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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.
Are automatic imitation and spatial compatibility mediated by different processes?
Automatic imitation or "imitative compatibility" is thought to be mediated by the mirror neuron system and to be a laboratory model of the motor mimicry that occurs spontaneously in naturalistic social interaction. Imitative compatibility and spatial compatibility effects are known to depend on different stimulus dimensions-body movement topography and relative spatial position. However, it is not yet clear whether these two types of stimulus-response compatibility effect are mediated by the same or different cognitive processes. We present an interactive activation model of imitative and spatial compatibility, based on a dual-route architecture, which substantiates the view they are mediated by processes of the same kind. The model, which is in many ways a standard application of the interactive activation approach, simulates all key results of a recent study by Catmur and Heyes (2011). Specifically, it captures the difference in the relative size of imitative and spatial compatibility effects; the lack of interaction when the imperative and irrelevant stimuli are presented simultaneously; the relative speed of responses in a quintile analysis when the imperative and irrelevant stimuli are presented simultaneously; and the different time courses of the compatibility effects when the imperative and irrelevant stimuli are presented asynchronously.
What's social about social learning?
Research on social learning in animals has revealed a rich variety of cases where animals--from caddis fly larvae to chimpanzees--acquire biologically important information by observing the actions of others. A great deal is known about the adaptive functions of social learning, but very little about the cognitive mechanisms that make it possible. Even in the case of imitation, a type of social learning studied in both comparative psychology and cognitive science, there has been minimal contact between the two disciplines. Social learning has been isolated from cognitive science by two longstanding assumptions: that it depends on a set of special-purpose modules--cognitive adaptations for social living; and that these learning mechanisms are largely distinct from the processes mediating human social cognition. Recent research challenges these assumptions by showing that social learning covaries with asocial learning; occurs in solitary animals; and exhibits the same features in diverse species, including humans. Drawing on this evidence, I argue that social and asocial learning depend on the same basic learning mechanisms; these are adapted for the detection of predictive relationships in all natural domains; and they are associative mechanisms--processes that encode information for long-term storage by forging excitatory and inhibitory links between event representations. Thus, human and nonhuman social learning are continuous, and social learning is adaptively specialized--it becomes distinctively "social"--only when input mechanisms (perceptual, attentional, and motivational processes) are phylogenetically or ontogenetically tuned to other agents.
Automatic imitation.
"Automatic imitation" is a type of stimulus-response compatibility effect in which the topographical features of task-irrelevant action stimuli facilitate similar, and interfere with dissimilar, responses. This article reviews behavioral, neurophysiological, and neuroimaging research on automatic imitation, asking in what sense it is "automatic" and whether it is "imitation." This body of research reveals that automatic imitation is a covert form of imitation, distinct from spatial compatibility. It also indicates that, although automatic imitation is subject to input modulation by attentional processes, and output modulation by inhibitory processes, it is mediated by learned, long-term sensorimotor associations that cannot be altered directly by intentional processes. Automatic imitation provides an important tool for the investigation of the mirror neuron system, motor mimicry, and complex forms of imitation. It is a new behavioral phenomenon, comparable with the Stroop and Simon effects, providing strong evidence that even healthy adult humans are prone, in an unwilled and unreasoned way, to copy the actions of others.
Imitative pecking by budgerigars, Melopsittacus undulatus, over a 24 h delay
On video, budgerigars observed a conspecific demonstrator depressing a stopper by pecking or by stepping and then feeding from the box below. The observers were given access to the stopper, immediately after observation or following a 24 h delay, and we recorded the proportion of their stopper removal responses that were made by pecking and by stepping. In experiments 1a and 1b, observers of pecking made a greater proportion of pecking responses than observers of stepping, and this effect did not vary between the immediate and delayed test groups. The results of experiment 2 replicated this effect with a delayed test, and suggested that it was due to imitation of pecking. Control birds that observed a demonstrator feeding, but did not see stopper removal, made a smaller proportion of pecking responses than pecking observers, but their behaviour did not differ from that of stepping observers. These findings are consistent with the associative sequence learning model of imitation, which suggests that the capacity to imitate a particular action depends on correlated experience of observing and executing that action. © 2009 The Association for the Study of Animal Behaviour.
The interaction between response effects during the acquisition of response priming.
In three experiments we investigated the role of associative learning in the acquisition of response priming by effect stimuli, by examining their interaction during response-effect learning. Having replicated the ability of visual effect stimuli to prime their associated responses, we paired a response with a compound consisting of visual and auditory effects before assessing the ability of the auditory effect stimulus to prime the response. This priming was reduced if the visual stimulus had been pre-trained as an effect of the response. By contrast, priming by the visual effect stimulus was potentiated when the auditory effect had been pre-trained. We interpret these interactions in terms of contemporary associative learning theory derived from studies of conditioning.
Stimulus Enhancement: Controls for Social Facilitation and Local Enhancement
In two experiments, magazine-trained observer rats were given access to a pair of levers and allowed to make 50 reinforced responses immediately after the left or the right lever had been pressed 100 times for food reward by a conspecific demonstrator. In Experiment 1, where observers were rewarded for responses on either lever, those that had been able to see their demonstrators responding showed a response bias in favor of the demonstrator's lever, while rats that had been separated from their demonstrators by a metal screen did not. In Experiment 2, each observer saw a "viewed demonstrator" responding on one lever and was tested, with reinforcement for responses on the viewed demonstrator's lever or on the other manipulandum in a second chamber where a "box demonstrator" had just finished responding. Observers rewarded for responding on the same lever as the viewed demonstrator (Group SAME) showed better discrimination at the beginning of the test session than observers rewarded for responses on the other lever (Group DIFFERENT). Within Group SAME, rats whose box demonstrator had responded on the reinforced lever showed better discrimination than those whose box demonstrator responded on the nonreinforced lever, but the reverse was true in Group DIFFERENT. Both experiments provide evidence of stimulus enhancement unconfounded by social facilitation and local enhancement, and the latter suggests that demonstrator-deposited scent cues can facilitate discrimination via local enhancement or on the basis of their sensory properties alone. © 2000 Academic Press.
Time course analyses confirm independence of imitative and spatial compatibility.
Imitative compatibility, or automatic imitation, has been used as a measure of imitative performance and as a behavioral index of the functioning of the human mirror system (e.g., Brass, Bekkering, Wohlschlager, & Prinz, 2000; Heyes, Bird, Johnson, & Haggard, 2005; Kilner, Paulignan, & Blakemore, 2003). However, the use of imitative compatibility as a measure of imitation has been criticized on the grounds that imitative compatibility has been confounded with simple spatial compatibility (Aicken, Wilson, Williams, & Mon-Williams, 2007; Bertenthal, Longo, & Kosobud, 2006; Jansson, Wilson, Williams, & Mon-Williams, 2007). Two experiments are reported in which, in contrast with previous studies, imitative compatibility was measured on both spatially compatible and spatially incompatible trials, and imitative compatibility was shown to be present regardless of spatial compatibility. Additional features of the experiments allowed measurement of the time courses of the imitative and spatial compatibility effects both within and across trials. It was found that imitative compatibility follows a different time course from spatial compatibility, providing further evidence for their independence and supporting the use of imitative compatibility as a measure of imitation.
Effector-dependent learning by observation of a finger movement sequence.
Can observational learning be effector dependent? In 3 experiments, observers watched a model respond to a 6-item unique sequence in a serial reaction time task. Their sequence knowledge was then compared with that of controls who had performed an unrelated task or observed a model responding to random targets. Observational learning was indicated when the introduction of a new sequence was associated with more reaction time elevation in observers than in controls. The authors found evidence of observational learning only when observers used the finger movement sequence that they observed during training, not when they responded at the same sequence of locations using different digits. Free generation and recognition tests also detected observational learning. These results imply that observational learning can be both explicit and effector dependent.
Motor learning by observation: evidence from a serial reaction time task.
This study sought evidence of observational motor learning, a type of learning in which observation of the skilled performance of another person not only facilitates motor skill acquisition but does so by contributing to the formation of effector-specific motor representations. Previous research has indicated that observation of skilled performance engages cognitive processes similar to those occurring during action execution or physical practice, but has not demonstrated that these include processes involved in effector-specific representation. In two experiments, observer subjects watched the experimenter performing a serial reaction time (SRT) task with a six-item unique sequence before sequence knowledge was assessed by response time and/or free generation measures. The results suggest that: (1) subjects can acquire sequence information by watching another person performing the task (Experiments 1-2); (2) observation results in as much sequence learning as task practice when learning is measured by reaction times (RTs) and more than task practice when sequence learning is measured by free generation performance (Experiment 2, Part 1); and (3) sequence knowledge acquired by model observation can be encoded motorically--that is, in an effector-specific fashion (Experiment 2, Part 2).
Where do mirror neurons come from?
Debates about the evolution of the 'mirror neuron system' imply that it is an adaptation for action understanding. Alternatively, mirror neurons may be a byproduct of associative learning. Here I argue that the adaptation and associative hypotheses both offer plausible accounts of the origin of mirror neurons, but the associative hypothesis has three advantages. First, it provides a straightforward, testable explanation for the differences between monkeys and humans that have led some researchers to question the existence of a mirror neuron system. Second, it is consistent with emerging evidence that mirror neurons contribute to a range of social cognitive functions, but do not play a dominant, specialised role in action understanding. Finally, the associative hypothesis is supported by recent data showing that, even in adulthood, the mirror neuron system can be transformed by sensorimotor learning. The associative account implies that mirror neurons come from sensorimotor experience, and that much of this experience is obtained through interaction with others. Therefore, if the associative account is correct, the mirror neuron system is a product, as well as a process, of social interaction.