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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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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.
Keywords: Animals; *Behavior, Animal; Evolution; Humans; *Imitative Behavior; Learning; Models, Animal
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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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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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Topál, J., Byrne, R. W., Miklósi, Á., & Csányi, V. (2006). Reproducing human actions and action sequences: “Do as I Do!” in a dog. Anim. Cogn., 9(4), 355–367.
Abstract: We present evidence that a dog (Philip, a 4-year-old tervueren) was able to use different human actions as samples against which to match his own behaviour. First, Philip was trained to repeat nine human-demonstrated actions on command ('Do it!'). When his performance was markedly over chance in response to demonstration by one person, testing with untrained action sequences and other demonstrators showed some ability to generalise his understanding of copying. In a second study, we presented Philip with a sequence of human actions, again using the 'Do as I do' paradigm. All demonstrated actions had basically the same structure: the owner picked up a bottle from one of six places; transferred it to one of the five other places and then commanded the dog ('Do it!'). We found that Philip duplicated the entire sequence of moving a specific object from one particular place to another more often than expected by chance. Although results point to significant limitations in his imitative abilities, it seems that the dog could have recognized the action sequence, on the basis of observation alone, in terms of the initial state, the means, and the goal. This suggests that dogs might acquire abilities by observation that enhance their success in complex socio-behavioural situations.
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Fripp, D., Owen, C., Quintana-Rizzo, E., Shapiro, A., Buckstaff, K., Jankowski, K., et al. (2005). Bottlenose dolphin (Tursiops truncatus) calves appear to model their signature whistles on the signature whistles of community members. Anim. Cogn., 8(1), 17–26.
Abstract: Bottlenose dolphins are unusual among non-human mammals in their ability to learn new sounds. This study investigates the importance of vocal learning in the development of dolphin signature whistles and the influence of social interactions on that process. We used focal animal behavioral follows to observe six calves in Sarasota Bay, Fla., recording their social associations during their first summer, and their signature whistles during their second. The signature whistles of five calves were determined. Using dynamic time warping (DTW) of frequency contours, the calves' signature whistles were compared to the signature whistles of several sets of dolphins: their own associates, the other calves' associates, Tampa Bay dolphins, and captive dolphins. Whistles were considered similar if their DTW similarity score was greater than those of 95% of the whistle comparisons. Association was defined primarily in terms of time within 50 m of the mother/calf pair. On average, there were six dolphins with signature whistles similar to the signature whistles of each of the calves. These were significantly more likely to be Sarasota Bay resident dolphins than non-Sarasota dolphins, and (though not significantly) more likely to be dolphins that were within 50 m of the mother and calf less than 5% of the time. These results suggest that calves may model their signature whistles on the signature whistles of members of their community, possibly community members with whom they associate only rarely.
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Scheumann, M., & Call, J. (2004). The use of experimenter-given cues by South African fur seals (Arctocephalus pusillus). Anim. Cogn., 7(4), 224–230.
Abstract: Dogs can use a variety of experimenter-given cues such as pointing, head direction, and eye direction to locate food hidden under one of several containers. Some authors have proposed that this is a result of the domestication process. In this study we tested four captive fur seals in a two alternative object choice task in which subjects had to use one of the following experimenter-given cues to locate the food: (1) the experimenter pointed and gazed at one of the objects, (2) the experimenter pointed at only one of the objects, (3) the experimenter gazed at only one of the objects, (4) the experimenter glanced at only one of the objects, (5) the experimenter pointed and gazed at one of the objects but was sitting closer to one object than to the other, (6) the experimenter pointed only with the index finger at one of the objects, (7) the experimenter presented a replica of one of the objects. The fur seals were able to use cues which involved a fully exposed arm or a head direction, but failed to use glance only, the index finger pointing and the object replica cues. The results showed that a domestication process was not necessary to develop receptive skills to cues given by an experimenter. Instead, we hypothesize that close interactions with humans prior to testing enabled fur seals to uses ome gestural cues without formal training. We also analyzed the behavior of the seals depending on the level of difficulty of the task. Behavioral signs of hesitation increased with task difficulty. This suggests that the fur seals were sensitive to task difficulty.
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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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