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Heyes, C. (2012). What's social about social learning? J Comp Psychol, 120.
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Heyes, C. M. (2002). Transformation and associative theories of imitation. In K. Dautenhahn, & C. L. Nehaniv (Eds.), Imitation in animals and artefacts (pp. 501–523). Cambridge, MA.: MIT Press.
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Ray, E. D., Gardner, M. R., & Heyes, C. M. (2000). Seeing how it's done: matching conditions for observer rats (Rattus norvegicus) in the bidirectional control. Anim. Cogn., 3(3), 147–157.
Abstract: In an attempt to increase the reliability of the demonstrator-consistent responding effect produced in the bidirectional control procedure, experiments 1-4 sought conditions that would magnify the matching effect. The aim was to produce a robust demonstrator-consistent responding effect in order that future analytic experiments could investigate the psychological processes responsible for this effect. The joystick responses of observer rats trained using the standard bidirectional control procedure parameters were compared with those of observers subject to conditions identified in the social learning literature as favourable for imitation. Unlike mice, observer rats in experiments 1 a and 1 b tended to push a joystick in the same direction as their demonstrators when the demonstrators were either familiar or unfamiliar males and females. Comparable demonstrator-consistent responding occurred following observation of a standard and a salient joystick response (experiment 2). Experiment 3 showed that the discriminative accuracy of a demonstrator's responding was important for matching behaviour, and suggested that matching might be enhanced with more than the conventional single observation session. Experiment 4 confirmed that the bidirectional control effect is sensitive to the amount of observational experience; after six observation sessions, demonstrator-inconsistent responding occurs. The results of experiments 1-3 are, and those of experiment 4 are not, compatible with the hypothesis that demonstrator-consistent responding in the bidirectional control is caused by olfactory cues deposited by demonstrators on the joystick.
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Heyes, C. M. (1993). Imitation, culture and cognition. Anim. Behav., 46(5), 999–1010.
Abstract: Abstract. This paper examines the significance of imitation in non-human animals with respect to the phylogenetic origins of culture and cognitive complexity. It is argued that both imitation (learning about behaviour through nonspecific observation) and social learning (learning about the environment through conspecific observation) can mediate social transmission of information, and that neither is likely to play an important role in supporting behavioural traditions or culture. Current evidence suggests that imitation is unlikely to do this because it does not insulate information from modification through individual learning in the retention period between acquisition and re-transmission. Although insignificant in relation to culture, imitation apparently involves complex and little-understood cognitive operations. It is unique in requiring animals spontaneously to equate extrinsic visual input with proprioceptive and/or kinaesthetic feedback from their own actions, but not in requiring or implicating self-consciousness, representation, metarepresentation or a capacity for goal-directed action.
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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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Heyes, C., & Galef, B. G. (Eds.). (1996). Social learning in animals: the roots of culture. San Diego, CA: Academic Press, Inc.
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Campbell, F. M., & Heyes, C. M. (2002). Rats smell: odour-mediated local enhancement, in a vertical movement two-action test. Anim. Behav., 63(6), 1055–1063.
Abstract: In two experiments, hungry rats, Rattus norvegicus, were present in one side of an operant chamber while a conspecific demonstrator in the adjacent compartment moved a single lever either up or down for a food reward. During a subsequent test session, in which these rats were allowed access to the lever for the first time, all responses were rewarded regardless of their direction. In experiment 1, rats that were prevented from observing the direction of lever movement by means of a screen showed a reliable demonstrator-consistent response bias, while rats that had observed the direction of lever movement and in addition had access to any odour cues deposited on the lever did not. In experiment 2, each rat observed another rat (the `viewed' demonstrator) moving a lever either up or down. They were then transferred into the test compartment of a different operant chamber in which another rat (the `box' demonstrator) had moved the lever in the same direction as the viewed demonstrator or in the opposite direction. These observer rats showed a reliable preference for their box demonstrator's direction, but responded in the opposite direction to their viewed demonstrator. Taken together, the results of these experiments suggest that directional responding by rats in a vertical movement two-action test is influenced by demonstrator-deposited odour cues in addition to visual experience of a demonstrator's behaviour. Furthermore, while odour-mediated local enhancement gave rise to demonstrator-consistent responding, visual observation of a conspecific appeared to have the reverse effect. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.
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Campbell, F. M., Heyes, C. M., & Goldsmith, A. R. (1999). Stimulus learning and response learning by observation in the European starling, in a two-object/two-action test. Anim. Behav., 58(1), 151–158.
Abstract: Juvenile European starlings, Sturnus vulgaris , were allowed to observe a conspecific demonstrator using its beak to remove one of two distinctively coloured objects (i.e. a red or a black plug) from a hole in the lid of a plastic box. Both plugs could be removed by either pulling up on a loop of string inserted through the centre of the plug, or pushing down on the plug. When subsequently allowed access to the plugs, and rewarded with food for all removal responses, regardless of the object to which they were made and their direction, observer birds removed the same plug in the same direction as their demonstrator. These results suggest that the two-object/two-action paradigm is a valuable procedure for testing for the simultaneous effects of learning about a stimulus and a response, an object and an action, through conspecific observation.
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Heyes, C. M., & Dawson, G. R. (1990). A demonstration of observational learning in rats using a bidirectional control. Q J Exp Psychol B, 42(1), 59–71.
Abstract: Hungry rats observed a conspecific demonstrator pushing a single manipulandum, a joystick, to the right or to the left for food reward and were then allowed access to the joystick from a different orientation. The effects of right-pushing vs left-pushing observation experience on (1) response acquisition, (2) reversal of a left-right discrimination, and (3) responding in extinction, were examined. Rats that had observed left-pushing made more left responses during acquisition than rats that had observed right-pushing, and rats that had observed demonstrators pushing in the direction that had previously been reinforced took longer to reach criterion reversal and made more responses in extinction than rats that had observed demonstrators pushing in the opposite direction to that previously reinforced. These results provide evidence that rats are capable of learning a response, or a response-reinforcer contingency, through conspecific observation.
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Reed, P., Skiera, F., Adams, L., & Heyes, C. M. (1996). Effects of Isolation Rearing and Mirror Exposure on Social and Asocial Discrimination Performance. Learn. Motiv., 27(2), 113–129.
Abstract: Four experiments examined the effects of rearing in isolation on rats performance on discrimination-based and social learning tasks. After demonstrating that the rearing procedures produced similar results in an open field task to those previously established (Experiment 1), rats were subjected to two discrimination tasks: an instrumental occasion setting procedure (Experiment 3) and a nonspatial win-stay/lose-shift versus win-shift/lose-stay procedure (Experiment 4). Deficits in acquisition of the necessary discriminations were noted in the rats raised in isolation, but there were no differences between isolation-reared and socially reared subjects in response acquisition per se. In Experiment 2, rats were presented with an observational learning task using the bidirectional control procedure. Socially reared rats had a tendency to imitate the behavior they had observed, but rats raised in isolation performed the opposite behavior to that observed, indicating a failure to use a conspecific as a reference point in the task. The presence of a mirror during rearing in isolation was also investigated, but was found to have little effect in attenuating the above deficits in behavior.
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