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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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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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Biederman, G. B., Robertson, H. A., & Vanayan, M. (1986). Observational learning of two visual discriminations by pigeons: a within-subjects design. J Exp Anal Behav, 46(1), 45–49.
Abstract: Pigeon's observational learning of successive visual discrimination was studied using within-subject comparisons of data from three experimental conditions. Two pairs of discriminative stimuli were used; each bird was exposed to two of the three experimental conditions, with different pairs of stimuli used in a given bird's two conditions. In one condition, observers were exposed to visual discriminative stimuli only. In a second condition, subjects were exposed to a randomly alternating sequence of two stimuli where the one that would subsequently be used as S+ was paired with the operation of the grain magazine. In a third experimental condition, subjects were exposed to the performance of a conspecific in the operant discrimination procedure. After exposures to conspecific performances, there was facilitation of discriminative learning, relative to that which followed exposures to stimulus and reinforcement sequences or exposures to stimulus sequences alone. Exposure to stimulus and food-delivery sequences enhanced performance relative to exposure to stimulus sequences alone. The differential effects of these three types of exposure were not attributable to order effects or to task difficulty; rather, they clearly were due to the type of exposure.
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Bering, J. M. (2004). A critical review of the “enculturation hypothesis”: the effects of human rearing on great ape social cognition. Anim. Cogn., 7(4), 201–212.
Abstract: Numerous investigators have argued that early ontogenetic immersion in sociocultural environments facilitates cognitive developmental change in human-reared great apes more characteristic of Homo sapiens than of their own species. Such revamping of core, species-typical psychological systems might be manifest, according to this argument, in the emergence of mental representational competencies, a set of social cognitive skills theoretically consigned to humans alone. Human-reared great apes' capacity to engage in “true imitation,” in which both the means and ends of demonstrated actions are reproduced with fairly high rates of fidelity, and laboratory great apes' failure to do so, has frequently been interpreted as reflecting an emergent understanding of intentionality in the former. Although this epigenetic model of the effects of enculturation on social cognitive systems may be well-founded and theoretically justified in the biological literature, alternative models stressing behavioral as opposed to representational change have been largely overlooked. Here I review some of the controversy surrounding enculturation in great apes, and present an alternative nonmentalistic version of the enculturation hypothesis that can also account for enhanced imitative performance on object-oriented problem-solving tasks in human-reared animals.
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Akins, C. K., & Zentall, T. R. (1996). Imitative learning in male Japanese quail (Coturnix japonica) using the two-action method. J Comp Psychol, 110(3), 316–320.
Abstract: The study of imitative learning in animals has suffered from the presence of a number of confounding motivational and attentional factors (e.g., social facilitation and stimulus enhancement). The two-action method avoids these problems by exposing observers to demonstrators performing a response (e.g., operating a treadle) using 1 of 2 distinctive topographies (e.g., by pecking or by stepping). Japanese quail (Coturnix japonica) observers exposed to conspecific demonstrators showed a high correlation between the topography of the response they observed and the response they performed. These data provide strong evidence for the existence of true imitative learning in an active, precocious bird under conditions that control for alternative accounts.
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Akins, C. K., Klein, E. D., & Zentall, T. R. (2002). Imitative learning in Japanese quail (Coturnix japonica) using the bidirectional control procedure. Anim Learn Behav, 30(3), 275–281.
Abstract: In the bidirectional control procedure, observers are exposed to a conspecific demonstrator responding to a manipulandum in one of two directions (e.g., left vs. right). This procedure controls for socially mediated effects (the mere presence of a conspecific) and stimulus enhancement (attention drawn to a manipulandum by its movement), and it has the added advantage of being symmetrical (the two different responses are similar in topography). Imitative learning is demonstrated when the observers make the response in the direction that they observed it being made. Recently, however, it has been suggested that when such evidence is found with a predominantly olfactory animal, such as the rat, it may result artifactually from odor cues left on one side of the manipulandum by the demonstrator. In the present experiment, we found that Japanese quail, for which odor cues are not likely to play a role, also showed significant correspondence between the direction in which the demonstrator and the observer push a screen to gain access to reward. Furthermore, control quail that observed the screen move, when the movement of the screen was not produced by the demonstrator, did not show similar correspondence between the direction of screen movement observed and that performed by the observer. Thus, with the appropriate control, the bidirectional procedure appears to be useful for studying imitation in avian species.
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