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Pompilio, L., & Kacelnik, A. (2005). State-dependent learning and suboptimal choice: when starlings prefer long over short delays to food. Anim. Behav., 70(3), 571–578.
Abstract: Recent studies have used labels such as `work ethics', `sunk costs' and `state-dependent preferences' for apparent anomalies in animals' choices. They suggest that preference between options relates to the options' history, rather than depending exclusively on the expected payoffs. For instance, European starlings, Sturnus vulgaris, trained to obtain identical food rewards from two sources while in two levels of hunger preferred the food source previously associated with higher hunger, regardless of the birds' state at the time of testing. We extended this experimentally and theoretically by studying starlings choosing between sources that differed not only in history but also in the objective properties (delay until reward) of the payoffs they delivered. Two options (PF and H) were initially presented in single-option sessions when subjects were, respectively, prefed or hungry. While option PF offered a delay until reward of 10 s in all treatments, option H delivered delays of 10, 12.5, 15 and 17.5 s in four treatments. When training was completed, we tested preference between the options. When delays in both options were equal (10 s), the birds strongly preferred H. When delay in H was 17.5 s, the birds were indifferent, with intermediate results for intermediate treatments. Preference was not mediated by disrupted knowledge of the delays. Thus, preferences were driven by past state-dependent gains, rather than by the joint effect of the birds' state at the time of choice and knowledge of the absolute properties of each alternative, as assumed in state-dependent, path-independent models of optimal choice.
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Kacelnik, A. (1990). R.C. Bolies and M.D. Beecher, Editors, Evolution and Learning, Lawrence Erlbaum, Hillsdale, New Jersey (1988), p. x. Anim. Behav., 40(3), 602–603.
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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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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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Brunner, D., Kacelnik, A., & Gibbon, J. (1992). Optimal foraging and timing processes in the starling, Sturnus vulgaris: effect of inter-capture interval. Anim. Behav., 44(4), 597–613.
Abstract: Laboratory experiments with starlings, Sturnus vulgaris, were conducted to investigate the interaction between timing and cost-benefit considerations. The design simulated an environment in which food was distributed in patches. Patches contained a random number of food items (N=0-4) separated by a fixed inter-capture interval or fixed interval. All patches ended with sudden depletion. The time elapsed since the last prey capture was the only way to detect the depletion of the patch. Once the patch was depleted a new patch could be reached by travelling between two perches. Three measures of timing were taken: (1) rate of working for food as function of `waiting' time in a patch, (2) the time of the last response in a patch or `giving-in' time, and (3) the time at which travel was initiated or `moving-on' time. The fixed interval that characterized patches was varied between conditions. The mean time of the peak in working rate was consistently centred around the fixed interval, while the other two measures of timing kept a roughly linear relation to the fixed interval, with slope greater than one. In accordance with Scalar Expectancy Theory, variability in the three forms of timing was proportional to the magnitude of the fixed interval. The birds seemed to take account of this increase in variability as shown by the mean value of their giving-up criterion. These results imply that information-processing constraints are important for modelling behavioural optimality.
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Cuthill, I. C., Kacelnik, A., Krebs, J. R., Haccou, P., & Iwasa, Y. (1990). Starlings exploiting patches: the effect of recent experience on foraging decisions. Anim. Behav., 40(4), 625–640.
Abstract: Laboratory and field experiments have shown that, as predicted by the marginal value model, starlings, Sturnus vulgaris, stay longer in a food patch when the average travel time between patches is long. A laboratory analogue of a patchy environment was used to investigate how starlings respond to rapidly fluctuating changes in travel time in order to find out the length of experience over which information is integrated. When there was a progressive increase in the amount of work required to obtain successive food items in a patch (experiment 1), birds consistently took more prey after long than after short travel times; travel experience before the most recent had no effect on the number of prey taken. Such behaviour does not maximize the rate of energy intake in this environment. The possibility that this is the result of a simple constraint on crop capacity is rejected as, when successive prey were equally easy to obtain up until a stepwise depletion of the patch (experiment 2), birds took equal numbers of prey per visit after long and short travel times: the rate-maximizing behaviour. A series of models are developed to suggest the possible constraints on optimal behaviour that affect starlings in the type of environment mimicked by experiment 1.
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Weir, A. A. S., & Kacelnik, A. (2006). A New Caledonian crow (Corvus moneduloides) creatively re-designs tools by bending or unbending aluminium strips. Anim. Cogn., 9(4), 317–334.
Abstract: Previous observations of a New Caledonian crow (Corvus moneduloides) spontaneously bending wire and using it as a hook [Weir et al. (2002) Science 297:981] have prompted questions about the extent to which these animals 'understand' the physical causality involved in how hooks work and how to make them. To approach this issue we examine how the same subject (“Betty”) performed in three experiments with novel material, which needed to be either bent or unbent in order to function to retrieve food. These tasks exclude the possibility of success by repetition of patterns of movement similar to those employed before. Betty quickly developed novel techniques to bend the material, and appropriately modified it on four of five trials when unbending was required. She did not mechanically apply a previously learned set of movements to the new situations, and instead sought new solutions to each problem. However, the details of her behaviour preclude concluding definitely that she understood and planned her actions: in some cases she probed with the unmodified tools before modifying them, or attempted to use the unmodified (unsuitable) end of the tool after modification. Gauging New Caledonian crows' level of understanding is not yet possible, but the observed behaviour is consistent with a partial understanding of physical tasks at a level that exceeds that previously attained by any other non-human subject, including apes.
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Brunner, D., Kacelnik, A., & Gibbon, J. (1996). Memory for inter-reinforcement interval variability and patch departure decisions in the starling,Sturnus vulgaris. Anim. Behav., 51(5), 1025–1045.
Abstract: An experiment with starlings was conducted to investigate the effect of variability in inter-reinforcement intervals on foraging decisions. The experimental design simulated an environment in which food was distributed in patches. Patches contained zero to four food items which could be collected by pecking at a key. All patches ended with sudden depletion. The time elapsed since the last reinforcement was the only way to detect the depletion of the patch. Once a patch was depleted, a new patch could be reached by completion of a travel requirement of 20 flights between two perches. Key pecks within a patch and the time of the last response in a patch (giving-in time) were recorded. The level of variability in the inter-reinforcement intervals was varied between different conditions. An increase in inter-reinforcement interval variability resulted in a flattening of response rate functions and giving-in time distributions, and in more asymmetry of the response functions, but not of the giving-in time distributions. Two theoretical models of decision making are presented, which differ in the assumptions about memory constraints. In one case, all inter-reinforcement intervals are remembered but in the other, only the intervals with extreme values are remembered. Both models accommodate response rates as a function of trial time, but only the second is compatible with the observed departure decision. Our results are compatible with net rate maximization.
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Weir, A. A. S., Chappell, J., & Kacelnik, A. (2002). Shaping of hooks in New Caledonian crows. Science, 297(5583), 981.
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Kenward, B., Rutz, C., Weir, A. A. S., & Kacelnik, A. (2006). Development of tool use in New Caledonian crows: inherited action patterns and social influences. Anim. Behav., 72(6), 1329–1343.
Abstract: New Caledonian crows, Corvus moneduloides, are the most advanced avian tool makers and tool users. We previously reported that captive-bred isolated New Caledonian crows spontaneously use twig tools and cut tools out of Pandanus spp. tree leaves, an activity possibly under cultural influence in the wild. However, what aspects of these behaviours are inherited and how they interact with individual and social experience remained unknown. To examine the interaction between inherited traits, individual learning and social transmission, we observed the ontogeny of twig tool use in hand-reared juveniles. Successful food retrieval was preceded by stereotyped object manipulation action patterns that resembled components of the mature behaviour, demonstrating that tool-oriented behaviours in this species are an evolved specialization. However, there was also an effect of social learning: juveniles that had received demonstrations of twig tool use by their human foster parent showed higher levels of handling and insertion of twigs than did their naive counterparts; a choice experiment showed that they preferred to handle objects that they had seen being manipulated by their human foster parent. Our observations are consistent with the hypothesis that individual learning, cultural transmission and creative problem solving all contribute to the acquisition of the tool-oriented behaviours in the wild, but inherited species-typical action patterns have a greater role than has been recognized.
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