Self-induced versus reactive triggering of synchronous movements in a deafferented patient and control subjects.

The present study investigates the contribution of tactile-kinesthetic information to the timing of movements. The relative timing of simultaneous tapping movements of finger and foot (hand-foot asynchrony) was examined in a simple reaction time task and in discrete self-initiated taps (Experiment 1), and in externally triggered synchronization tapping (Experiment 2). We compared the performance of a deafferented participant (IW) to the performance of two control groups of different ages. The pattern of results in control groups replicates previous findings: Whereas positive hand-foot asynchronies (hand precedes foot) are observed in a simultaneous reaction to an auditory stimulus, hand-foot asynchronies are negative with discrete self-initiated as well as auditorily paced sequences of synchronized finger and foot taps. In the first case, results are explained by a simultaneous triggering of motor commands. In contrast, self-initiated and auditorily paced movements are assumed to be controlled in terms of their afferent consequences, as provided by tactile-kinesthetic information. The performance of the deafferented participant differed from that of healthy participants in some aspects. As expected on the basis of unaffected motor functions, the participant was able to generate finger and foot movements in reaction to an external signal. In spite of the lack of movement-contingent sensory feedback, the deafferented participant showed comparable timing errors in self-initiated and regularly paced tapping as observed in control participants. However, in discrete self-initiated taps IW's hand-foot asynchronies were considerably larger than in control participants, while performance did not differ from that of controls in continuous movement generation. These findings are discussed in terms of an internal generation of the movement's sensory consequences (forward-modeling).

Interactions between perception and action in a reaction task with overlapping S-R assignments.

We have used a novel task to study relationships between perception and action. Four experiments studied stimulus-response (S-R) relationships under conditions in which stimuli and responses were functionally unrelated (i.e., not assigned to each other by instruction) and merely overlapped in time. On each trial, participants carried out movements on a graphic tablet while observing motions displayed on a computer screen. The movement on trial n was specified by the motion observed on the previous trial n-1, whereas the motion observed on trial n specified the movement to be performed on trial n + 1. Results showed that stimulus motion had a contrast-like impact on response movement. Watching a small motion while performing a medium-sized movement increased movement size, whereas watching a large motion led to a decrease (Experiment 1). Further experiments showed that the contrast pattern was not affected by the mode of motion presentation (Experiment 2), or by the interval between motion and movement execution (Experiment 3). Contrast was also observed in the reverse direction, i.e., from action to perception (Experiment 4). We propose that the contrast effect is due to a mechanism for selective code modification. This mechanism acts to increase the distinctiveness of simultaneously activated perception and action codes in a common representational domain.

Tapping with peripheral nerve block. a role for tactile feedback in the timing of movements.

This study examines the impact of peripheral nerve block, that is, the elimination of tactile feedback on synchronization performance. In a tapping experiment in which subjects were instructed to tap in synchrony with an auditory pacing signal, three different tasks were studied under conditions with and without peripheral nerve block: standard tapping with tactile contact, isometric tapping, and contact-free tapping. In addition, the maximum tapping rate was registered both with and without peripheral nerve block. It was found that the anticipatory error, usually observed in synchronization tasks, was affected by the peripheral nerve block in the standard tapping and the isometric tapping task. In both tasks, local anesthesia led to an increase in asynchrony between the pacing signal and the tap. Performance remained unimpaired in those tasks in which tactile information was assumed to play a minor role (maximum tapping rate and contact-free tapping). The results clearly demonstrate the importance of tactile feedback for the timing of movements. The predictions of a model assuming a strong correlation between the amount of sensory feedback and the size of the negative asynchrony in synchronization tasks were examined and discussed.

The Theory of Event Coding (TEC): a framework for perception and action planning.

Traditional approaches to human information processing tend to deal with perception and action planning in isolation, so that an adequate account of the perception-action interface is still missing. On the perceptual side, the dominant cognitive view largely underestimates, and thus fails to account for, the impact of action-related processes on both the processing of perceptual information and on perceptual learning. On the action side, most approaches conceive of action planning as a mere continuation of stimulus processing, thus failing to account for the goal-directedness of even the simplest reaction in an experimental task. We propose a new framework for a more adequate theoretical treatment of perception and action planning, in which perceptual contents and action plans are coded in a common representational medium by feature codes with distal reference. Perceived events (perceptions) and to-be-produced events (actions) are equally represented by integrated, task-tuned networks of feature codes--cognitive structures we call event codes. We give an overview of evidence from a wide variety of empirical domains, such as spatial stimulus-response compatibility, sensorimotor synchronization, and ideomotor action, showing that our main assumptions are well supported by the data.

An analysis of ideomotor action.

This article presents a framework based on the work of R. H. Lotze (1852), W. James (1890), and A. G. Greenwald (1970) for understanding ideomotor actions that tend to arise when individuals watch others perform certain actions. Two principles of ideomotor action induction are distinguished: perceptual induction, in which people tend to perform the movements they see, and intentional induction, in which people tend to perform movements suited to achieve what they would like to see. In 3 experiments, ideomotor hand, head, and foot movements were studied while participants watched a ball traveling toward a target. Results showed strong support for intentional induction, weaker support for perceptual induction, and a strong impact of the effector studied. The representational basis of action induction (stimulus vs. goal representations) and the automaticity of the underlying processing (task-dependent vs. task-independent induction) were considered.

Correspondence effects with manual gestures and postures: a study of imitation.

In this study, the authors applied methods and theories from research of stimulus-response compatibility (SRC) to action imitation. In 6 experiments, they adopted the logic of the Simon paradigm (B. Hommel & W. Prinz, 1996) to explore interference between task-relevant symbolic stimulus features (color) and task-irrelevant iconic stimulus features (2 hand gestures and 2 postures). The same 2 hand gestures served as responses. Pronounced correspondence effects for both gestures and postures showed up throughout. In line with theories of SRC, the authors account for these correspondence effects in terms of overlap arising between stimulus and response features in a common representational domain. As a specific extension of this approach, they propose 2 functionally independent mechanisms: One operates movement-based when dynamic information is provided, and the other operates state-based with static postures as stimuli. Implications for theories of both SRC and action imitation are discussed.

Neuromagnetic correlates of sensorimotor synchronization.

Sensorimotor synchronization tasks, in which subjects have to tap their finger in synchrony with an isochronous auditory click, typically reveal a synchronization error with the tap preceding the click by about 20 to 50 msec. Although extensive behavioral studies and a number of different explanatory accounts have located the cause of this so-called "negative asynchrony" on different levels of processing, the underlying mechanisms are still not completely understood. Almost nothing is known about the central processes, in particular, which sensory or motor events are synchronized by subjects. The present study examined central-level processing in synchronization tasks with magnetoencephalography (MEG). Eight subjects synchronized taps with their right index finger to an isochronous binaural pacing signal presented at an interstimulus interval of 800 msec. To gain information on central temporal coupling between "tap" and "click," evoked responses were averaged time-locked to the auditory signal and the tap onset. Tap-related responses could be explained with a three dipole model: One source, peaking at approximately 77 msec before tap onset, was localized in contralateral primary motor cortex (MI); the two other sources, peaking approximately at tap onset and 75 msec after tap onset, in contralateral primary somatosensory cortex (SI). Temporal coupling of these sources was compared in relation to different trigger points. The second SI source was equally well time-locked to the tap and to the auditory click. Furthermore, analysis of the time locking of this source activity as a function of the temporal order of tap and click showed that the second event - irrespective whether tap or click - was decisive in triggering the second SI source. This suggests that subjects use mainly sensory feedback in judging and evaluating whether they are "keeping time."

Sensorimotor synchronization: the impact of temporally displaced auditory feedback.

When subjects are asked to tap in synchrony to a regular sequence of stimulus events (e.g., clicks), performance is not perfect in that, usually, an anticipation of the tap is observed. The present study examines the influence of temporally displaced auditory feedback on the size of this anticipatory error. Whereas earlier studies have shown that this asynchrony exhibits a linear increase in size as a function of an increasing delay in such additional auditory feedback, this study compared the impact of shifting feedback forward in time (i.e., feedback presented before the tap) with that of delayed auditory feedback. Results showed that the impact of feedback displacement on the amount of asynchrony differed for positive and negative displacements. Delayed feedback led to an increase in asynchrony, whereas negative displacements had (almost) no effect. This finding is related to a model assuming that the various feedback components arising from the tap (tactile, kinesthetic, auditory) are integrated to form one central representation, and that the timing of this central representation arises from a linear combination of the components involved.

Localizing the first position of a moving stimulus: the Fröhlich effect and an attention-shifting explanation.

When subjects are asked to determine where a fast-moving stimulus enters a window, they typically do not localize the stimulus at the edge, but at some later position within that window (Fröhlich effect). We report five experiments that explored this illusion. An attentional account is tested, assuming that the entrance of the stimulus in the window initiates a focus shift toward it. While this shift is under way, the stimulus moves into the window. Because the first phenomenal (i.e., explicitly reportable) representation of the stimulus will not be available before the end of the focus shift, the stimulus is perceived at some later position. In Experiment 1, we established the Fröhlich effect and showed that it size depends on stimulus parameters such as movement speed and movement direction. In Experiments 2 and 3, we examined the influence of eye movements and tested whether the effect changed when the stimuli were presented within a structural background or when they started from different eccentricities. In Experiments 4 and 5, specific predictions from the attentional model were tested: In Experiment 4 we showed that the processing of the moving stimulus benefits from a preceding peripheral cue indicating the starting position of the subsequent movement, which induces a preliminary focus shift to the position where the moving stimulus would appear. As a consequence the Fröhlich effect was reduced. Using a detection task in Experiment 5, we showed that feature information about the moving stimulus is lost when it falls into the critical interval of the attention shift. In conclusion, the present attentional account shows that selection mechanisms are not exclusively space based; rather, they can establish a spatial representation that is also used for perceptual judgement--that is, selection mechanisms can be space establishing as well.

Delayed auditory feedback in synchronization.

In two experiments, the effect of feedback delay on synchronization performance was examined. In Experiment 1, whether feedback delay has an effect on the asynchrony between tap and click was investigated, and a transfer effect between conditions, observed in a preliminary experiment, was studied. Experiment 2 served as a control experiment that assured that the effects were not caused by order effects. A linear relationship between the size of the delay and the asynchrony between tap and click was observed; that is, with increasing delay, the size of the asynchrony increased as well. The results support an extended version of the Paillard-Fraisse hypothesis that accounts for the results observed in synchronization tasks in which more than a single source of feedback from the tap is available. The results indicate that all tap-related information is integrated into a joint event code representing that tap at a central level. These codes are superimposed on click-related codes and are therefore responsible for the observed asynchronies. Moreover, the results suggest that tap-related codes arise from a linear combination of their tactile-kinesthetic and auditory components.

Synchronizing actions with events: the role of sensory information.

Tasks requiring the subject to tap in synchrony to a regular sequence of stimulus events (e.g., clicks) usually elicit a response pattern in which the tap precedes the click by about 30-50 msec. This "negative asynchrony" was examined, first, by instructing subjects to use different effectors for tapping (hand vs. foot; Experiments 1 and 2), and second, by administering extrinsic auditory feedback in addition to the intrinsic tactile/kinesthetic feedback (Experiment 2). Experiment 3 controlled whether the results observed in Experiment 2 were due to purely sensory factors within the auditory modality. Results suggest that taps are synchronized with clicks at the central level by superimposing two sensory codes in time: the tactile/kinesthetic code that represents the tap (the afferent movement code) and the auditory code that represents the click (the afferent code that results from the guiding signal). Because the processing times involved in code generation are different for these two central codes, the tap has to lead over the click.