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Imitation: Is Cognitive Neuroscience Solving the Correspondence Problem?

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Imitation: Is Cognitive Neuroscience Solving the Correspondence Problem? ARTICLE IN PRESS TICS 384 Review TRENDS in Cognitive Sciences Vol.xx No.xx Monthxxxx Imitation: is cognitive neuroscience solving the correspondence problem? Marcel Brass1 and Cecilia Heyes2 1Department of Cognitive Neurology, Max Planck Institute for Human Cognitive and Brain Sciences, Stephanstr. 1A, 04103 Leipzig, Germany 2Department of Psychology, University College London, Gower Street, London WC1E 6BT, UK Imitation poses a unique problem: how does the imi- framework (generalist theories) or whether it depends on tator know what pattern of motor activation will make a special purpose mechanism (specialist theories). We their action look like that of the model? Specialist then review research on the role of learning in imitation theories suggest that this correspondence problem has and observation of biological motion. Finally, we discuss a unique solution; there are functional and neurological a problem for generalist theories: If imitation depends on mechanisms dedicated to controlling imitation. General- ist theories propose that the problem is solved by general mechanisms of associative learning and action control. Box 1. Mirror neurons: What do they do? What are they for? Recent research in cognitive neuroscience, stimulated Mirror neurons in the premotor area F5 of monkeys are active both by the discovery of mirror neurons, supports generalist when the animal observes and when it executes a specific action solutions. Imitation is based on the automatic activation (for a review see [52,53]. The discovery of these cells has had a revolutionary impact, turning perception–action interaction into a of motor representations by movement observation. focus of intensive, interdisciplinary research worldwide. Naturally These externally triggered motor representations are there has been a great deal of speculation about the function of then used to reproduce the observed behaviour. This mirror neurons, including proposals that they mediate, not only imitative capacity depends on learned perceptual-motor imitation, but also action understanding, empathy, language deve- links. Finally, mechanisms distinguishing self from other lopment [54,55], and action simulation [56]. However, at present, direct experimental evidence for the involvement of mirror neurons are implicated in the inhibition of imitative behaviour. in one or all these functions is relatively weak. Progress in research on the function of mirror neurons might have been hampered by a failure to distinguish clearly between two questions: What do they do?, What are they for? Imitation might well Introduction be one of the things that mirror neurons do; under some conditions, Could you learn to tango by telephone? Maybe, but it in some species, mirror neurons could be involved in the generation of imitative behaviour. However, mirror neurons could do imitation would be much easier to learn by watching the steps than without being for imitation; they could be involved in generating by listening to instructions. When movements are visible, imitative behaviour without imitation being the function that we can copy them; we can just do what we see. Imitation – favoured their evolution by natural selection. In other words, imi- copying body movement – appears to be simple. However, tation and other functions of mirror neurons could be exaptations the ease with which humans imitate raises a question, rather than adaptations [57]. Generalist theories of imitation (see text) imply that mirror sometimes known as the correspondence problem [1], that neurons – and other neural systems that are active during both is proving difficult to answer: When we observe another action observation and action execution – can do imitation but are person moving we do not see the muscle activation under- not for imitation. They suggest that mirror neurons acquire their lying their movement but rather the external conse- properties in the course of ontogeny as a side-effect of the operation quences of that activation. So how does the observer’s of general associative learning and action control processes (see [58] for a similar view). Once established, mirror neurons might be motor system ‘know’ which muscle activations will lead to involved in the generation of imitative behaviour (and in several the observed movement? other functions), but, according to generalist theories, the properties It is important to solve the correspondence problem of mirror neurons are not innate, and the learning and action-control because imitation provides a foundation for language processes that lead to their formation evolved in response to much acquisition, skill learning, socialisation, and encultura- more general adaptive problems. Empirically, the question of whether there are neurons with mirror properties present at birth tion. A review of current progress on this problem is timely remains unresolved. However, the finding that mirror neurons for because the discovery of mirror neurons has stimulated tool use can develop during ontogeny [59] suggests that learning considerable interest in action observation and imitation plays a crucial role. among cognitive neuroscientists, and the function of these Monkeys have mirror neurons but, according to some authors, they do not imitate. If this is true, it is evidence against the hypo- intriguing cells is itself a focus of controversy (Box 1). In thesis that mirror neurons are for imitation [53], but it is consistent this review, we outline recent accounts of the correspon- with the generalist perspective. The environments in which humans dence problem, considering whether imitation can be develop might be such that they yield mirror neurons with greater understood within a general learning and motor control imitation-relevant specificity, and/or human imitation might repre- sent the interplay between mirror neurons and more complex Corresponding author: Brass, M. ([email protected]). cognitive abilities [26,51]. www.sciencedirect.com 1364-6613/$ - see front matter Q 2005 Elsevier Ltd. All rights reserved. doi:10.1016/j.tics.2005.08.007 ARTICLE IN PRESS TICS 384 2 Review TRENDS in Cognitive Sciences Vol.xx No.xx Monthxxxx shared representations of perception and action, how can Generalist theories assume that imitation is mediated we distinguish between internally generated and extern- by general learning and motor control mechanisms. There ally triggered motor representations. are two, complementary generalist theories under active development: ideomotor theory (IM) (Figure 1b), which subsumes imitation within a general account of motor Generalist and specialist theories of imitation control [5–7], and the associative sequence learning model Current theories of imitation offer specialist or generalist (ASL) (Figure 1c), which claims that the capacity to solutions to the correspondence problem (see Figure 1). imitate is a product of general processes of associative Specialist theories suggest that imitation is mediated by a learning [1,8]. IM assumes that all actions are represented special purpose mechanism. The most prominent special- in the form of ‘images’ of the sensory feedback they ist theory is the active intermodal matching (AIM) model produce, and that such representations are used to initiate [2,3] (Figure 1a). AIM proposes that, when a body move- and control body movements. From the perspective of ment is observed with the intention to imitate, the initial, IM observing somebody else executing an action leads to visual representation of the movement is converted into a an activation of an internal motor representation in the ‘supramodal’ representation which contains information about ‘organ relations’. AIM does not specify exactly how observer because the observed action is similar to the this information is encoded, but it implies that the code is content of the equivalent motor representation. This pre- the lingua franca of the perceptual-motor world. Supra- activated motor representation is then used to imitate the modal coding allows a visual representation of an observed observed behaviour. Hence, the ease with which humans action to be matched up with a pattern of motor activation imitate is due to the general organization of motor control that can produce the same action. Historically, discussion rather than a special purpose mechanism dedicated to of the AIM model has centred on the controversial ques- imitation. In the goal directed theory of imitation (GOADI) tion of whether newborn infants can imitate facial ges- Bekkering and colleagues have extended this approach by tures (for critical discussion of this issue see [1,4]). We do claiming that goals play a crucial role in the represen- not believe that further discussion of this issue here will tation and imitation of actions [9,10]. facilitate understanding of the correspondence problem. It ASL complements IM by explaining imitation, not only is preferable, in our view, to evaluate theories of imitation of ‘transparent’ actions such as finger movements, but also in terms of their capacity to account for a broad range of ‘opaque’ actions such as facial expressions, where the empirical findings. observer’s image of the model is not normally similar to (a) AIM (b) IM Sensory Motor Stimulus representation representation Motor program Visual perception of target adult facial acts Priming via similarity Supramodal of Priming representation acts via similarity Equivalence detector (c) ASL Horizontal Proprioceptive Sensory Sensory Sensory information 1 2 n Vertical Infant motor acts Motor1 Motor2 Motorn
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