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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.
Keywords: Animals; *Behavior, Animal; Columbidae; Cues; *Imitative Behavior; *Learning; Odors; Sound
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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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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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Biro, D., Inoue-Nakamura, N., Tonooka, R., Yamakoshi, G., Sousa, C., & Matsuzawa, T. (2003). Cultural innovation and transmission of tool use in wild chimpanzees: evidence from field experiments. Anim. Cogn., 6(4), 213–223.
Abstract: Chimpanzees (Pan troglodytes) are the most proficient and versatile users of tools in the wild. How such skills become integrated into the behavioural repertoire of wild chimpanzee communities is investigated here by drawing together evidence from three complementary approaches in a group of oil-palm nut- ( Elaeis guineensis) cracking chimpanzees at Bossou, Guinea. First, extensive surveys of communities adjacent to Bossou have shown that population-specific details of tool use, such as the selection of species of nuts as targets for cracking, cannot be explained purely on the basis of ecological differences. Second, a 16-year longitudinal record tracing the development of nut-cracking in individual chimpanzees has highlighted the importance of a critical period for learning (3-5 years of age), while the similar learning contexts experienced by siblings have been found to result in near-perfect (13 out of 14 dyads) inter-sibling correspondence in laterality. Third, novel data from field experiments involving the introduction of unfamiliar species of nuts to the Bossou group illuminates key aspects of both cultural innovation and transmission. We show that responses of individuals toward the novel items differ markedly with age, with juveniles being the most likely to explore. Furthermore, subjects are highly specific in their selection of conspecifics as models for observation, attending to the nut-cracking activities of individuals in the same age group or older, but not younger than themselves. Together with the phenomenon of inter-community migration, these results demonstrate a mechanism for the emergence of culture in wild chimpanzees.
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Nicol, C. J. (2004). Development, direction, and damage limitation: social learning in domestic fowl. Learn Behav, 32(1), 72–81.
Abstract: This review highlights two areas of particular interest in the study of social learning in fowl. First, the role of social learning in the development of feeding and foraging behavior in young chicks and older birds is described. The role of the hen as a demonstrator and possible teacher is considered, and the subsequent social influence of brood mates and other companions on food avoidance and food preference learning is discussed. Second, the way in which work on domestic fowl has contributed to an understanding of the importance of directed social learning is examined. The well-characterized hierarchical social organization of small chicken flocks has been used to design studies which demonstrate that the probability of social transmission is strongly influenced by social relationships between birds. The practical implications of understanding the role of social learning in the spread of injurious behaviors in this economically important species are briefly considered.
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Zentall, T. R. (2004). Action imitation in birds. Learn Behav, 32(1), 15–23.
Abstract: Action imitation, once thought to be a behavior almost exclusively limited to humans and the great apes, surprisingly also has been found in a number of bird species. Because imitation has been viewed by some psychologists as a form of intelligent behavior, there has been interest in how it is distributed among animal species. Although the mechanisms responsible for action imitation are not clear, we are now at least beginning to understand the conditions under which it occurs. In this article, I try to identify and differentiate the various forms of socially influenced behavior (species-typical social reactions, social effects on motivation, social effects on perception, socially influenced learning, and action imitation) and explain why it is important to differentiate imitation from other forms of social influence. I also examine some of the variables that appear to be involved in the occurrence of imitation. Finally, I speculate about why a number of bird species, but few mammal species, appear to imitate.
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Whiten, A., Horner, V., Litchfield, C. A., & Marshall-Pescini, S. (2004). How do apes ape? Learn. Behav., 32(1), 36–52.
Abstract: In the wake of telling critiques of the foundations on which earlier conclusions were based, the last 15 years have witnessed a renaissance in the study of social learning in apes. As a result, we are able to review 31 experimental studies from this period in which social learning in chimpanzees, gorillas, and orangutans has been investigated. The principal question framed at the beginning of this era, Do apes ape? has been answered in the affirmative, at least in certain conditions. The more interesting question now is, thus, How do apes ape? Answering this question has engendered richer taxonomies of the range of social-learning processes at work and new methodologies to uncover them. Together, these studies suggest that apes ape by employing a portfolio of alternative social-learning processes in flexibly adaptive ways, in conjunction with nonsocial learning. We conclude by sketching the kind of decision tree that appears to underlie the deployment of these alternatives.
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Caldwell, C. A., & Whiten, A. (2004). Testing for social learning and imitation in common marmosets, Callithrix jacchus, using an artificial fruit. Anim. Cogn., 7(2), 77–85.
Abstract: We tested for social learning and imitation in common marmosets using an artificial foraging task and trained conspecific demonstrators. We trained a demonstrator marmoset to open an artificial fruit, providing a full demonstration of the task to be learned. Another marmoset provided a partial demonstration, controlling for stimulus enhancement effects, by eating food from the outside of the apparatus. We thus compared three observer groups, each consisting of four animals: those that received the full demonstration, those that received the partial demonstration, and a control group that saw no demonstration prior to testing. Although none of the observer marmosets succeeded in opening the artificial fruit during the test periods, there were clear effects of demonstration type. Those that saw the full demonstration manipulated the apparatus more overall, whereas those from the control group manipulated it the least of the three groups. Those from the full-demonstration group also contacted the particular parts of the artificial fruit that they had seen touched (localised stimulus enhancement) to a greater extent than the other two groups. There was also an interaction between the number of hand and mouth touches made to the artificial fruit for the full- and partial-demonstration groups. Whether or not these data represent evidence for imitation is discussed. We also propose that the clear differences between the groups suggest that social learning mechanisms provide real benefits to these animals in terms of developing novel food-processing skills analogous to the one presented here.
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