who were more authoritative but expressed less positive emotion within the family. Finally, there was an interaction between children’s prosocial lie-telling behavior and their evaluations of the protagonist’s and recipient’s feelings. Findings contribute to understanding the trajectory of children’s prosocial lie-telling, their reasons for telling such lies, and their knowledge about interpersonal communication.     

GROUP FORAGING SENSITIVITY TO PREDICTABLE AND UNPREDICTABLE CHANGES IN FOOD DISTRIBUTION: PAST EXPERIENCE OR PRESENT CIRCUMSTANCES?

The ideal free distribution theory (Fretwell & Lucas, 1970) predicts that the ratio of foragers at two patches will equal the ratio of food resources obtained at the two patches. The theory assumes that foragers have "perfect knowledge" of patch profitability and that patch choice maximizes fitness. How foragers assess patch profitability has been debated extensively. One assessment strategy may be the use of past experience with a patch. Under stable environmental conditions, this strategy enhances fitness. However, in a highly unpredictable environment, past experience may provide inaccurate information about current conditions. Thus, in a nonstable environment, a strategy that allows rapid adjustment to present circumstances may be more beneficial. Evidence for this type of strategy has been found in individual choice. In the present experiments, a flock of pigeons foraged at two patches for food items and demonstrated results similar to those found in individual choice. Experiment 1 utilized predictable and unpredictable sequences of resource ratios presented across days or within a single session. Current foraging decisions depended on past experience, but that dependence diminished when the current foraging environment became more unpredictable. Experiment 2 repeated Experiment I with a different flock of pigeons under more controlled circumstances in an indoor coop and produced similar results.     

Blind to Bias? Young Children Do Not Anticipate that Sunk Costs Lead to Irrational Choices

Young children anticipate that others act rationally in light of their beliefs and desires, and environmental constraints. However, little is known about whether children anticipate others’ irrational choices. We investigated young children's ability to predict that sunk costs can lead to irrational choices. Across four experiments, 5- to 6-year-olds (total N = 185) and adults (total N = 117) judged which of two identical objects an agent would keep, one obtained at a high cost or one obtained at a low cost. In Experiment 1, adults predicted that the agent would choose the high-cost object over the low-cost one, whereas children responded at chance. Experiment 2 replicated these findings in children, but also included another condition which showed they were sensitive to future costs. They predicted that an agent would be more likely to seek out a low-cost item than a high-cost item. Experiments 3 and 4 then found that children do not anticipate the sunk cost bias in first person scenarios, or in interpersonal sunk cost scenarios, where costs are sunk by others. Taken together, our findings suggest that young children may struggle to understand and predict irrational behavior. The findings also reveal an asymmetry between how they consider sunk costs and future costs in understanding actions. We propose that this asymmetry might arise because children do not consider sunk costs as wasted.     

Testing a stochastic foraging model in an operant simulation: Agreement with qualitative but not quantitative predictions

An operant simulation of foraging through baited and empty patches was studied with 4 pigeons. On a three-key panel, side keys were designated as patches, and successive opportunities to complete 16 fixed-ratio 10 schedules on side keys were defined as encounters with feeders. In a random half of the patches in any session, some of the fixed-ratio 10 schedules yielded reinforcement (baited feeders) and the other schedules yielded nonreinforcement (empty feeders). In the other half of the patches, all feeders were empty. Pigeons could travel between patches at any time by completing a fixed-ratio schedule on the center key. An optimal foraging model was tested in Experiments 1 and 2 by varying center-key travel time and number of baited feeders in baited patches. The ordinal predictions that number of feeders visited in empty patches would increase with travel time and decrease as number of baited feeders increased were supported, but pigeons visited far more feeders in empty patches than the optimal number predicted by the model to maximize energy/time. In Experiment 3, evidence was found to suggest that the number of empty feeders encountered before the first baited feeder in baited patches is an important factor controlling leaving empty patches.     

Comparing Locomotion With Lever-press Travel In An Operant Simulation Of Foraging

An operant model of foraging was studied. Rats searched for food by pressing on the left lever, the patch, which provided one, two, or eight reinforcers before extinction (i.e., zero reinforcers). Obtaining each reinforcer lowered the probability of receiving another reinforcer, simulating patch depletion. Rats traveled to another patch by pressing the right lever, which restored reinforcer availability to the left lever. Travel requirement changed by varying the probability of reset for presses on the right lever; in one condition, additional locomotion was required. That is, rats ran 260 cm from the left to the right lever, made one response on the right lever, and ran back to a fresh patch on the left lever. Another condition added three hurdles to the 260-cm path. The lever-pressing and simple locomotion conditions generated equivalent travel times. Adding the hurdles produced longer times in patches than did the lever-pressing and simple locomotion requirements. The results contradict some models of optimal foraging but are in keeping with McNair's (1982) optimal giving-up time model and add to the growing body of evidence that different environments may produce different foraging strategies.     

Dynamic averaging and foraging decisions in horses (Equus callabus)

The variability of most environments taxes foraging decisions by increasing the uncertainty of the information available. One solution to the problem is to use dynamic averaging, as do some granivores and carnivores. Arguably, the same strategy could be useful for grazing herbivores, even though their food renews and is more homogeneously distributed. Horses (Equus callabus) were given choices between variable patches after short or long delays. When patch information was current, horses returned to the patch that was recently best, whereas those without current information matched choices to the long-term average values of the patches. These results demonstrate that a grazing species uses dynamic averaging and indicate that, like granivores and carnivores, they can use temporal weighting to optimize foraging decisions.     

Changes in feeding and foraging patterns as an antipredator defensive strategy: a laboratory simulation using aversive stimulation in a closed economy.

The effects of the risk of electric shock on the meal patterns of rats living in an operant chamber were investigated. Rats could obtain food by working on a response lever that provided reinforcement according to chained fixed-ratio continuous reinforcement schedules that allowed the animals control over meal size. Using a two-compartment operant chamber with a safe nesting area and manipulanda area with a grid floor, shock could be correlated with responding on the schedule. Shocks (less than or equal to 1.25 per hour) were scheduled to occur randomly throughout the day, independent of the rat's behavior. Shock caused a reorganization of meal patterns such that the animals took less frequent but larger meals. This pattern reduced the time the animals spent at risk without compromising caloric balance. Similar changes in feeding pattern were obtained in both hooded and albino rats. Exposure to shock in a separate chamber did not produce these behavioral modifications. The magnitude of shock-induced alterations of meal patterns was greater with chained fixed-ratio 90 continuous reinforcement than with chained fixed-ratio 10 continuous reinforcement. Additionally, the rats seemed to be able to reduce food intake but increase caloric efficiency, such that the reduced food intake did not have deleterious effects on maintenance of body weight. These behavioral modifications reduced the number of shocks received from that which would have been expected if meal pattern changes had not occurred. We suggest that this technique may provide a useful laboratory simulation of the impact that the risk of predation has on foraging behavior.     

Altruism between romantic partners: Biological offspring as a genetic bridge between altruist and recipient

When the cost of altruism is low, individuals are more likely to help non-kin (i.e., friends and romantic partners) than kin. This trend is thought to reflect the fact that people tend to be emotionally closer with friends and romantic partners than kin. However, as the cost of altruism increases, altruistic preference shifts to kin. The present study highlights this phenomenon by examining altruism between siblings, romantic partners, romantic partners who have biological children together, and romantic partners who have adopted children together. Participants (n = 203) completed a questionnaire about altruism in low-, medium-, and high-cost situations. Participants gave more low-cost help to their romantic partners (regardless of whether they had a child together) than their siblings. More medium-cost help was given to romantic partners who had a child (biological and adopted) than siblings and romantic partners without children. In the high-cost condition, the estimated altruistic tendencies were stronger toward siblings and romantic partners who have a biological child than toward romantic partners with no children and partners with adopted children. Participants also believed they were more altruistic than their siblings and romantic partners.     

The currency of procurement cost

As the number of instrtumental responses required to procure access to food is increased, animals decrease the frequency of initiating meals and increase meal size, conserving total intake while limiting the increase in the overall cost of feeding. In two studies, one using wheel turns and one using bar presses as the instrumental response, we asked whether freely feeding laboratory rats measure cost according to the energy or the time they expend. In each study we varied both the price (i.e., number of wheel turns or bar presses) and the force required to make a response (i.e., torque on the wheel or weight of the bar). Price affected both procurement time (from the first to the last procurement response) and procurement work, whereas torque and bar weight affected work without altering time in most cases. Meal patterns were altered by all manipulations of price, but changes in torque and bar weight had little effect on meal patterns, except in the conditions in which they altered procurement time. These results suggest that time is a critical currency of procurement cost in rats.