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Dorrance, B. R., & Zentall, T. R. (2002). Imitation of conditional discriminations in pigeons (Columba livia). J Comp Psychol, 116(3), 277–285.
Abstract: In the present experiments, the 2-action method was used to determine whether pigeons could learn to imitate a conditional discrimination. Demonstrator pigeons (Columba livia) stepped on a treadle in the presence of 1 light and pecked at the treadle in the presence of another light. Demonstration did not seem to affect acquisition of the conditional discrimination (Experiment 1) but did facilitate its reversal of the conditional discrimination (Experiments 2 and 3). The results suggest that pigeons are not only able to learn a specific behavior by observing another pigeon, but they can also learn under which circumstances to perform that behavior. The results have implications for proposed mechanisms of imitation in animals.
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Ray, E. D., & Heyes, C. M. (2002). Do rats in a two-action test encode movement egocentrically or allocentrically? Anim. Cogn., 5(4), 245–252.
Abstract: Two-action tests of imitation compare groups that observe topographically different responses to a common manipulandum. The general aim of the two experiments reported here was to find a demonstrator-consistent responding effect in a procedure that could be elaborated to investigate aspects of what was learned about the demonstrated lever response. Experiment 1 was a pilot study with rats of a variant of the two-action method of investigating social learning about observed responses. Groups of observer rats ( Rattus norvegicus) saw a demonstrator push a lever up or down for a food reward. When these observers were subsequently given access to the lever and rewarded for responses in both directions, their directional preferences were compared with two 'screen control' groups that were unable to see their demonstrators' behaviour. Demonstrator-consistent responding was found to be restricted to observers that were able to see demonstrator performance, suggesting that scent cues alone were insufficient to cue a preference for the demonstrators' response direction and thereby that the rats learned by observation about body movements (imitation) or lever movement (emulation). Experiment 2 assessed responding on two levers, one that had been manipulated by the demonstrator, and a second, transposed lever positioned some distance away. Demonstrator-consistent responding was abolished when actions were observed and performed in different parts of the apparatus, suggesting that observed movement was encoded allocentrically with respect to the apparatus rather than egocentrically with respect to the actor's body. With particular reference to the influence of scent cues, the results are discussed in relation to the strengths and weaknesses of this and other varieties of the two-action procedure as tests of imitation in animals and human infants.
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Caldwell, C. A., & Whiten, A. (2002). Evolutionary perspectives on imitation: is a comparative psychology of social learning possible? Anim. Cogn., 5(4), 193–208.
Abstract: Studies of imitation in animals have become numerous in recent times, but do they contribute to a comparative psychology of social learning? We review this burgeoning field to identify the problems and prospects for such a goal. Difficulties of two main kinds are identified. First, researchers have tackled questions about social learning from at least three very different theoretical perspectives, the “phylogenetic”, “animal model”, and “adaptational”. We examine the conflicts between them and consider the scope for integration. A second difficulty arises in the methodological approaches used in the discipline. In relation to one of these – survey reviews of published studies – we tabulate and compare the contrasting conclusions of nine articles that together review 36 studies. The basis for authors' disagreements, including the matters of perceptual opacity, novelty, sequential structure, and goal representation, are examined. In relation to the other key method, comparative experimentation, we identify 12 studies that have explicitly compared species' imitative ability on similar tasks. We examine the principal problems of comparing like with like in these studies and consider solutions, the most powerful of which we propose to be the use of a systematic range of task designs, rather than any single “gold standard” task.
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Bjorklund, D. F., Yunger, J. L., Bering, J. M., & Ragan, P. (2002). The generalization of deferred imitation in enculturated chimpanzees (Pan troglodytes). Anim. Cogn., 5(1), 49–58.
Abstract: Deferred imitation of object-related actions and generalization of imitation to similar but not identical tasks was assessed in three human-reared (enculturated) chimpanzees, ranging in age from 5 to 9 years. Each ape displayed high levels of deferred imitation and only slightly lower levels of generalization of imitation. The youngest two chimpanzees were more apt to generalize the model's actions when they had displayed portions of the target behaviors at baseline, consistent with the idea that learning is more likely to occur when working within the “zone of proximal development.” We argue that generalization of imitation is the best evidence to date of imitative learning in chimpanzees.
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Klein, E. D., & Zentall, T. R. (2003). Imitation and affordance learning by pigeons (Columba livia). J Comp Psychol, 117(4), 414–419.
Abstract: The bidirectional control procedure was used to determine whether pigeons (Columba livia) would imitate a demonstrator that pushed a sliding screen for food. One group of observers saw a trained demonstrator push a sliding screen door with its beak (imitation group), whereas 2 other groups watched the screen move independently (possibly learning how the environment works) with a conspecific either present (affordance learning with social facilitation) or absent (affordance learning alone). A 4th group could not see the screen being pushed (sound and odor control). Imitation was evidenced by the finding that pigeons that saw a demonstrator push the screen made a higher proportion of matching screen pushes than observers in 2 appropriate control conditions. Further, observers that watched a screen move without a demonstrator present made a significantly higher proportion of matching screen pushes than would be expected by chance. Thus, these pigeons were capable of affordance learning.
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Horowitz, A. C. (2003). Do humans ape? Or do apes human? Imitation and intention in humans (Homo sapiens) and other animals. J Comp Psychol, 117(3), 325–336.
Abstract: A. Whiten, D. M. Custance, J.-C. Gomez, P. Teixidor, and K. A. Bard (1996) tested chimpanzees' (Pan troglodytes) and human children's (Homo sapiens) skills at imitation with a 2-action test on an “artificial fruit.” Chimpanzees imitated to a restricted degree; children were more thoroughly imitative. Such results prompted some to assert that the difference in imitation indicates a difference in the subjects' understanding of the intentions of the demonstrator (M. Tomasello, 1996). In this experiment, 37 adult human subjects were tested with the artificial fruit. Far from being perfect imitators, the adults were less imitative than the children. These results cast doubt on the inference from imitative performance to an ability to understand others' intentions. The results also demonstrate how any test of imitation requires a control group and attention to the level of behavioral analysis.
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Stoinski, T. S., & Whiten, A. (2003). Social learning by orangutans (Pongo abelii and Pongo pygmaeus) in a simulated food-processing task. J Comp Psychol, 117(3), 272–282.
Abstract: Increasing evidence for behavioral differences between populations of primates has created a resurgence of interest in examining mechanisms of information transfer between individuals. The authors examined the social transmission of information in 15 captive orangutans (Pongo abelii and Pongo pygmaeus) using a simulated food-processing task. Experimental subjects were shown 1 of 2 methods for removing a suite of defenses on an “artificial fruit.” Control subjects were given no prior exposure before interacting with the fruit. Observing a model provided a functional advantage in the task, as significantly more experimental than control subjects opened the fruit. Within the experimental groups, the authors found a trend toward differences in the actual behaviors used to remove 1 of the defenses. Results support observations from the wild implying horizontal transfer of information in orangutans and show that a number of social learning processes are likely to be involved in the transfer of knowledge in this species.
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Matsuzawa, T. (2003). The Ai project: historical and ecological contexts. Anim. Cogn., 6(4), 199–211.
Abstract: This paper aims to review a long-term research project exploring the chimpanzee mind within historical and ecological contexts. The Ai project began in 1978 and was directly inspired by preceding ape-language studies conducted in Western countries. However, in contrast with the latter, it has focused on the perceptual and cognitive capabilities of chimpanzees rather than communicative skills between humans and chimpanzees. In the original setting, a single chimpanzee faced a computer-controlled apparatus and performed various kinds of matching-to-sample discrimination tasks. Questions regarding the chimpanzee mind can be traced back to Wolfgang Koehler's work in the early part of the 20th century. Yet, Japan has its unique natural and cultural background: it is home to an indigenous primate species, the Japanese snow monkey. This fact has contributed to the emergence of two previous projects in the wild led by the late Kinji Imanishi and his students. First, the Koshima monkey project began in 1948 and became famous for its discovery of the cultural propagation of sweet-potato washing behavior. Second, pioneering work in Africa, starting in 1958, aimed to study great apes in their natural habitat. Thanks to the influence of these intellectual ancestors, the present author also undertook the field study of chimpanzees in the wild, focusing on tool manufacture and use. This work has demonstrated the importance of social and ecological perspectives even for the study of the mind. Combining experimental approaches with a field setting, the Ai project continues to explore cognition and behavior in chimpanzees, while its focus has shifted from the study of a single subject toward that of the community as a whole.
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Hirata, S., & Celli, M. L. (2003). Role of mothers in the acquisition of tool-use behaviours by captive infant chimpanzees. Anim. Cogn., 6(4), 235–244.
Abstract: This article explores the maternal role in the acquisition of tool-use behaviours by infant chimpanzees ( Pan troglodytes). A honey-fishing task, simulating ant/termite fishing found in the wild, was introduced to three dyads of experienced mother and naive infant chimpanzees. Four fishing sites and eight sets of 20 objects to be used as tools, not all appropriate, were available. Two of the mothers constantly performed the task, using primarily two kinds of tools; the three infants observed them. The infants, regardless of the amount of time spent observing, successfully performed the task around the age of 20-22 months, which is earlier than has been recorded in the wild. Two of the infants used the same types of tools that the adults predominantly used, suggesting that tool selectivity is transmitted. The results also show that adults are tolerant of infants, even if unrelated; infants were sometimes permitted to lick the tools, or were given the tools, usually without honey, as well as permitted to observe the adult performances closely.
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Sousa, C., Okamoto, S., & Matsuzawa, T. (2003). Behavioural development in a matching-to-sample task and token use by an infant chimpanzee reared by his mother. Anim. Cogn., 6(4), 259–267.
Abstract: We investigated the behavioural and cognitive development of a captive male infant chimpanzee, Ayumu, raised by his mother, Ai. Here we report Ayumu's achievements up to the age of 2 years and 3 months, in the context of complex computer-controlled tasks. From soon after birth, Ayumu had been present during an experiment performed by his mother. The task consisted of two phases, a matching-to-sample task in which she received token rewards, and the insertion of these tokens into a vending machine to obtain food rewards. Ayumu himself received no reward or encouragement from humans for any of the actions he exhibited during the experiment. At the age of 9 months and 3 weeks, Ayumu performed his first matching-to-sample trial. At around 1 year and 3 months, he began to perform them consistently. Also during this period, he frequently stole food rewards from his mother. At 2 years and 3 months, Ayumu succeeded for the first time in inserting a token into the vending machine. Once he had succeeded in using a token, he performed both phases of the task in sequence 20 times consecutively. The infant's behaviour was not shaped by food rewards but by a strong motivation to copy his mother's behaviour. Our observations of Ayumu thus mirror the learning processes shown by wild chimpanzees.
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