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Subiaul, F., Cantlon, J. F., Holloway, R. L., & Terrace, H. S. (2004). Cognitive imitation in rhesus macaques. Science, 305(5682), 407–410.
Abstract: Experiments on imitation typically evaluate a student's ability to copy some feature of an expert's motor behavior. Here, we describe a type of observational learning in which a student copies a cognitive rule rather than a specific motor action. Two rhesus macaques were trained to respond, in a prescribed order, to different sets of photographs that were displayed on a touch-sensitive monitor. Because the position of the photographs varied randomly from trial to trial, sequences could not be learned by motor imitation. Both monkeys learned new sequences more rapidly after observing an expert execute those sequences than when they had to learn new sequences entirely by trial and error.
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Zentall, T. R. (2006). Imitation: definitions, evidence, and mechanisms. Anim. Cogn., 9(4), 335–353.
Abstract: Imitation can be defined as the copying of behavior. To a biologist, interest in imitation is focused on its adaptive value for the survival of the organism, but to a psychologist, the mechanisms responsible for imitation are the most interesting. For psychologists, the most important cases of imitation are those that involve demonstrated behavior that the imitator cannot see when it performs the behavior (e.g., scratching one's head). Such examples of imitation are sometimes referred to as opaque imitation because they are difficult to account for without positing cognitive mechanisms, such as perspective taking, that most animals have not been acknowledged to have. The present review first identifies various forms of social influence and social learning that do not qualify as opaque imitation, including species-typical mechanisms (e.g., mimicry and contagion), motivational mechanisms (e.g., social facilitation, incentive motivation, transfer of fear), attentional mechanisms (e.g., local enhancement, stimulus enhancement), imprinting, following, observational conditioning, and learning how the environment works (affordance learning). It then presents evidence for different forms of opaque imitation in animals, and identifies characteristics of human imitation that have been proposed to distinguish it from animal imitation. Finally, it examines the role played in opaque imitation by demonstrator reinforcement and observer motivation. Although accounts of imitation have been proposed that vary in their level of analysis from neural to cognitive, at present no theory of imitation appears to be adequate to account for the varied results that have been found.
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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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Gajdon, G. K., Fijn, N., & Huber, L. (2006). Limited spread of innovation in a wild parrot, the kea (Nestor notabilis). Anim. Cogn., 9(3), 173–181.
Abstract: In the local population of kea in Mount Cook Village, New Zealand, some keas open the lids of rubbish bins with their bill to obtain food scraps within. We investigated the extent to which this innovation has spread in the local population, and what factors limit the acquisition of bin opening. Only five males of 36 individually recognised birds were observed to have performed successful bin opening. With one exception there were always other keas present, watching successful bin opening. Seventeen additional individuals were seen to have benefitted from lid opening. Their foraging success was less than that of the bin openers. Social status of bin openers did not differ from scrounging males. Among the individuals that were regularly seen at the site of the bins but were not successful in bin opening, social status and the ratio of feeding directly from open bins correlated with the amount of opening attempts. We conclude that scrounging facilitated certain behavioural aspects of bin opening rather than inhibiting them. The fact that only 9% of opening attempts were successful, and the long period of time required to increase efficiency in lid opening shows that mainly individual experience, and to a lesser extent insight and social learning, play key roles in acquisition of the opening technique. The results indicate that the spread of innovative solutions of challenging mechanical problems in animals may be restricted to only a few individuals.
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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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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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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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Mercado, E. 3rd, Herman, L. M., & Pack, A. A. (2005). Song copying by humpback whales: themes and variations. Anim. Cogn., 8(2), 93–102.
Abstract: Male humpback whales (Megaptera novaeangliae) produce long, structured sequences of sound underwater, commonly called “songs.” Humpbacks progressively modify their songs over time in ways that suggest that individuals are copying song elements that they hear being used by other singers. Little is known about the factors that determine how whales learn from their auditory experiences. Song learning in birds is better understood and appears to be constrained by stable core attributes such as species-specific sound repertoires and song syntax. To clarify whether similar constraints exist for song learning by humpbacks, we analyzed changes over 14 years in the sounds used by humpback whales singing in Hawaiian waters. We found that although the properties of individual sounds within songs are quite variable over time, the overall distribution of certain acoustic features within the repertoire appears to be stable. In particular, our findings suggest that species-specific constraints on temporal features of song sounds determine song form, whereas spectral variability allows whales to flexibly adapt song elements.
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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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Whiten, A., Custance, D. M., Gomez, J. C., Teixidor, P., & Bard, K. A. (1996). Imitative learning of artificial fruit processing in children (Homo sapiens) and chimpanzees (Pan troglodytes). J Comp Psychol, 110(1), 3–14.
Abstract: Observational learning in chimpanzees and young children was investigated using an artificial fruit designed as an analog of natural foraging problems faced by primates. Each of 3 principal components could be removed in 2 alternative ways, demonstration of only one of which was watched by each subject. This permitted subsequent imitation by subjects to be distinguished from stimulus enhancement. Children aged 2-4 years evidenced imitation for 2 components, but also achieved demonstrated outcomes through their own techniques. Chimpanzees relied even more on their own techniques, but they did imitate elements of 1 component of the task. To our knowledge, this is the first experimental evidence of chimpanzee imitation in a functional task designed to simulate foraging behavior hypothesized to be transmitted culturally in the wild.
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