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Hand to mouth: automatic imitation across effector systems.
The effector dependence of automatic imitation was investigated using a stimulus-response compatibility (SRC) procedure during which participants were required to make an open or closed response with their hand or their mouth. The correct response for each trial was indicated by a pair of letters in Experiments 1 and 2 and by a colored square in Experiment 3. Each of these imperative stimuli was accompanied by task-irrelevant action images depicting a hand or mouth opening or closing. In relation to the response, the irrelevant stimulus was movement compatible or movement incompatible, and effector compatible or effector incompatible. A movement compatibility effect was observed for both hand and mouth responses. These movement compatibility effects were present when the irrelevant stimulus was effector compatible and when it was effector incompatible, but were smaller when the irrelevant stimulus and response effectors were incompatible. Consistent with the associative sequence learning (ASL) model of imitation, these findings indicate that automatic imitation is partially effector dependent and therefore that the effector dependence of intentional imitation reflects, at least in part, the nature of the mechanisms that mediate visuomotor translation for imitation.
Action preparation helps and hinders perception of action.
Several theories of the mechanisms linking perception and action require that the links are bidirectional, but there is a lack of consensus on the effects that action has on perception. We investigated this by measuring visual event-related brain potentials to observed hand actions while participants prepared responses that were spatially compatible (e.g., both were on the left side of the body) or incompatible and action type compatible (e.g., both were finger taps) or incompatible, with observed actions. An early enhanced processing of spatially compatible stimuli was observed, which is likely due to spatial attention. This was followed by an attenuation of processing for both spatially and action type compatible stimuli, likely to be driven by efference copy signals that attenuate processing of predicted sensory consequences of actions. Attenuation was not response-modality specific; it was found for manual stimuli when participants prepared manual and vocal responses, in line with the hypothesis that action control is hierarchically organized. These results indicate that spatial attention and forward model prediction mechanisms have opposite, but temporally distinct, effects on perception. This hypothesis can explain the inconsistency of recent findings on action-perception links and thereby supports the view that sensorimotor links are bidirectional. Such effects of action on perception are likely to be crucial, not only for the control of our own actions but also in sociocultural interaction, allowing us to predict the reactions of others to our own actions.
The discrimination of natural movement by budgerigars (Melopsittacus undulates) and pigeons (Columba livia).
Three experiments examined the ability of birds to discriminate between the actions of walking forwards and backwards as demonstrated by video clips of a human walking a dog. Experiment 1 revealed that budgerigars (Melopsittacus undulates) could discriminate between these actions when the demonstrators moved consistently from left to right. Test trials then revealed that the discrimination transferred, without additional training, to clips of the demonstrators moving from right to left. Experiment 2 replicated the findings from Experiment 1 except that the demonstrators walked as if on a treadmill in the center of the display screen. The results from the first 2 experiments were replicated with pigeons in Experiment 3. The results cannot be explained if it is assumed that animals rely on static cues, such as those derived from individual postures, in order to discriminate between the actions of another animal. Instead, this type of discrimination appears to be controlled by dynamic cues derived from changes in the posture of the demonstrators.
General processes, rather than "goals," explain imitation errors.
The goal-directed theory of imitation (GOADI) states that copying of action outcomes (e.g., turning a light switch) takes priority over imitation of the means by which those outcomes are achieved (e.g., choice of effector or grip). The object < effector < grip error pattern in the pen-and-cups task provides strong support for GOADI. Experiment 1 replicated this effect using video stimuli. Experiment 2 showed that shifting the color cue from objects to effectors makes imitation of effector selection more accurate than imitation of object and grip selection. Experiment 3 replicated this result when participants were required to describe actions. Experiment 4 indicated that, when participants are imitating and describing actions, enhancing grip discriminability makes grip selection the most accurately executed component of the task. Consistent with theories that hypothesize that imitation relies on task-general mechanisms (e.g., the associative sequence learning model, ideomotor theory), these findings suggest that imitation is no more or less goal directed than other tasks involving action observation.
Bottom-up, not top-down, modulation of imitation by human and robotic models.
Visual observation of human actions provokes more motor activation than observation of robotic actions. We investigated the extent to which this visuomotor priming effect is mediated by bottom-up or top-down processing. The bottom-up hypothesis suggests that robotic movements are less effective in activating the 'mirror system' via pathways from visual areas via the superior temporal sulcus to parietal and premotor cortices. The top-down hypothesis postulates that beliefs about the animacy of a movement stimulus modulate mirror system activity via descending pathways from areas such as the temporal pole and prefrontal cortex. In an automatic imitation task, subjects performed a prespecified movement (e.g. hand opening) on presentation of a human or robotic hand making a compatible (opening) or incompatible (closing) movement. The speed of responding on compatible trials, compared with incompatible trials, indexed visuomotor priming. In the first experiment, robotic stimuli were constructed by adding a metal and wire 'wrist' to a human hand. Questionnaire data indicated that subjects believed these movements to be less animate than those of the human stimuli but the visuomotor priming effects of the human and robotic stimuli did not differ. In the second experiment, when the robotic stimuli were more angular and symmetrical than the human stimuli, human movements elicited more visuomotor priming than the robotic movements. However, the subjects' beliefs about the animacy of the stimuli did not affect their performance. These results suggest that bottom-up processing is primarily responsible for the visuomotor priming advantage of human stimuli.
Imitation: is cognitive neuroscience solving the correspondence problem?
Imitation poses a unique problem: how does the imitator know what pattern of motor activation will make their action look like that of the model? Specialist theories suggest that this correspondence problem has a unique solution; there are functional and neurological mechanisms dedicated to controlling imitation. Generalist theories propose that the problem is solved by general mechanisms of associative learning and action control. Recent research in cognitive neuroscience, stimulated by the discovery of mirror neurons, supports generalist solutions. Imitation is based on the automatic activation of motor representations by movement observation. These externally triggered motor representations are then used to reproduce the observed behaviour. This imitative capacity depends on learned perceptual-motor links. Finally, mechanisms distinguishing self from other are implicated in the inhibition of imitative behaviour.
Four routes of cognitive evolution.
Four routes of cognitive evolution are distinguished: phylogenetic construction, in which natural selection produces qualitative change to the way a cognitive mechanism operates (language); phylogenetic inflection, in which natural selection biases the input to a cognitive mechanism (imprinting and spatial memory); ontogenetic construction, in which developmental selection alters the way a cognitive mechanism operates (face recognition and theory of mind); and ontogenetic inflection, in which developmental selection changes the input to a cognitive mechanism (imitation). This framework integrates findings from evolutionary psychology (i.e., all research on the evolution of mentality and behavior). In contrast with human nativist evolutionary psychology, it recognizes the adaptive significance of developmental processes, conserves the distinction between cognitive and noncognitive mechanisms, and encompasses research on human and nonhuman animals.
Imitation of lateralised body movements: doing it the hard way.
Two experiments examined imitation of lateralised body movement sequences presented at six viewing angles (0 degrees , 60 degrees , 120 degrees , 180 degrees , 240 degrees , and 300 degrees rotation relative to the participant's body). Experiment 1 found that, when participants were instructed simply to "do what the model does", at all viewing angles they produced more actions using the same side of the body as the model (anatomical matches), than actions using the opposite side (anatomical non-matches). In Experiment 2 participants were instructed to produce either anatomical matches or anatomical non-matches of observed actions. When the model was viewed from behind (0 degrees ), the anatomically matching group were more accurate than the anatomically non-matching group, but the non-matching group was superior when the model faced the participant (180 degrees and 240 degrees ). No reliable differences were observed between groups at 60 degrees , 120 degrees , and 300 degrees . In combination, the results of Experiments 1 and 2 suggest that, when they are confronting a model, people choose to imitate the hard way; they attempt to match observed actions anatomically, in spite of the fact that anatomical matching is more subject to error than anatomical non-matching.
Through the looking glass: counter-mirror activation following incompatible sensorimotor learning.
The mirror system, comprising cortical areas that allow the actions of others to be represented in the observer's own motor system, is thought to be crucial for the development of social cognition in humans. Despite the importance of the human mirror system, little is known about its origins. We investigated the role of sensorimotor experience in the development of the mirror system. Functional magnetic resonance imaging was used to measure neural responses to observed hand and foot actions following one of two types of training. During training, participants in the Compatible (control) group made mirror responses to observed actions (hand responses were made to hand stimuli and foot responses to foot stimuli), whereas the Incompatible group made counter-mirror responses (hand to foot and foot to hand). Comparison of these groups revealed that, after training to respond in a counter-mirror fashion, the relative action observation properties of the mirror system were reversed; areas that showed greater responses to observation of hand actions in the Compatible group responded more strongly to observation of foot actions in the Incompatible group. These results suggest that, rather than being innate or the product of unimodal visual or motor experience, the mirror properties of the mirror system are acquired through sensorimotor learning.
Infants' behavioral reenactment of "failed attempts": exploring the roles of emulation learning, stimulus enhancement, and understanding of intentions.
Two studies were conducted to examine whether infants' reenactment of intended but unconsummated acts in A. N. Meltzoff's (1995) failed-attempt paradigm is due to reading the adult's underlying intention or to the effects of nonimitative social learning processes. Two novel conditions that emphasized the object affordances and the spatial contiguity of the object sets were devised. When infants' first actions only were counted, infants who observed the full-demonstration model produced more target acts. When all target acts produced within the 20-s response period were counted, infants in the emulation-learning and spatial contiguity conditions produced as many target acts as infants in the full-demonstration and failed-attempt conditions. This pattern of findings suggests that nonimitative social learning processes may influence infants' response in the behavioral reenactment paradigm.
Acquisition of automatic imitation is sensitive to sensorimotor contingency.
The associative sequence learning model proposes that the development of the mirror system depends on the same mechanisms of associative learning that mediate Pavlovian and instrumental conditioning. To test this model, two experiments used the reduction of automatic imitation through incompatible sensorimotor training to assess whether mirror system plasticity is sensitive to contingency (i.e., the extent to which activation of one representation predicts activation of another). In Experiment 1, residual automatic imitation was measured following incompatible training in which the action stimulus was a perfect predictor of the response (contingent) or not at all predictive of the response (noncontingent). A contingency effect was observed: There was less automatic imitation indicative of more learning in the contingent group. Experiment 2 replicated this contingency effect and showed that, as predicted by associative learning theory, it can be abolished by signaling trials in which the response occurs in the absence of an action stimulus. These findings support the view that mirror system development depends on associative learning and indicate that this learning is not purely Hebbian. If this is correct, associative learning theory could be used to explain, predict, and intervene in mirror system development.
Blind imitation in pigeons, Columba livia
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. © 2006 The Association for the Study of Animal Behaviour.
Sequence learning by action, observation and action observation
The serial reaction time (SRT) task was used to compare learning of a complex sequence by action (participants responded to sequential stimuli), by observation (participants watched but did not respond to sequential stimuli), and by action-observation (participants watched an expert model responding to sequential stimuli). Each of these groups was compared with an untrained control group. Experiment 1 indicated that both observation and action-observation were sufficient to support learning of a 12-item second-order conditional (SOC) sequence. Experiment 2 confirmed these findings, and showed that, as indexed by reaction time (RT), the extent of learning by observation and by action-observation was comparable to that of action-based learning. Using a recognition test, Experiment 2 and 3 also provided evidence that, whereas learning by stimulus observation was explicit, learning by actionobservation was implicit. These findings are consistent with a connection between motor systems and implicit learning, but do not support the hypothesis that overt action is necessary for implicit learning. © 2005 The British Psychological Society.
Action observation supports effector-dependent learning of finger movement sequences.
Practising a motor skill can result in effector-dependent learning (learning that does not transfer from the set of muscles used in training to a new set of muscles). Proceeding from neurophysiological evidence of motor activation during action observation, this study asked whether observational learning, learning through observation of skilled performance, can also be effector-dependent. Adult human participants observed a model's right hand as the model responded to an eight-item sequence in a serial reaction time (SRT) task. Their sequence learning was then compared in two tests with that of controls who had observed the model's right hand responding to random targets during training. All participants performed the SRT task with their right hand in the first test and with their left hand in the second. Evidence of observational learning was obtained in the right hand test but not in the left hand test. This implies that sequence learning based on observation of right hand performance did not transfer to the left hand, and therefore that observational learning can support effector-dependent learning of finger movement sequences. A second experiment used the same procedure to assess learning by a group of participants who observed a sequence of response locations only. This group did not observe the model's responses. Results suggested that action observation was necessary for the effector-dependent observational learning demonstrated in Experiment 1.
Sequence learning by action, observation and action observation.
The serial reaction time (SRT) task was used to compare learning of a complex sequence by action (participants responded to sequential stimuli), by observation (participants watched but did not respond to sequential stimuli), and by action-observation (participants watched an expert model responding to sequential stimuli). Each of these groups was compared with an untrained control group. Experiment 1 indicated that both observation and action-observation were sufficient to support learning of a 12-item second-order conditional (SOC) sequence. Experiment 2 confirmed these findings, and showed that, as indexed by reaction time (RT), the extent of learning by observation and by action-observation was comparable to that of action-based learning. Using a recognition test, Experiment 2 and 3 also provided evidence that, whereas learning by stimulus observation was explicit, learning by action-observation was implicit. These findings are consistent with a connection between motor systems and implicit learning, but do not support the hypothesis that overt action is necessary for implicit learning.
Spatial S-R compatibility effects in an intentional imitation task.
The active intermodal mapping hypothesis suggests that intentional imitation is mediated by a highly efficient, special-purpose mechanism of actor-centered movement encoding. In the present study, using methods from stimulus-response (S-R) compatibility research, we found no evidence to support this hypothesis. In two experiments, the performance of adult participants instructed to imitate actor-centered spatial properties of head, arm, and leg movements was affected by task-irrelevant, egocentric spatial cues. In Experiment 1, participants imitated using the same side of their bodies as did the model, and performance was less accurate when egocentric stimulus location was response incompatible than when it was response compatible. This effect was reversed in Experiment 2 when participants imitated using the opposite side of their bodies. These findings, in line with general process theories of imitation, imply that intentional imitation is mediated by the same processes that mediate responding to inanimate stimuli on the basis of arbitrary S-R mappings.
Mesmerising mirror neurons.
Mirror neurons have been hailed as the key to understanding social cognition. I argue that three currents of thought-relating to evolution, atomism and telepathy-have magnified the perceived importance of mirror neurons. When they are understood to be a product of associative learning, rather than an adaptation for social cognition, mirror neurons are no longer mesmerising, but they continue to raise important questions about both the psychology of science and the neural bases of social cognition.
Automatic imitation of intransitive actions.
Previous research has indicated a potential discontinuity between monkey and human ventral premotor-parietal mirror systems, namely that monkey mirror systems process only transitive (object-directed) actions, whereas human mirror systems may also process intransitive (non-object-directed) actions. The present study investigated this discontinuity by seeking evidence of automatic imitation of intransitive actions--hand opening and closing--in humans using a simple reaction time (RT), stimulus-response compatibility paradigm. Left-right and up-down spatial compatibility were controlled by ensuring that stimuli were presented and responses executed in orthogonal planes, and automatic imitation was isolated from simple and complex orthogonal spatial compatibility by varying the anatomical identity of the stimulus hand and response hemispace, respectively. In all conditions, action compatible responding was faster than action incompatible responding, and no effects of spatial compatibility were observed. This experiment therefore provides evidence of automatic imitation of intransitive actions, and support for the hypothesis that human and monkey mirror systems differ with respect to the processing of intransitive actions.
Stimulus-driven selection of routes to imitation.
Several models have proposed that an action can be imitated via one of two routes: a direct visuospatial route, which can in principle mediate imitation of both meaningful (MF) and meaningless (ML) actions, and an indirect semantic route, which can be used only for MF actions. The present study investigated whether selection between the direct and indirect routes is strategic or stimulus driven. Tessari and Rumiati (J Exp Psychol Hum Percept Perform 30:1107-1116, 2004) have previously shown, using accuracy measures, that imitation of MF actions is superior to imitation of ML actions when the two action types are presented in separate blocks, and that the advantage of MF over ML items is smaller or absent when they are presented in mixed blocks. We first replicated this finding using an automated reaction time (RT), as well as accuracy, measure. We then examined imitation of MF and ML actions in the mixed condition as a function of the action type presented in the previous trial and in relation to the number of previous test trials. These analyses showed that (1) for both action types, performance was worse immediately after ML than MF trials, and (2) even at the beginning of the mixed condition, responding to MF actions was no better than responding to ML items. These results suggest that the properties of the action stimulus play a substantial role in determining whether imitation is mediated by the direct or the indirect route, and that effects of block composition on imitation need not be generated through strategic switching between routes.