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.     

Common marmosets (<Emphasis Type="Italic">Callithrix jacchus</Emphasis>) do not utilize social information in three simultaneous social foraging tasks

Social foraging is suggested to increase foraging efficiency, as individuals might benefit from public information acquired by monitoring the foraging activities of other group members. We conducted a series experiments with captive common marmosets (Callithrix jacchus) to investigate to what extent marmosets utilize social information about food location when foraging simultaneously with conspecifics. Subjects were confronted with dominant and subordinate demonstrators in three experiments which differed in the amount of information about food location available to the demonstrators. In all three experiments, the focal subjects’ performance in the social condition was not enhanced in comparison to a non-social control condition. Because we could rule out kleptoparasitism and aggressive displacements as explanations, we argue that the subjects’ tendency for scramble competition by avoiding others and dispersing over the foraging area seems to inhibit or mask the acquisition of social information about the location of rewarded patches.     

The role of landmarks in cotton-top tamarin spatial foraging: evidence for geometric and non-geometric features

We report experiments on captive cotton-top tamarins (Saguinus oedipus) designed to explore two components of spatial foraging. First, do tamarins have the capacity to extract geometric information concerning the spatial relationship between a landmark and a piece of food located above or below it? Second, when tamarins use a landmark to find a target location, what non-geometric features of the landmark do they encode? To explore these problems, we created an artificial jungle environment and trained subjects to find food either above or below a target object (i.e., landmark). Once subjects successfully located the food, we transformed various features associated with the landmark, including its color, orientation, and shape; we also manipulated the landmark-food reward distance, the overall shape of the jungle, and the number and position of landmarks. Results showed that the tamarins' success in finding the food reward was not affected by landmark color, orientation, number, or overall shape of the jungle, suggesting that with respect to the particular test conditions, these features are not relevant to the representation of a landmark. Subjects also generalized to novel landmark-food distances, suggesting that they had integrated geometric (i.e., above/below) with non-geometric (i.e., color/shape) features. Performance was negatively affected by changes to the shape of the landmark, indicating that this feature is critical to the representation of a landmark. Accepted after revision: 7 August 2001 Electronic Publication     

Carbon dioxide narcosis modifies the patch leaving decision of foraging parasitoids

Gleaning information is a way for foragers to adjust their behavior in order to maximize their fitness. Information decreases the uncertainty about the environment and could help foragers to accurately estimate environmental characteristics. In a patchy resource, information sampled during previous patch visits is efficient only if it is retained in the memory and retrieved upon arrival in a new patch. In this study, we tested whether the braconid Asobara tabida, a parasitoid of Drosophila larvae, retains information gleaned on patch quality in the memory and adjusts its foraging behavior accordingly. Females were anesthetized with CO₂ after leaving a first patch containing a different number of hosts and were allowed to visit a second patch containing only kairomones. CO₂ is known to erase unconsolidated information from the memory. We show that in the absence of a short CO₂ narcosis, females responded according to their previous experience, whereas anesthetized females did not. The anesthetized females stayed a given time in the second patch irrespective of what they encountered before. CO₂ narcosis had no effect on the residence time of the non-experienced females in a patch containing hosts or only kairomones in comparison with the non-anesthetized females that had a previous foraging experience. We conclude that CO₂ narcosis erases the effect of the previous patch quality, perhaps due to a memory disruption. Direct information processing is likely to be involved in parasitoid decision making through retention of the information on the previous patch quality into a CO₂ sensitive memory.     

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.     

Honeybee methodology, cognition, and theory: recording local directional decisions in interpatch foraging and interpreting their theoretical relevance

Investigations made into the cognitive decision making of honeybees (Apis mellifera) traveling from one flower patch to another flower patch (interpatch foraging) are few. To facilitate such research, we present methods to artificially emulate interpatch foraging and quantify the immediate decision making of honeybees (within 50 cm) choosing to fly an interpatch path. These “Interpatch Methods” are validated, applied, and shown to produce novel information for the field of honeybee spatial cognition. Generally, we demonstrate that a single foraging cohort of honeybees is shown to be capable of making decisions based upon different sets of learned cues, in the exact same context. Specifically, both terminal beacon orientation cues and compass navigation cues can guide the cognitive decision making of interpatch foraging honeybees; our bees chose both cues equally. Finally, the theoretical importance of decision making for interpatch paths is compared with the other foraging paths (outward and homing) with respect to the information available to recruited foragers and scout foragers. We conclude that the ability to analyze interpatch foraging is critical for a more complete understanding of honeybee foraging cognition and that our methods are capable of providing that understanding.     

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.     

Female degus (Octodon degus) monitor their environment while foraging socially

Vigilance or scanning involves interruptions in foraging behavior when individuals lift their heads and conduct visual monitoring of the environment. Theoretical considerations assume that foraging with the "head down", and scanning ("head up") are mutually exclusive activities, such that foraging precludes vigilance. We tested this generalization in a socially foraging, small mammal model, the diurnal Chilean degu (Octodon degus). We studied spontaneous bouts of scanning of captive degus when foraging in pairs of female sibs and non-sibs. We examined the extent to which foraging (head down postures) and scanning (head up postures) were mutually exclusive in subjects exposed to none, partial, and complete lateral visual obstruction of their partners. In addition, we monitored the orientation of their bodies to examine the target of attention while foraging and scanning. Lastly, we examined the temporal occurrence of scanning events to assess the extent of scanning coordination, and whether this coordination is kin-biased. Visual obstruction had a significant influence on degu vigilance. Focal degus increased their quadrupedal and semi-erect scanning when foraging under a partially obstructed view of their partners. Degus oriented their bodies toward partners when foraging and scanning. Despite this, degus did not coordinate scanning bouts; instead, they scanned independently from one another. Relatedness among cage mates did not influence any aspect of degu behavior. Contrary to theoretical expectations, these results indicate that foraging and vigilance are not mutually exclusive, and that kinship per se does not influence scanning behavior and coordination.     

Patch choice as a function of procurement cost and encounter rate.

The effects of patch encounter rate on patch choice and meal patterns were studied in rats foraging in a laboratory environment offering two patch types that were encountered sequentially and randomly. The cost of procuring access to one patch was greater than the other. Patches were either encountered equally often or the high-cost patch was encountered more frequently. As expected, rats exploited the low-cost patch on almost 100% of encounters and exploited the high-cost patch on a percentage of encounters that was inversely proportional to its cost. Meal size was the same at both patches. Surprisingly, when low-cost patches were rare, the rats did not increase their use of high-cost patches. This resulted in spending more time and energy searching for patches and a higher average cost per meal. The rats responded to this increased cost by reducing the frequency and increasing the size of meals at both patches and thereby limited total daily foraging cost and conserved total intake.     

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.     

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.     

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.     

Experimental setting affects the performance of guppies in a numerical discrimination task

A recent study found that guppies (Poecilia reticulata) can be trained to discriminate 4 versus 5 objects, a numerical discrimination typically achieved only by some mammals and birds. In that study, guppies were required to discriminate between two patches of small objects on the bottom of the tank that they could remove to find a food reward. It is not clear whether this species possesses exceptional numerical accuracy compared with the other ectothermic vertebrates or whether its remarkable performance was due to a specific predisposition to discriminate between differences in the quality of patches while foraging. To disentangle these possibilities, we trained guppies to the same numerical discriminations with a more conventional two-choice discrimination task. Stimuli were sets of dots presented on a computer screen, and the subjects received a food reward upon approaching the set with the larger numerosity. Though the cognitive problem was identical in the two experiments, the change in the experimental setting led to a much poorer performance as most fish failed even the 2 versus 3 discrimination. In four additional experiments, we varied the duration of the decision time, the type of stimuli, the length of training, and whether correction was allowed in order to identify the factors responsible for the difference. None of these parameters succeeded in increasing the performance to the level of the previous study, although the group trained with three-dimensional stimuli learned the easiest numerical task. We suggest that the different results with the two experimental settings might be due to constraints on learning and that guppies might be prepared to accurately estimate patch quality during foraging but not to learn an abstract stimulus–reward association.     

Contextual interference effects in sequence learning for young and older adults

Practice of different tasks in a random order induces better retention than practicing them in a blocked order, a phenomenon known as the contextual interference (CI) effect. Our purpose was to investigate whether the CI effect exists in sequence learning, such that practicing different sequences in a random order will result in better learning of sequences than practicing them in blocks, and whether this effect is affected by aging. Subjects practiced a serial reaction time task where a set of three 4-element sequences were arranged in blocks or in a random order on 2 successive days. Subjects were divided into 4 groups based on a 2-GROUP (young or old) by 2-ORDER (random or blocked practice) between-subject design. Three days after practice (Day 5), subjects were tested with practiced and novel sequences to evaluate sequence-specific learning. The results replicate the CI effect in sequence learning in both young and older adults. Older adults retained sequences better when trained in a random condition than in a blocked condition, although the random condition incurs greater task switching costs in older adults during practice. Our study underscores the distinction between age-related effects on learning vs. performance, and offers practical implications for enhancing skill learning in older adults.     

Reading Nonwords Aloud: Evidence for dynamic control in skilled readers

In two experiments, we examined whether the dynamics of the reading system are adjusted on a trial-by-trial basis, despite the use of stimuli that all require the same spelling?Csound translation routine. Subjects read aloud easy and more difficult nonwords in a predictable alternating sequence (AABB). Dynamic control was inferred via the observation of switch costs in response times and/or accuracy (A to B and B to A) for both types of items. Consistent with online control, switch costs were observed for both kinds of items. Various ways in which the reading system could adjust in response to such stimuli are considered.     

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.     

Discrimination learning in a foraging situation

Rats were allowed to forage in a simulated natural environment made up of eight food sources (patches) each containing a fixed number of pellets. Two of the eight contained an extra supply of peanuts. The peanut patches were signaled by an olfactory/visual cue located at the bottom of the ladder leading to the patch. In successive phases the number of sessions per day, height of the patches, and availability of peanuts were manipulated. Subjects showed evidence of discrimination learning under these conditions, although the degree of discriminatory behavior varied as a function of environmental manipulations. Assessment of behavior within foraging sessions showed that subjects systematically changed their patterns of utilization of patches across time. Sampling or exploration, as well as food reinforcement, seem implicated in these results.     

“To eat or not to be eaten?” Collective risk-monitoring in groups

The importance of risk-monitoring has been increasing in many key aspects of our modern lives. This paper examines how individuals monitor such risks collectively by extending a behavioral ecological model of animal foraging to human groups. Just as animals must forage for food under predatory risk, humans must divide valuable material and psychological resources between foraging activity and risk-monitoring activity. We predicted that game-theoretic aspects of the group situation complicate such a trade-off decision in resource allocation, eventually yielding a mixed equilibrium in a group. When the equilibrium is reached, only a subset of members engage in the risk-monitoring activity while others free-ride, concentrating mainly on their own foraging activity. Laboratory groups engaging in foraging under moderate risk provided a support to this prediction. When the risk-level was set higher, however, “herding behavior” (conforming to the dominant behavior) interfered with the emergence of equilibrium. Implications for risk management are discussed.     

Learning a keying sequence you never executed: Evidence for independent associative and motor chunk learning

A substantial amount of research has addressed how people learn and control movement sequences. Recent results suggested that practice with discrete key pressing sequences results in two types of sequence learning: associative learning and motor chunk development (Verwey & Abrahamse, 2012). In the present study, we addressed whether in keying sequences of limited length associative learning develops also when the use of the chunking mode is prevented by introducing during practice random deviants. In line with the notion of two different learning mechanisms, the present results indicate that associative sequence learning develops when motor chunks cannot be developed during practice. This confirms the notion that motor chunks do not rely on these associations. In addition, experience with a particular execution mode during the practice phase seems to benefit subsequent use of that mode with unfamiliar and random sequences. Also, participants with substantial video-gaming experience were faster in executing discrete keying sequences in the chunking mode. These last two results may point to the development of a general ability to produce movement sequences in the chunking mode.     

Sampling and decision rules used by honey bees in a foraging arena

Animals must continuously choose among various available options to exploit the most profitable resource. They also need to keep themselves updated about the values of all available options, since their relative values can change quickly due to depletion or exploitation by competitors. While the sampling and decision rules by which foragers profitably exploit a flower patch have attracted a great deal of attention in theory and experiments with bumble bees, similar rules for honey bee foragers, which face similar foraging challenges, are not as well studied. By presenting foragers of the honey bee Apis cerana with choice tests in a foraging arena and recording their behavior, we investigate possible sampling and decision rules that the foragers use to choose one option over another and to track other options. We show that a large part of the sampling and decision-making process of a foraging honey bee can be explained by decomposing the choice behavior into dichotomous decision points and incorporating the cost of sampling. The results suggest that a honey bee forager, by using a few simple rules as part of a Bayesian inference process, is able to effectively deal with the complex task of successfully exploiting foraging patches that consist of dynamic and multiple options.