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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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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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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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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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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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Roberts, J., Hunter, M. L., & Kacelnik, A. (1981). The ground effect and acoustic communication. Anim. Behav., 29(2), 633–634.
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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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Fawcett, T. W., Skinner, A. M. J., & Goldsmith, A. R. (2002). A test of imitative learning in starlings using a two-action method with an enhanced ghost control. Anim. Behav., 64(4), 547–556.
Abstract: Imitative learning, in which an individual learns to reproduce the behaviour pattern of another, has attracted considerable attention as a potentially powerful form of social learning. Despite extensive research, however, it has proved difficult to demonstrate in nonhuman animals. We investigated the ability of European starlings, Sturnus vulgaris, to imitate the behaviour of a conspecific. Subjects watched a trained conspecific manipulating a plug for access to a food reward, using either a pushing or a pulling action. When later tested with the same apparatus these birds completed the task using the same action they had previously observed. In a second experiment, a separate group of starlings saw the plug move upwards or downwards automatically and a nearby conspecific obtain a food reward. When given access to the task these starlings failed to move the plug in the direction they had seen. Our experiment is an improvement on previous bidirectional control designs and provides strong evidence that starlings are capable of imitation. We advocate further use of this experimental design in attempts to demonstrate imitative learning. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.
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Coleman, K., & Wilson, D. S. (1998). Shyness and boldness in pumpkinseed sunfish: individual differences are context-specific. Anim. Behav., 56(4), 927–936.
Abstract: Natural selection often promotes a mix of behavioural phenotypes in a population. Adaptive variation in the propensity to take risks might explain individual differences in shyness and boldness in humans and other species. It is often implicitly assumed that shyness and boldness are general personality traits expressed across many situations. From the evolutionary standpoint, however, individual differences that are adaptive in one context (e.g. predator defence) may not be adaptive in other contexts (e.g. exploration of the physical environment or intraspecific social interactions). We measured the context specificity of shyness and boldness in a natural population of juvenile pumpkinseed sunfish,Lepomis gibbosus, by exposing the fish to a potentially threatening stimulus (a red-tipped metrestick extended towards the individual) and a nonthreatening stimulus (a novel food source). We also related these measures of shyness and boldness to behaviours observed during focal observations, both before and after the introduction of a predator (largemouth bass,Micropterus salmoides). Consistent individual differences were found within both contexts, but individual differences did not correlate across contexts. Furthermore, fish that were scored as intermediate in their response to the metrestick behaved most boldly as foragers and in response to the bass predators. These results suggest that shyness and boldness are context-specific and may not exist as a one-dimensional behavioural continuum even within a single context.
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Cloutier, S., Newberry, R. C., Honda, K., & Alldredge, J. R. (2002). Cannibalistic behaviour spread by social learning. Anim. Behav., 63(6), 1153–1162.
Abstract: We hypothesized that social learning is involved in the spread of cannibalism in domestic fowlGallus gallus domesticus . To investigate this hypothesis without harming birds, we used an inanimate chicken model as our cannibalism stimulus. We randomly assigned flocks of 12 White Leghorn pullets to one of two treatments: (1) flocks with two trained demonstrators (N=9) and (2) control flocks (N=8). Demonstrators were trained to pierce a membrane covering a dish of chicken blood and consume the blood. To assess the effect of access to the cannibalism stimulus during demonstrations, we randomly assigned observer pairs to one of two observer treatments: (1) observe stimulus through a wire mesh partition and (2) observe stimulus within the same enclosure. We conducted five 10-min demonstration sessions, each followed by a 10-min test of each observer pair in the absence of demonstrators, over a period of 15 days when the birds were 41-55 days of age, and two further tests at 63-64 and 91-92 days of age. Pairs that observed demonstrators piercing a membrane and consuming blood were more likely to perform this task when tested than control pairs. Learning of the task was enhanced by direct access to the cannibalism stimulus rather than observing it through a wire mesh partition. Blood consumption during tests was increased by direct access to the cannibalism stimulus during demonstration sessions. The birds made bigger holes in the membrane when tested after observing trained demonstrators and after having direct access to the stimulus. Our results provide the first experimental evidence that social learning can contribute to the spread of cannibalistic behaviour in domestic fowl. We suggest that stimulus enhancement and observational conditioning were the social-learning mechanisms involved. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.
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