Motor cortex activation induced by a mirror: evidence from lateralized readiness potentials

Similar motor regions are activated during voluntarily executed or observed movements. We investigated whether observing movements of one's own hand through a mirror will generate activations in the cortical motor regions of both the moving and nonmoving hands. Using the lateralized readiness potential (LRP), an electrophysiological correlate of premotor activation in the primary motor cortex, we recorded evoked responses to movements while subjects were viewing the performing (right) hand through a mirror placed sagittally, giving the impression that the left hand was performing the task. Reliable LRPs were recorded in relation to the seen hand, indicating motor cortex activity in the contralateral hemisphere of the inactive hand while the opposite hand was performing the movement.     

Delayed flanker effects on lateralized readiness potentials

When participants were required to respond to a relevant central target and ignore irrelevant flanking stimuli, the flankers produced a response compatibility effect. Electrophysiological studies have shown that irrelevant flanker stimuli can affect the motor system. The present experiments further examined the characteristics of flanker effects on the motor system. Sixty participants responded in the flanker task to arrows (experiment 1) or letters (experiment 2). To examine time and extent of flanker effects on the motor system, target onset was delayed with blocked or random stimulus-onset asynchronies (SOA). With SOA of 0 and 100 ms, flanker effects on behavioral measures were reduced in random as compared to blocked conditions, but enhanced with SOA of 400 ms. With SOA of 400 ms, flanker effects on the early lateralized readiness potentials (LRP) were reduced in blocked as compared to random conditions, indicating that the onset of flanker effects on the LRP was delayed. Response-locked LRPs suggest that flanker and target stimuli activate the motor system successively. Findings challenge current theories of the flanker compatibility effect.     

Response activation impairments in schizophrenia: evidence from the lateralized readiness potential

Previous research has demonstrated deficits in preresponse motor activity in schizophrenia, as evidenced by a reduced lateralized readiness potential (LRP). The LRP deficit could be due to increased activation of the incorrect response (e.g., failure to suppress competition) or to reduced activation of the correct response (e.g., a low-level impairment in response preparation). To distinguish these possibilities, we asked whether the LRP impairment is increased under conditions of strong response competition. We manipulated the compatibility of stimulus-response mappings (Experiment 1) and the compatibility of the target with flankers (Experiment 2). In both experiments, the patient LRP was reduced as much under conditions of low response competition as under high competition. These results are incompatible with a failure of patients to suppress competition and are instead consistent with a deficit in activating the correct response.     

Asymmetric switch cost: an investigation using lateralized readiness potentials

Switching to a dominant task incurs larger costs than switching to a non-dominant task. This study investigated whether this cost asymmetry derives from the inhibition of the dominant task rule that occurred during the previous trial. Participants were presented with a five-letter array consisting of (left) and (right), and asked to respond to the central target letter after being informed about the task rule with a pre-cue. The rule was switched between dominant (left-hand to, right-hand to) and non-dominant tasks (right-hand to, left-hand to) every two trials. Reaction times revealed the asymmetrical switch cost and the effect of target-flanker congruency. Stimulus-locked lateralized readiness potentials showed that the dominant task rule was inhibited during the non-dominant task, whereas this inhibition was not carried over to the dominant task trial.     

The structure of motor programming: Evidence from reaction times and lateralized readiness potentials

There is a widely accepted notion that movement elements are assembled prior to movement execution in a central motor programming stage. However, it is not clear how this stage is structured—whether it is a unitary stage allowing different motor parameters to cross talk or whether there are several independent processes dealing with each motor parameter. We addressed this question by orthogonally manipulating two movement‐related factors: response sequence complexity and movement duration. Both factors yielded main effects on reaction time but no interaction. Additive effects of both factors on the onsets of response‐ but not stimulus‐synchronized lateralized readiness potentials suggest separable motoric loci of sequence complexity and duration. These findings are at variance with the notion of a unitary movement programming stage.     

Partial information can be transmitted in an auditory channel: inferences from lateralized readiness potentials

With the two-choice go/no-go paradigm, we investigated whether timbre attribute can be transmitted as partial information from the stimulus identification stage to the response preparation stage in auditory tone processing. We manipulated two attributes of the stimulus: timbre (piano vs. violin) and acoustic intensity (soft vs. loud) to ensure an earlier processing of timbre than intensity. We associated the timbre attribute more with go trials. Results showed that lateralized readiness potentials (LRPs) were consistently elicited in no-go trials. This showed that the timbre attribute had been transmitted to the response preparation stage before the intensity attribute was processed in the stimuli identification stage. Such a result provides evidence for the continuous model and asynchronous discrete coding (ADC) model in information processing. We suggest that partial information can be transmitted in an auditory channel.     

Tracing sequential waves of rapid visuomotor activation in lateralized readiness potentials

Feedforward activation processes are widely regarded as crucial for the automatic initiation of motor responses, whereas recurrent processes are often regarded as crucial for visual awareness. Here, we used a set of behavioral criteria to evaluate whether rapid processing in the human visuomotor system proceeds as would be expected of a feedforward system that works independent of visual awareness. We measured lateralized readiness potentials (LRPs) for key-press responses to color targets ("masks") preceded by masked color primes mapped onto the same or opposite response, and traced the time-course of motor activation as a function of different prime and mask characteristics. LRP time-courses showed that initial motor activation occurred in prime direction and was time-locked to prime onset. Response activation was then captured on-line by the mask signal, with motor activation now time-locked to the mask and proceeding in mask direction. Crucially, the time-course of early activation by the prime was independent of all mask characteristics. This invariance in early priming effects contrasted with large differences in visual awareness for the prime produced by different masks. Results suggest that primed responses to color stimuli are controlled by feedforward waves of activation sequentially elicited by prime and mask signals traveling rapidly enough to escape the recurrent processes leading to backward masking.     

Locus of the effect of temporal preparation: evidence from the lateralized readiness potential

It is well established that reaction time (RT) is shorter when a response signal is preceded by a warning signal, because the warning signal causes the participant to prepare for the upcoming response. A review of chronometric and psychophysiological studies reveals the prevailing view that this temporal preparation operates mainly at a motor level speeding up rather late processes. To assess the locus of this preparation effect, we conducted two experiments employing the lateralized readiness potential (LRP). Contrary to this prevailing view, the results of both experiments clearly indicate that temporal preparation enhances the processing speed of relatively early processes, because a manipulation of temporal uncertainty affected RT, the P300 latency, and the stimulus-to-LRP interval but not the LRP-to-keypress interval.     

The lateralized readiness potential and response kinetics in response-time tasks

Previous studies have found that the magnitude of the lateralized readiness potential (LRP) at the time of response initiation is constant across spontaneous variations in response time in both cued and uncued, speeded tasks. Other studies have found that the LRP is also unaffected by instructed changes in peak response force and time to peak force in cued, self-paced tasks, but that the LRP is sensitive to instructed changes in force gain rate in uncued, self-paced tasks. The present study examined the LRP in an uncued, speeded task as a function of response time and several measures of response kinetics. The magnitude of the LRP at the time of electromyographic onset was constant across spontaneous variations in all measures. The peak of the contingent negative variation did vary as a function of peak response force and integrated force to peak, but not response time. These findings support the idea that the LRP in speeded tasks is a selective, on-line index of the preparation associated with using a particular hand, and is not an index of the elements of motor programming that determine subsequent response kinetics.     

Covert motor activity on NoGo trials in a task sharing paradigm: evidence from the lateralized readiness potential

Previous studies on task sharing propose that a representation of the co-actor's task share is generated when two actors share a common task. An important function of co-representation seems to lie in the anticipation of others' upcoming actions, which is essential for one's own action planning, as it enables the rapid selection of an appropriate response. We utilized measures of lateralized motor activation, the lateralized readiness potential (LRP), in a task sharing paradigm to address the questions (1) whether the generation of a co-representation involves motor activity in the non-acting person when it is other agent's turn to respond, and (2) whether co-representation of the other's task share is generated from one's own egocentric perspective or from the perspective of the actor (allocentric). Results showed that although it was the other agent's turn to respond, the motor system of the non-acting person was activated prior to the other's response. Furthermore, motor activity was based on egocentric spatial properties. The findings support the tight functional coupling between one's own actions and actions produced by others, suggesting that the involvement of the motor system is crucial for social interaction.     

Impaired response selection in schizophrenia: Evidence from the P3 wave and the lateralized readiness potential

Reaction times (RTs) are substantially prolonged in schizophrenia patients, but the latency of the P3 component is not. This suggests that the RT slowing arises from impairments in a late stage of processing. To test this hypothesis, 20 schizophrenia patients and 20 control subjects were tested in a visual oddball paradigm that was modified to allow measurement of the lateralized readiness potential (LRP), an index of stimulus-response translation processes. Difference waves were used to isolate the LRP and the P3 wave. Patients and control subjects exhibited virtually identical P3 difference waves, whereas the LRP difference wave was reduced in amplitude and delayed in latency in the patients. These results indicate that, at least in simple tasks, the delayed RTs observed in schizophrenia are primarily a consequence of impairments in the response selection and preparation processes that follow perception and categorization.     

Preparation of response force and movement direction: onset effects on the lateralized readiness potential

Two experiments assessed the preparatory effects of advance information about response force and movement direction on the lateralized readiness potential (LRP). In a choice reaction time (RT) task, an imperative stimulus required an isometric flexion or an extension of the left or right index finger. Prior information about response force or about movement direction reduced RT and shortened the interval from the onset of the imperative stimulus up to the onset of the LRP. Advance information, however, about direction but not about force decreased the interval from LRP onset to the onset of the overt response. The identical pattern of results was obtained in a second experiment, in which each participant performed both precue conditions. The findings of both experiments support the notion that response force is specified before movement direction. These results are consistent with the view accordingly different mechanisms are involved in the specification of muscle force and movement direction.     

The lateralized readiness potential: relationship between human data and response activation in a connectionist model.

Psychophysiological measures such as the lateralized readiness potential (LRP) have been used to study information processing in the Eriksen flankers task. The data provided by these measures are consistent with a continuous flow theory, which proposes that the output of stimulus evaluation is continuously available to the response channels. Cohen et al. (1992) realized this theory in a connectionist model and showed that its behavior corresponded to that of human subjects in the flankers task. We report here a modification of the model and an analysis of the degree to which simulated LRPs (based on the activation functions of the response units of the model) resemble the actual LRPs of human subjects in the same task. Across a variety of different experimental conditions and outcomes, there was a marked correspondence between the simulated and actual LRPs. These observations strengthen the propriety of the connectionist model and of the continuous flow theory on which it is based.     

Age differences in task switching and response monitoring: evidence from ERPs

This study investigates age differences in the flexible adaptation to changing demands on task switching and conflict processing. We applied a cued task-switching version of the Stroop task and manipulated the ratio of conflict trials. During task preparation, the P300 varied as a function of conflict ratio and a later positive component was larger for switch than non-switch trials. Stimulus-related conflict processing as indicated by a negativity for incompatible trials (Ni) was delayed for older adults. Moreover, the Ni varied as a function of conflict ratio and was larger for switch than for non-switch trials. Age differences were also obtained in the correct response negativity (CRN). CRN was larger on incompatible trials and this CRN-compatibility effect was enhanced when incompatible trials were infrequent in younger, but not in older adults. Our findings suggest impairments of older adults primarily in response-related conflict processing and in the flexible adaptation to changing task contexts.     

Maintenance of response readiness in patients with Parkinson's disease: evidence from a simple reaction time task

The authors explored the effect of Parkinson's disease (PD) on the generation and maintenance of response readiness in a simple reaction time task. They compared performance of idiopathic PD patients without dementia, age-matched controls, and younger controls over short (1-, 3-, and 6-s) and long (12- and 18-s) foreperiod intervals. After each trial, the authors probed memory for visual information that also had to be maintained during the trial interval. Patients and controls did not differ overall in their ability to maintain readiness over long delays. However, within the PD group only, errors in maintaining visual information were correlated with difficulty in maintaining readiness, suggesting that systems impaired in PD may facilitate the maintenance of processing in both motor and cognitive domains.     

The application of jackknife-based onset detection of lateralized readiness potential in correlative approaches

The onset of the lateralized readiness potential (LRP) represents an interesting parameter within mental chronometry. According to Miller, Patterson, and Ulrich (1998), reliability increases when LRP onsets are detected in jackknifed LRP waveforms. Until now, jackknifed LRP onsets could be analyzed in factorial designs only. The aim of the present study was to extend the application of jackknifing to correlative approaches (e.g., in personality research). Using different onset scoring techniques, in several simulations run on realistic LRP data jackknifing and single-subject procedures were compared regarding their estimates of a simulated true correlation. For all scoring techniques but regression, jackknifed coefficients were on average higher than the respective single-subject coefficients, and statistical power was higher. Overall, combining jackknifing with a 50%-relative onset criterion is recommended.     

Genetic variation in dopamine moderates neural response during reward anticipation and delivery: evidence from event-related potentials

Neuroimaging studies have found moderating effects of dopamine genes during both the anticipation and delivery of rewards, particularly the catechol-O-methyltransferase (COMT) genotype. Event-related potential studies, meanwhile, have focused on the stimulus-preceding negativity (SPN) and the feedback negativity (FN) during reward anticipation and delivery, respectively. In anticipation of uncertain outcomes, we observed an increased SPN among Met homozygotes. We also observed an increased FN among Met homozygotes in response to outcome delivery, an effect that was driven primarily by an increased response to monetary gains. The COMT genotype moderates event-related potential responses during both the anticipation and delivery of uncertain reward, suggesting that the SPN and FN are sensitive to dopaminergically mediated and reward-related neural activity.     

Early emotion word processing: evidence from event-related potentials

Behavioral and electrophysiological responses were monitored to 80 controlled sets of emotionally positive, negative, and neutral words presented randomly in a lexical decision paradigm. Half of the words were low frequency and half were high frequency. Behavioral results showed significant effects of frequency and emotion as well as an interaction. Prior research has demonstrated sensitivity to lexical processing in the N1 component of the event-related brain potential (ERP). In this study, the N1 (135-180 ms) showed a significant emotion by frequency interaction. The P1 window (80-120 ms) preceding the N1 as well as post-N1 time windows, including the Early Posterior Negativity (200-300 ms) and P300 (300-450 ms), were examined. The ERP data suggest an early identification of the emotional tone of words leading to differential processing. Specifically, high frequency negative words seem to attract additional cognitive resources. The overall pattern of results is consistent with a time line of word recognition in which semantic analysis, including the evaluation of emotional quality, occurs at an early, lexical stage of processing.     

Error-related brain potentials are differentially related to awareness of response errors: evidence from an antisaccade task

The error negativity (Ne/ERN) and error positivity (Pe) are two components of the event-related brain potential (ERP) that are associated with action monitoring and error detection. To investigate the relation between error processing and conscious self-monitoring of behavior, the present experiment examined whether an Ne and Pe are observed after response errors of which participants are unaware. Ne and Pe measures, behavioral accuracy, and trial-to-trial subjective accuracy judgments were obtained from participants performing an antisaccade task, which elicits many unperceived, incorrect reflex-like saccades. Consistent with previous research, subjectively unperceived saccade errors were almost always immediately corrected, and were associated with faster correction times and smaller saccade sizes than perceived errors. Importantly, irrespective of whether the participant was aware of the error or not, erroneous saccades were followed by a sizable Ne. In contrast, the Pe was much more pronounced for perceived than for unperceived errors. Unperceived errors were characterized by the absence of posterror slowing. These and other results are consistent with the view that the Ne and Pe reflect the activity of two separate error monitoring processes, of which only the later process, reflected by the Pe, is associated with conscious error recognition and remedial action.