Cognitive Control Structures in the Imitation Learning of Spatial Sequences and Rhythms-An fMRI Study.

Imitation learning involves the acquisition of novel motor patterns based on action observation (AO). We used event-related functional magnetic resonance imaging to study the imitation learning of spatial sequences and rhythms during AO, motor imagery (MI), and imitative execution in nonmusicians and musicians. While both tasks engaged the fronto-parietal mirror circuit, the spatial sequence task recruited posterior parietal and dorsal premotor regions more strongly. The rhythm task involved an additional network for auditory working memory. This partial dissociation supports the concept of task-specific mirror mechanisms. Two regions of cognitive control were identified: 1) dorsolateral prefrontal cortex (DLPFC) was found to be more strongly activated during MI of novel spatial sequences, which allowed us to extend the 2-level model of imitation learning by Buccino et al. (2004) to spatial sequences. 2) During imitative execution of both tasks, the posterior medial frontal cortex was robustly activated, along with the DLPFC, which suggests that both regions are involved in the cognitive control of imitation learning. The musicians' selective behavioral advantage for rhythm imitation was reflected cortically in enhanced sensory-motor processing during AO and by the absence of practice-related activation differences in DLPFC during rhythm execution.

Effect Anticipation Affects Perceptual, Cognitive, and Motor Phases of Response Preparation: Evidence from an Event-Related Potential (ERP) Study.

The anticipation of action effects is a basic process that can be observed even for key-pressing responses in a stimulus-response paradigm. In Ziessler et al.'s (2012) experiments participants first learned arbitrary effects of key-pressing responses. In the test phase an imperative stimulus determined the response, but participants withheld the response until a Go-stimulus appeared. Reaction times (RTs) were shorter if the Go-stimulus was compatible with the learned response effect. This is strong evidence that effect representations were activated during response planning. Here, we repeated the experiment using event-related potentials (ERPs), and we found that Go-stimulus locked ERPs depended on the compatibility relationship between the Go-stimulus and the response effect. In general, this supports the interpretation of the behavioral data. More specifically, differences in the ERPs between compatible and incompatible Go-stimuli were found for the early perceptual P1 component and the later frontal P2 component. P1 differences were found only in the second half of the experiment and for long stimulus onset asynchronies (SOAs) between imperative stimulus and Go-stimulus, i.e., when the effect was fully anticipated and the perceptual system was prepared for the effect-compatible Go-stimulus. P2 amplitudes, likely associated with evaluation and conflict detection, were larger when Go-stimulus and effect were incompatible; presumably, incompatibility increased the difficulty of effect anticipation. Onset of response-locked lateralized readiness potentials (R-LRPs) occurred earlier under incompatible conditions indicating extended motor processing. Together, these results strongly suggest that effect anticipation affects all (i.e., perceptual, cognitive, and motor) phases of response preparation.

Using tools with real and imagined tool movements.

WHEN USING LEVER TOOLS, SUBJECTS HAVE TO DEAL WITH TWO, NOT NECESSARILY CONCORDANT EFFECTS OF THEIR MOTOR BEHAVIOR: the body-related proximal effects, like tactile sensations from the moving hand, and/or more external distal effects, like the moving effect points of the lever. As a consequence, spatial compatibility relationships between stimulus (S; at which the effect points of the lever aim at), responding hand (R) and effect point of the lever (E) play a critical role in response generation. In the present study we examine whether the occurrence of compatibility effects needs real tool movements or whether a similar response pattern can be already evoked by pure mental imaginations of the tool effects. In general, response times and errors observed with real and imagined tool movements showed a similar pattern of results, but there were also differences. With incompatible relationships and thus more difficult tasks, response times were reduced with imagined tool movements than compared with real tool movements. On the contrary, with compatible relationships and thus high overlap between proximal and distal action effects, response times were increased with imagined tool movements. Results are only in parts consistent with the ideomotor theory of motor control.

The activation of effect codes in response preparation: new evidence from an indirect priming paradigm.

Evidence for the anticipation of environmental effects as an integral part of response planning comes mainly from experiments in which the effects were physically presented. Thus, in these studies it cannot be excluded that effect codes were activated during response preparation only because the effects were displayed as external stimuli before response execution. In order to provide more clear-cut evidence for the anticipation of response effects in action planning, we performed a series of three experiments using a new paradigm, where displaying effect codes before the response was avoided. Participants first learned arbitrary effects of key-pressing responses. In the following test phase they were instructed to execute a response only if a Go stimulus was presented after a variable stimulus onset asynchrony (SOA). The Go stimulus was either compatible or incompatible with the effect, but independent of the response. In Experiment 1 we tested the paradigm with two responses and two effects. We found a significant compatibility effect: If the Go stimulus was compatible with the response effect, responses were initiated faster than in incompatible trials. In Experiment 2 response effects were only present in the acquisition phase, but not in the test phase. The compatibility effect disappeared, indicating that the results of Experiment 1 were indeed related to the anticipation of the forthcoming response effects. In Experiment 3 we extended this paradigm by using a larger number of stimuli and response alternatives. Again we found a robust compatibility effect, which can only be explained if the effect representations are active before response execution. The compatibility effects in Experiments 1 and 3 did not depend on the SOA. The fact that the Go stimulus affected response preparation at any time indicates that the role of effect anticipation is not limited to response selection.

The temporal dynamics of effect anticipation in course of action planning.

"Strong" versions of the ideomotor theory of action control claim that anticipations of the environmental effects that actions bring about are mandatory for response selection. This is considered to be the one and only way of how actions can be voluntarily selected. We studied this notion in a series of four experiments where we adapted the flanker paradigm to investigate the involvement of effect codes in the preparation of motor responses. Participants first learned that their responses to stimulus letters were contingently followed by the presentation of a new letter on the screen. In the second phase of the experiments, the action-demanding letters were presented together with the effects of the correct response, effects of other responses, or neutral letters. Varying the stimulus onset asynchrony (SOA) between target stimuli and the flanking effect stimuli provides the opportunity to investigate the temporal dynamics of the activation of effect codes. Hence, flanker stimuli were presented before, simultaneously with, or after the onset of the target. The results indicate that effect-related information from the flanker stimuli is involved in the preparation process, but mainly in later phases of response preparation. The observed pattern of results suggests that, at least under conditions where responses are determined by stimuli, effect codes are activated in course of response planning to enable the evaluation of the executed response and the monitoring of response execution, but they do not automatically activate the responses themselves.

Binding in voluntary action control.

The last decade has seen a proliferation of empirical studies that seek to understand how the cognitive system links voluntary motor actions with their perceptual effects. A view that has found considerable support in this research is the ideomotor approach to action control which holds that actors select, initiate, and execute a movement by activating anticipatory codes of the movement's sensory effects. We, first review the empirical evidence from different paradigms showing that effects of voluntary actions become anticipated during response production. In a second step we survey empirical data investigating the nature of the mechanisms that link voluntary motor actions with their intended and expected perceptual effects. We argue that the integration, or binding, of perceptual and motor codes occurs in action planning where features of intended effects are selectively bound to features of the actions that are selected to achieve these effects in the environment. As a final step we will summarize empirical findings that may elucidate the particular roles of effect-code activation in response production and control.

Anticipated action consequences as a nexus between action and perception: evidence from event-related potentials.

We used high-density event-related potentials (ERP) in a modified flanker paradigm to study the role of anticipated action consequences in action planning and the role of anticipation in the perception of action consequences. Prior to the experiment, participants were trained to classify target letters in a four-alternative forced-choice task; another letter was presented as an effect following each response. After participants had thus acquired the response-effect contingencies, in the experiment effect letters were presented as flankers to target letters. Effect-compatible flankers were letters that were learned as effects of the correct response to the target; effect-incompatible ones were learned as effects of other responses; neutral flankers were never presented as action effects. To help distinguish early and late effects of flankers on target processing, flankers were presented either simultaneously with the target or after a delay. We found that effect-incompatible flankers resulted in longer, than other flankers, time between the onset of the response-locked lateralized readiness potential and the response, indicating extended motor processing. ERP evoked by the effect-incompatible flankers differed from those evoked by other flankers in early perceptual component P1 and in later frontal component P2 reflecting stimulus evaluation and conflict detection. These results show that anticipating action consequences involves brain systems ranging from perceptual to executive; anticipated action effects constitute a link between perception and action.

The role of anticipation and intention in the learning of effects of self-performed actions.

The anticipative learning model for acquiring action-effect relations states that the acquisition of action-effect relations depends on processes that are part of action planning, in particular the anticipation of possible effects. Experiment 1 shows that response planning is indeed crucial for the learning of response effects. In this experiment distractors (tones) were presented either during response preparation or in the time interval between response execution and the presentation of a response effect. Response-effect learning was impaired when the distractors were presented during response preparation. The finding is consistent with the assumption that the distractors impaired the anticipation of potential effects and therefore reduced effect learning. In Experiment 2 all responses had two effects. Participants were instructed to produce one of the effects. Under this condition, response-effect learning was only found for the instructed effect, not for the non-instructed effect. The two experiments thus support the view that response-effect learning is selective and depends on the anticipation of potential effects during response planning. The results are discussed in terms of a model that explains both the learning of response-effect relations and the use of these effects for action control within the same theoretical framework.

Sequence learning in Parkinson's disease: the effect of spatial stimulus-response compatibility.

Patients with Parkinson's disease (PD) have repeatedly demonstrated reduced sequence-specific learning effects in serial reaction time tasks (SRTs). Previous research with PD patients has mainly employed the 'classical' SRT task, involving a spatially compatible assignment of stimuli and responses. From cognitive research, it is known that spatial compatibility triggers rapid, automatic responses in the direction of the stimulus. Automatic responding has shown to be disinhibited in PD patients and may therefore interfere with stimulus anticipation during the learning process. The aim of the present study was to test this hypothesis by investigating if reduced sequence-specific learning depends on spatial stimulus-response compatibility. PD patients and age-matched controls were examined either with an SRT variant involving central stimulus presentation, thereby preventing automatic linking of stimulus and response locations, or with a spatially compatible SRT task. Patients showed reduced sequence-specific learning effects only when the stimulus-response assignment was spatially compatible. This pattern of results confirms the hypothesis that sequence learning deficits in PD may result from a predominance of automatic response activation over learning-based stimulus anticipations during the learning phase.