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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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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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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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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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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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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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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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Call, J., Carpenter, M., & Tomasello, M. (2005). Copying results and copying actions in the process of social learning: chimpanzees (Pan troglodytes) and human children (Homo sapiens). Anim. Cogn., 8(3), 151–163.
Abstract: There is currently much debate about the nature of social learning in chimpanzees. The main question is whether they can copy others' actions, as opposed to reproducing the environmental effects of these actions using their own preexisting behavioral strategies. In the current study, chimpanzees (Pan troglodytes) and human children (Homo sapiens) were shown different demonstrations of how to open a tube-in both cases by a conspecific. In different experimental conditions, demonstrations consisted of (1) action only (the actions necessary to open the tube without actually opening it); (2) end state only (the open tube, without showing any actions); (3) both of these components (in a full demonstration); or (4) neither of these components (in a baseline condition). In the first three conditions subjects saw one of two different ways that the tube could open (break in middle; caps off ends). Subjects' behavior in each condition was assessed for how often they opened the tube, how often they opened it in the same location as the demonstrator, and how often they copied the demonstrator's actions or style of opening the tube. Whereas chimpanzees reproduced mainly the environmental results of the demonstrations (emulation), human children often reproduced the demonstrator's actions (imitation). Because the procedure used was similar in many ways to the procedure that Meltzoff (Dev Psych 31:1, 1995) used to study the understanding of others' unfulfilled intentions, the implications of these findings with regard to chimpanzees' understanding of others' intentions are also discussed.
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Dorrance, B. R., & Zentall, T. R. (2001). Imitative learning in Japanese quail (Coturnix japonica) depends on the motivational state of the observer quail at the time of observation. J Comp Psychol, 115(1), 62–67.
Abstract: The 2-action method was used to examine whether imitative learning in Japanese quail (Coturnix japonica) depends on the motivational state of the observer quail at the time of observation of the demonstrated behavior. Two groups of observers were fed before observation (satiated groups), whereas 2 other groups of observers were deprived of food before observation (hungry groups). Quail were tested either immediately following observation or after a 30-min delay. Results indicated that quail in the hungry groups imitated, whereas those in the satiated groups did not, regardless of whether their test was immediate or delayed. The results suggest that observer quail may not learn (through observation) behavior that leads to a reinforcer for which they are unmotivated at the time of test. In addition, the results show that quail are able to delay the performance of a response acquired through observation (i.e., they show deferred imitation).
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