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Todd, I. A., & Kacelnik, A. (1993). Psychological mechanisms and the Marginal Value Theorem: dynamics of scalar memory for travel time. Anim. Behav., 46(4), 765–775.
Abstract: Abstract. The relation between memory for travel time and foraging decisions was studied experimentally. The temporal properties of two environments with patchily distributed food were simulated in the laboratory using pigeons, Columba livia, as subjects. The two environments differed in mean travel time, while the coefficient of variation of travel time and the decelerated function relating cumulative food gain to time in the patch were held constant within and between environments. Each environment contained a uniform mixture of five travel times experienced in a random order. Two of the five travel times were common in both environments. Effects of travel time were studied by comparing prey collected per patch visit (PPV) after various travel times within each environment, and by comparing patch exploitation after equal travel times between environments. Within the environment with long mean travel time (LMT) PPV was positively correlated with the last and the penultimate travel times but not with travel times before that. The increase in PPV per second of last travel time was six times greater than the increase per second of penultimate travel time, implying very steep memory discounting. In the environment with short mean travel time (SMT), there was no correlation between PPV and previous travel times. However, comparisons between environments of visits following travel times common to both environments (thus removing the effect of the last travel time) showed that substantially more prey were taken after equal travel times in the LMT than in the SMT environment. This difference cannot be accounted for by the within-environment effect of penultimate travel time, implying that there is a different, less steeply devalued, effect of the mixture of travel times. A model of information processing based on combining Scalar Expectancy Theory with the predictions of rate maximization under the Marginal Value Theorem is presented. The model can approximate the results obtained in this and previous experiments and provides a framework for further analysis of memory mechanisms of foraging behaviour.
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Bateson, M., & Kacelnik, A. (1997). Starlings' preferences for predictable and unpredictable delays to food. Anim. Behav., 53(6), 1129–1142.
Abstract: Risk-sensitive foraging theory is based on the premise that unpredictable runs of good or bad luck can cause a variable food source to differ in fitness value from a fixed food source yielding the same average rate of gain but no unpredictability. Thus, risk-sensitive predictions are dependent on the food intake from variable sources being not only variable but also unpredictable or `risky' in outcome. This study tested whether unpredictability is a component of the value that foraging starlings,Sturnus vulgarisattribute to food sources that are variable in the delay to obtain food. Two groups of birds chose between a fixed and a variable delay option; the variable option was unpredictable in the risky group and predictable in the risk-free group in the overall rate of intake it yielded. In both groups the fixed option was adjusted by titration to quantify the magnitude of preference for predictable and unpredictable variance. On negative energy budgets both groups were significantly risk-prone, with the risky group being significantly more risk-prone than the risk-free group. Switching the birds to positive budgets by doubling the size of each food reward had no significant effect on preference, and similar trends to those found with negative budgets were observed. These results are not readily explained by risk-sensitive foraging theory, but may be explained by the algorithm used by the birds to attribute value to average expected rewards.
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Schuck-Paim, C., & Kacelnik, A. (2002). Rationality in risk-sensitive foraging choices by starlings. Anim. Behav., 64(6), 869–879.
Abstract: Normative models of choice usually predict preferences between alternatives by computing their value according to some criterion, then identifying the alternative with greatest value. An important consequence of this procedure is captured in the economic concept of rationality, defined through a number of principles that are necessary for the existence of an ordinal scale of value upon which organisms base their choices. Violations of these principles, such as some recently reported breaches of transitivity and regularity in birds and honeybees, have strong implications for the understanding of decision mechanisms in humans and nonhumans alike. We investigated rationality in risky choice using European starlings, Sturnus vulgaris. Birds had to choose between two or three food sources, each associated with a different variance in delay to reward. In three experiments, starlings were strongly risk prone, showing regular and consistent preferences in binary and trinary choices. Preferences also satisfied weak and strong stochastic transitivity. Our results extend the generality of previous research in risk-sensitive foraging to situations where more than two alternatives are present and suggest that violations of rationality in risk-sensitive choices may be expressed only under restricted sets of conditions. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.
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Caraco, T., Kacelnik, A., Mesnick, N., & Smulewitz, M. (1992). Short-term rate maximization when rewards and delays covary. Anim. Behav., 44(Part 3), 441–447.
Abstract: In nature foragers must exploit resources that vary randomly in both the energy acquired per item (reward) and the time required to pursue, capture and process an item (delay). Furthermore, rewards and delays associated with particular resources may often covary significantly. An analytical model asks how variance-covariance levels for rewards and delays could influence choice of resources when lack of information or cognitive limitation implies that a consumer attempts to maximize its short-term rate of energy gain. Both greater expected reward and reduced expected delay clearly should enhance preference for a resource. The model predicts that increased delay variance and reduced reward-delay covariance should increase a forager's preference for a resource. A forager should be risk-averse towards reward variance when the reward-delay covariance is positive, but should become risk-prone towards reward variance when the reward-delay covariance is negative.
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Kacelnik, A. (1979). The foraging efficiency of great tits (Parus major L.) in relation to light intensity. Anim. Behav., 27(Part 1), 237–241.
Abstract: I report an experiment aimed at testing whether foraging efficiency of great tits is limited by light intensity at the time of the dawn chorus. Captive great tits hunting for prey under different luminance conditions were less successful in finding prey when foraging, hunted for a lower proportion of their time, and handled individual prey items for longer when luminance was under approximately 7 cd/m2. This luminance is not reached in the field until after the time of the dawn chorus, suggesting that in the early morning foraging is limited by light intensity. I suggest that a satisfactory functional explanation of the dawn chorus must take into account the comparatively low foraging opportunity early in the morning, as well as the factors affecting the opportunity for singing and other territorial activities.
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Roberts, J., Kacelnik, A., & Hunter, M. L. (1979). A model of sound interference in relation to acoustic communication. Anim. Behav., 27(Part 4), 1271–1273.
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Kacelnik, A. (1987). Information primacy or preference for familiar foraging techniques? A critique of Inglis & Ferguson. Anim. Behav., 35(3), 925–926.
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Cuthill, I., & Kacelnik, A. (1990). Central place foraging: a reappraisal of the `loading effect'. Anim. Behav., 40(6), 1087–1101.
Abstract: Animals that provision a central place usually bring back larger loads when foraging far from home. This positive correlation between average load size and distance is typically explained as rate-maximizing behaviour in the face of a trade-off between travel costs and a decelerating rate of prey gain in food patches (the `loading effect'). By using feeders to provide wild parent starlings, Sturnus vulgaris, with constant rates of prey loading, a positive load-distance correlation was shown to exist in the absence of a loading effect (experiment I). However, in a laboratory simulation where no load was transported (experiment II). the average number of prey eaten in patch visits by self-feeding starlings was invariant with travel distance, so the explanation of the load-distance correlation in experiment I must lie in featues peculiar to central place foraging. Bottlenecks in ingestion by chicks and interruption by visual detection of nest disturbance (experiment III) were rejected as causes of the correlation. Risks of dropping prey in flight appeared low, but the risk of kleptoparasitism received weak support. The travel-load size correlation may be an adaptive response to load transport costs, as return travel times increased with the load size being carried (experiment IV).
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Reboreda, J. C., & Kacelnik, A. (1990). On cooperation, tit-for-tat and mirros. Anim. Behav., 40(6), 1188–1189.
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Kacelnik, A., & Todd, I. A. (1992). Psychological mechanisms and the Marginal Value Theorem: effect of variability in travel time on patch exploitation. Anim. Behav., 43(2), 313–322.
Abstract: The Marginal Value Theorem (MVT) describes the behaviour that maximizes the ratio of expected gain over expected foraging time in a patchy environment. When travel time is variable, the MVT rationale and its predictions are sensitive only to the mean travel time and not to the spread or skew of the distribution. Two mechanistic arguments contradict these predictions. First, tests of the MVT have previously shown that there is a disproportionate influence of the last travel time, and second, psychological models of information processing suggest that memory for time intervals is strongly dependent on the scatter of the distribution experienced. These mechanistic concepts, combined with Jensen's inequality, suggest that patch exploitation should decrease as the scatter of the travel distribution increases. In a Skinner box experiment with pigeons, Columba livia, the problem was examined by simulating three environments with identical patches and the same mean travel time, but different travel time variability. Patch exploitation decreased with increasing variance in travel time. The results are used to argue in favour of the inclusion of realistic psychological properties as constraints in functional models of behaviour. Although both the MVT and the mechanistic models account for some features of the results, none of them can explain all the findings.
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