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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. (2012). What's social about social learning? J Comp Psychol, 120.
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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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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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Heyes, C. M. (1995). Imitation and flattery: a reply to Byrne & Tomasello. Anim. Behav., 50(5), 1421–1424.
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Heyes, C. M. (1994). Social learning in animals: categories and mechanisms. Biol. Rev., 69(2), 207–231.
Abstract: There has been relatively little research on the psychological mechanisms of social learning. This may be due, in part, to the practice of distinguishing categories of social learning in relation to ill-defined mechanisms (Davis, 1973; Galef, 1988). This practice both makes it difficult to identify empirically examples of different types of social learning, and gives the false impression that the mechanisms responsible for social learning are clearly understood. It has been proposed that social learning phenomena be subsumed within the categorization scheme currently used by investigators of asocial learning. This scheme distinguishes categories of learning according to observable conditions, namely, the type of experience that gives rise to a change in an animal (single stimulus vs. stimulus-stimulus relationship vs. response-reinforcer relationship), and the type of behaviour in which this change is detected (response evocation vs. learnability) (Rescorla, 1988). Specifically, three alignments have been proposed: (i) stimulus enhancement with single stimulus learning, (ii) observational conditioning with stimulus-stimulus learning, or Pavlovian conditioning, and (iii) observational learning with response-reinforcer learning, or instrumental conditioning. If, as the proposed alignments suggest, the conditions of social and asocial learning are the same, there is some reason to believe that the mechanisms underlying the two sets of phenomena are also the same. This is so if one makes the relatively uncontroversial assumption that phenomena which occur under similar conditions tend to be controlled by similar mechanisms. However, the proposed alignments are intended to be a set of hypotheses, rather than conclusions, about the mechanisms of social learning; as a basis for further research in which animal learning theory is applied to social learning. A concerted attempt to apply animal learning theory to social learning, to find out whether the same mechanisms are responsible for social and asocial learning, could lead both to refinements of the general theory, and to a better understanding of the mechanisms of social learning. There are precedents for these positive developments in research applying animal learning theory to food aversion learning (e.g. Domjan, 1983; Rozin & Schull, 1988) and imprinting (e.g. Bolhuis, de Vox & Kruit, 1990; Hollis, ten Cate & Bateson, 1991). Like social learning, these phenomena almost certainly play distinctive roles in the antogeny of adaptive behaviour, and they are customarily regarded as 'special kinds' of learning (Shettleworth, 1993).(ABSTRACT TRUNCATED AT 400 WORDS)
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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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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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McGregor, A., Saggerson, A., Pearce, J., & Heyes, C. (2006). Blind imitation in pigeons, Columba livia. Anim. Behav., 72(2), 287–296.
Abstract: Pigeons that had been trained with a food reward both to peck at and to step on a horizontal plate were allowed to observe a conspecific demonstrator pecking at or stepping on the plate before a test in which the observers were not rewarded for either pecking or stepping. In experiment 1, the demonstrators were not rewarded while being observed. In spite of this, the observers provided evidence of imitation: those that had observed pecking made a greater proportion of pecking responses on test than observers of stepping. In experiment 2, each observer was exposed to a pecking or a stepping conspecific on two occasions. On one occasion, the demonstrator received a food reward for each demonstrated response (continuous reinforcement condition), and on the other the demonstrator's responses were rewarded only rarely (variable interval condition). The observers provided equally strong evidence of imitation in each of these conditions; on test, they made proportionally more of the observed response both when the demonstrators had been richly rewarded and when they had been rarely rewarded. These results show that pigeons engage in `blind' imitation, that is, their imitative behaviour is not always guided by observational learning about response outcomes.
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