Imitative response tendencies following observation of intransitive actions

Clear and unequivocal evidence shows that observation of object affordances or transitive actions facilitates the activation of a compatible response. By contrast, the evidence showing response facilitation following observation of intransitive actions is less conclusive because automatic imitation and spatial compatibility have been confounded. Three experiments tested whether observation of a finger movement (i.e., an intransitive action) in a choice reaction-time task facilitates the corresponding finger movement response because of imitation, a common spatial code, or some combination of both factors. The priming effects of a spatial and an imitative stimulus were tested in combination (Experiment 1), in opposition (Experiment 2), and independently (Experiment 3). Contrary to previous findings, the evidence revealed significant contributions from both automatic imitation and spatial compatibility, but the priming effects from an automatic tendency to imitate declined significantly across a block of trials whereas the effects of spatial compatibility remained constant or increased slightly. These differential effects suggest that priming associated with automatic imitation is mediated by a different regime than priming associated with spatial compatibility.     

Is automatic imitation a specialized form of stimulus–response compatibility? Dissociating imitative and spatial compatibilities

In recent years research on automatic imitation has received considerable attention because it represents an experimental platform for investigating a number of interrelated theories suggesting that the perception of action automatically activates corresponding motor programs. A key debate within this research centers on whether automatic imitation is any different than other long-term S-R associations, such as spatial stimulus-response compatibility. One approach to resolving this issue is to examine whether automatic imitation shows similar response characteristics as other classes of stimulus-response compatibility. This hypothesis was tested by comparing imitative and spatial compatibility effects with a two alternative forced-choice stimulus-response compatibility paradigm. The stimulus on each trial was a left or right hand with either the index or middle finger tapping down. Speeded responses were performed with the index or middle finger of the right hand in response to the identity or the left-right spatial position of the stimulus finger. Two different tasks were administered: one that involved responding to the stimulus (S-R) and one that involved responding to the opposite stimulus (OS-R; i.e., the one not presented on that trial). Based on previous research and a connectionist model, we predicted standard compatibility effects for both spatial and imitative compatibility in the S-R task, and a reverse compatibility effect for spatial compatibility, but not for imitative compatibility, in the OS-R task. The results from the mean response times, mean percentage of errors, and response time distributions all converged to support these predictions. A second noteworthy result was that the recoding of the finger identity in the OS-R task required significantly more time than the recoding of the left-right spatial position, but the encoding time for the two stimuli in the S-R task was equivalent. In sum, this evidence suggests that the processing of spatial and imitative compatibility is dissociable with regard to two different processes in dual processing models of stimulus-response compatibility.     

Compatibility between observed and executed finger movements: comparing symbolic, spatial, and imitative cues

Intuitively, one can assume that imitating a movement is an easier task than responding to a symbolic stimulus like a verbal instruction. Support for this suggestion can be found in neuropsychological research as well as in research on stimulus-response compatibility. However controlled experimental evidence for this assumption is still lacking. We used a stimulus-response compatibility paradigm to test the assumption. In a series of experiments, it was tested whether observed finger movements have a stronger influence on finger movement execution than a symbolic or spatial cue. In the first experiment, we compared symbolic cues with observed finger movements using an interference paradigm. Observing finger movements strongly influenced movement execution, irrespective of whether the finger movement was the relevant or the irrelevant stimulus dimension. In the second experiment, effects of observed finger movements and spatial finger cues were compared. The observed finger movement dominated the spatial finger cue. A reduction in the similarity of observed and executed action in the third experiment led to a decrease of the influence of observed finger movement, which demonstrates the crucial role of the imitative relation of observed and executed action for the described effects. The results are discussed in relation to recent models of stimulus-response compatibility. Neurocognitive support for the strong relationship between movement observation and movement execution is reported.     

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.     

Differential dynamics of spatial and non-spatial stimulus-response compatibility effects: a dual task LRP study

Choice reaction times are shorter when stimulus and response features are compatible rather than incompatible. Recent studies revealed that spatial compatibility effects in Simon tasks are strongly attenuated when there is temporal overlap with a different high-priority task. In contrast, non-spatial variants of the Simon task appear to be unaffected by task overlap. The present study used the lateralized readiness potential (LRP) within a dual task design to elucidate the dynamics underlying these differential effects for a color and a spatial variant of the Simon task. In the color version there was no sign of early response priming by irrelevant stimulus features in the LRP. The color compatibility effect was independent of task overlap and reflected in the LRP onset latency. In contrast, in the spatial version, priming by irrelevant stimulus location showed up and was mirrored by early LRP activation. Response priming and the corresponding Simon effect, however, were present only in case of little temporal overlap with the primary task. The absence of spatial compatibility effects at strong temporal overlap suggests that response conflicts due to stimulus-related priming depend on the availability of processing resources.     

The influence of dimensional overlap on location-related priming in the Simon task

Choice reaction times are shorter when stimulus and response locations are compatible than when they are incompatible as in the Simon effect. Recent studies revealed that Simon effects are strongly attenuated when there is temporal overlap with a different high-priority task, accompanied by a decrease of early location-related response priming as reflected in the lateralized readiness potential (LRP). The latter result was obtained in a study excluding overlap of stimulus location with any other dimension in the tasks. Independent evidence suggests that location-related priming might be present in conditions with dimensional overlap. Here we tested this prediction in a dual-task experiment supplemented with recording LRPs. The secondary task was either a standard Simon task where irrelevant stimulus location overlapped with dimensions of the primary task or a Stroop-like Simon task including additional overlap of irrelevant and relevant stimulus attributes. At high temporal overlap, there was no Simon effect nor was there stimulus-related response priming in either condition. Therefore stimulus-triggered response priming seems to be abolished in conditions of limited capacity even if the likelihood of an S–R compatibility effect is maximized.     

Probing Representations of Gymnastics Movements: A Visual Priming Study

In this study, we designed a visual short‐term priming paradigm to investigate the mechanisms underlying the priming of movements and to probe movement representations in motor experts and matched controls. We employed static visual stimuli that implied or not human whole‐body movements, that is, gymnastics movements and static positions. Twelve elite female gymnasts and twelve matched controls performed a speeded two‐choice response time task. The participants were presented with congruent and incongruent prime‐target pairs and had to decide whether the target stimulus represented a gymnastics movement or a static position. First, a visual priming effect was observed in the two groups. Second, a stimulus–response rote association could not easily account for our results. Novel primes never presented as targets could also prime the targets. Third, by manipulating three levels of prime‐target relations in moving congruent pairs, we demonstrated that the more similar prime‐target pairs, the greater the facilitation in target. Lastly, gymnastics motor expertise impacted on priming effects.     

Learning associations between action and perception: effects of incompatible training on body part and spatial priming

Observation of another person executing an action primes the same action in the observer's motor system. Recent evidence has shown that these priming effects are flexible, where training of new associations, such as making a foot response when viewing a moving hand, can reduce standard action priming effects (Gillmeister, Catmur, Liepelt, Brass, & Heyes, 2008). Previously, these effects were obtained after explicit learning tasks in which the trained action was cued by the content of a visual stimulus. Here we report similar learning processes in an implicit task in which the participant's action is self-selected, and subsequent visual effects are determined by the nature of that action. Importantly, we show that these learning processes are specific to associations between actions and viewed body parts, in that incompatible spatial training did not influence body part or spatial priming effects. Our results are consistent with models of visuomotor learning that place particular emphasis on the repeated experience of watching oneself perform an action.     

Impact of Parkinson’s disease and dopaminergic medication on adaptation to explicit and implicit visuomotor perturbations

The capacity to learn new visuomotor associations is fundamental to adaptive motor behavior. Evidence suggests visuomotor learning deficits in Parkinson’s disease (PD). However, the exact nature of these deficits and the ability of dopamine medication to improve them are under-explored. Previous studies suggested that learning driven by large and small movement errors engaged distinct neural mechanisms. Here, we investigated whether PD patients have a generalized impairment in visuomotor learning or selective deficits in learning from large explicit errors which engages cognitive strategies or small imperceptible movement errors involving primarily implicit learning processes. Visuomotor learning skills of non-medicated and medicated patients were assessed in two reaching tasks in which the size of visuospatial errors experienced during learning was manipulated using a novel three-dimensional virtual reality environment. In the explicit perturbation task, the visuomotor perturbation was applied suddenly resulting in large consciously detected initial spatial errors, whereas in the implicit perturbation task, the perturbation was gradually introduced in small undetectable steps such that subjects never experienced large movement errors. A major finding of this study was that PD patients in non-medicated and medicated conditions displayed slower learning rates and smaller adaptation magnitudes than healthy subjects in the explicit perturbation task, but performance similar to healthy controls in the implicit perturbation task. Also, non-medicated patients showed an average reduced deadaptation relative to healthy controls when exposed to the large errors produced by the sudden removal of the perturbation in both the explicit and implicit perturbation tasks. Although dopaminergic medication consistently improved motor signs, it produced a variable impact on learning the explicit perturbation and deadaptation and unexpectedly worsened performance in some patients. Considered together, these results indicate that PD selectively impairs the ability to learn from large consciously detected visuospatial errors. This finding suggests that basal ganglia-related circuits are important neural structures for adaptation to sudden perturbations requiring awareness and high-cost action selection. Dopaminergic treatment may selectively compromise the ability to learn from large explicit movement errors for reasons that remain to be elucidated.     

Visual objects can potentiate a grasping neural simulation which interferes with manual response execution

Previous studies on visuomotor priming have provided insufficient information to determine whether the reach-to-grasp potentiation of a non-target object produces a specific effect during response execution. In order to answer this question, subjects were instructed to reach and grasp a response device with either a power or a precision grip, depending on whether the stimulus they saw was empty or full. Stimuli consisted of containers (graspable with either a power or a precision grip), with non-graspable stimuli added as a control condition (geometrical shapes). The image of the non-target object was removed during the execution phase. Results demonstrate slower execution responses related to motor incompatibility, though conversely, no faster responses with motor compatibility. Moreover, any visuomotor priming effect required that the container be displayed during response execution. These data suggest that during response execution, motor incompatibility produces a disruptive effect likely due to competition between two cerebral events: motor control of the actual response execution and visual object reach-to-grasp neural simulation.     

Exploring visuomotor priming following biological and non-biological stimuli

Observation of human actions influences the observer’s own motor system, termed visuomotor priming, and is believed to be caused by automatic activation of mirror neurons. Evidence suggests that priming effects are larger for biological (human) as opposed to non-biological (object) stimuli and enhanced when viewing stimuli in mirror compared to anatomical orientation. However, there is conflicting evidence concerning the extent of differences between biological and non-biological stimuli, which may be due to stimulus related confounds. Over three experiments, we compared how visuomotor priming for biological and non-biological stimuli was affected over views, over time and when attention to the moving stimulus was manipulated. The results indicated that the strength of priming for the two stimulus types was dependent on attentional location and load. This highlights that visuomotor priming is not an automatic process and provides a possible explanation for conflicting evidence regarding the differential effects of biological and non-biological stimuli.     

Motor priming by movement observation with contralateral concurrent action execution

In the present study, the influence of simultaneous action execution on motor priming was investigated during movement observation using a simple-reaction task. Although previous studies have reported various effects of priming on motor performance, it has not yet been clarified how an additional source conveying kinetic information would modulate the priming effects. In the experiment, participants were asked to respond to an auditory cue by flexing their wrist while observing a line movement, which was slowly swinging like an inverted pendulum. In addition to the observation of line movement, the participants executed wrist flexion-extension actions synchronizing with line movement. The hand involved in pre-response wrist action varied with the priming condition: no movement execution (observation only), contralateral hand, and ipsilateral hand. In the contralateral condition, the stimulus-response congruency of movement direction was conflicted depending on the frame of reference (visual vs. anatomical coordinates). We found that all three priming conditions produced the compatibility effect, and the effect size did not differ between them. Importantly, in the contralateral condition, participants responded faster when the direction of line movement was congruent with the response movement in the anatomical coordinates. That is, the reaction time was shorter when pre-response action execution was in the flexion phase, even though the direction of observed movement and the response action were incongruent from the participants’ view. These results suggest that kinetic information has a great contribution to the motor priming system, which can reverse the vision-based compatibility effect.     

The role of emotional context in facilitating imitative actions

In two experiments, we explored whether emotional context influences imitative action tendencies. To this end, we examined how emotional pictures, presented as primes, affect imitative tendencies using a compatibility paradigm. In Experiment 1, when seen index finger movements (lifting or tapping) and pre-instructed finger movements (tapping) were the same (tapping–tapping, compatible trials), participants were faster than when they were different (lifting–tapping, incompatible trials). This compatibility effect was enhanced when the seen finger movement was preceded by negative primes compared with positive or neutral primes. In Experiment 2, using only negative and neutral primes, the influence of negative primes on the compatibility effect was replicated with participants performing two types of pre-instructed finger movements (tapping and lifting). This emotional modulation of the compatibility effect was independent of the participants' trait anxiety level. Moreover, the emotional modulation pertained primarily to the compatible conditions, suggesting facilitated imitation due to negatively valent primes rather than increased interference. We speculate that negative stimuli increase imitative tendencies as a natural response in potential flight-or-fight situations.     

Inverse cue priming is not limited to masks with relevant features

Apart from positive priming effects, masked prime stimuli can impair responses to a subsequent target stimulus which shares response-critical features in contrast to a target assigned to the opposite response. This counterintuitive phenomenon is called inverse priming (or negative compatibility effect). Here we examine the generality of this phenomenon beyond priming of motor responses. We used a non-motor cue-priming paradigm to study the underlying mechanism of inverse priming for relevant features masks which include task-relevant stimulus features and for irrelevant masks which omit task-relevant features. We found inverse cue-priming effects with both types of masks. With task-irrelevant masks inverse cue-priming was emphasized in those participants being unable to perceive the prime. The existence of inverse non-motor priming under conditions where simple perceptual interactions between the stimuli are ruled out as the source of inverse priming is at odds with the view that inverse priming reflects motor inhibition. Alternatives are discussed.     

Does a tool eliminate spatial compatibility effects?

Responding to a stimulus is faster and more accurate when stimulus location and response location spatially correspond than when they do not correspond (stimulus–response compatibility). In five experiments this standard compatibility effect is examined when using a T-shaped lever as a tool. Handling the lever allowed distinguishing body-related action effects (e.g., the tactile feedback from the moving finger) from external action effects (e.g., reaching at the stimulus with the lever's end-point). Results showed that the spatial relationship between stimulus and the direction of the hand movement (S-R compatibility) as well as the relationship between the stimulus and the functional end-points of the tool (S-E compatibility) determine performance. More precisely, responses were fast and less error prone when both kinds of compatibility did correspond than when they did not correspond.     

Approach and avoidance movements are unaffected by cognitive conflict: A comparison of the Simon effect and stimulus–response compatibility

Participants in this study reached from central fixation to a lateral position that either contained or was opposite to the stimulus. Cognitive conflict was induced when the stimulus and response directions did not correspond. In the Simon task, the response direction was cued by the color of the lateral stimulus, and corresponding and noncorresponding trials varied randomly in the same block of trials, resulting in high uncertainty and long reaction times (RTs). In the stimulus-response compatibility (SRC) task, participants reached toward or away from the stimulus in separate blocks of trials, resulting in low uncertainty and short RTs. In the SRC task, cognitive conflict in noncorresponding trials slowed down RTs but hardly affected reach trajectories. In the Simon task, both RTs and reach trajectories were strongly influenced by stimulus-response correspondence. Despite the overall longer RTs in the Simon task, reaches were less direct and deviated toward the stimulus in noncorresponding trials. Thus, cognitive conflict was resolved before movement initiation in the SRC task, whereas it leaked into movement execution in the Simon task. Current theories of the Simon effect, such as the gating of response activation or response code decay, are inconsistent with our results. We propose that the SRC task was decomposed as approaching and avoiding the stimulus, which is sustained by stereotyped visuomotor routines. With complex stimulus-response relationships (Simon task), responses had to be coded as leftward and rightward, with more uncertainty about how to execute the action. This uncertainty permitted cognitive conflict to leak into the movement execution.     

Joint response–effect compatibility

When performing jointly on a task, human agents are assumed to represent their coactor’s share of this task, and research in various joint action paradigms has focused on representing the coactor’s stimulus–response assignments. Here we show that the response–effect (R–E) contingencies exploited by a coactor also affect performance, and thus might be represented as if they were used by oneself. Participants performed an R–E compatibility task, with keypresses producing spatially compatible or incompatible action effects. We did not observe any R–E compatibility effects when the task was performed in isolation (individual go–no-go). By contrast, small but reliable R–E compatibility effects emerged when the same task was performed in a joint setting. These results indicate that the knowledge of a coactor’s R–E contingencies can influence whether self-produced action effects are used for one’s own motor control.     

Time course of free-choice priming effects explained by a simple accumulator model

Unconscious visual stimuli can be processed by human observers and modulate their behavior. This has been shown for masked prime stimuli that influence motor responses to subsequent target stimuli. Beyond this, masked stimuli can also affect participants' behavior when they are free to choose one of two response alternatives. This finding demonstrates that an apparently free-choice between alternative behaviors can be subject to influences that are outside of awareness. We report three experiments which exhibit that the temporal dynamic of free-choice priming effects corresponds to that of forced-choice priming effects. Forced-choice priming effects were relatively robust against variations of prime stimuli but sensitive to physical features of target stimuli. Free-choice priming effects, in contrast, depended largely on the stimulus-response compatibility of the prime. A simple accumulator model which accounts for forced-choice response priming can also explain free-choice priming effects by the assumption that unconscious stimuli can initiate motor responses when participants are engaged in a speeded choice-reaction time task. According to our analyses free-choice priming results from a response selection mechanism which integrates conscious and unconscious information from external, stimulus driven sources and also from internal sources.     

Correlations between spatial compatibility effects: are arrows more like locations or words?

Responses are more efficient when their spatial mappings with features of targets are compatible compared to when they are incompatible, even when those features are irrelevant to task performance. Currently, a debate exists as to whether spatial information conveyed by different stimulus modes leads to qualitatively different spatial representations. We investigated the relations between three of the most commonly used spatial stimulus modes-arrows, locations, and location words-using correlations of compatibility effects between each of these modes as well as compatibility effects at different segments of their response time distributions. Our results show that when spatial information is irrelevant to task performance (the Simon task), the compatibility effects elicited by arrows and words are more strongly related with each other than with those of locations. However, when spatial information is task relevant (the stimulus-response compatibility SRC task), the compatibility effects elicited by arrows and locations are more related, and both are less related to the effect elicited by words. We suggest that these changing relations between stimulus modes are strategically determined based on which spatial coding technique for arrows is most advantageous to task performance. Furthermore, the varying relations between these spatial compatibility effects indicate that the compatibility effect with one stimulus mode is not always predictive of the compatibility effect in another mode.     

Validity and boundary conditions of automatic response activation in the Simon task.

Three experiments were conducted to determine whether spatial stimulus-response compatibility effects are caused by automatic response activation by stimulus properties or by interference between codes during translation of stimulus into response coordinates. The main evidence against activation has been that in a Simon task with hands crossed, responses are faster at the response location ipsilateral to the stimulus though manipulated by the hand contralateral to the stimulus. The experiments were conducted with hands in standard and in crossed positions and electroencephalogram measures showed coactivation of the motor cortex induced by stimulus position primarily during standard hand positions with visual stimuli. Only in this condition did the Simon effect decay with longer response times. The visual Simon effect appeared to be due to specific mechanisms of visuomotor information transmission that are not responsible for the effects obtained with crossed hands or auditory stimuli.