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.

ERP correlates of associative learning.

We examined changes of event-related potentials (ERPs) while participants learned stimulus-to-stimulus relations in an S1-S2 task. The design allowed for separating processes of associative learning from nonspecific effects. Participants had to respond to S2 by a left or right key-press dependent on S2 identity (letter W or M). Preparation for S2 could be improved by using the associative information given by S1. The S1 was an arrow pointing to the left or right. In combination with its color, arrow direction was informative about location and identity of S2, but participants were not informed about the relevance of color. Arrows in two of the colors were fully predictive for the S2 whereas the third color gave no valid information. This third stimulus controlled for habituation and procedural learning. Six blocks with 200 trials each and all three S1 colors in random order were presented. Behavioral and ERP differences in each block between "learning" and control trials were used to identify processes of associative learning. Several effects of associative learning were identified indicating the involvement of specific stages of information processing: a continuous increase of P3 amplitude evoked by S1 was accompanied by a decrease of P3 evoked by S2. These changes reflected the modifications of stimulus weights for response selection and the strengthened association between the two stimulus complexes in the time course of learning. The related motor preparation benefited from learning too, expressed in a decrease of CNV amplitude and an increase of LRP amplitude. Finally a decrease of N1 amplitude evoked by S2 indicated the reduced need to allocate spatial attention to the S2 location according to the learned meaning of S1.

The influence of time pressure and cue validity on response force in an S1-S2 paradigm.

Hypotheses about variations of response force have emphasised the influences of arousal and of motor preparation. To study both types of influences in one experiment, the effects of time pressure and of validity of S1 were investigated in tasks wherein a first stimulus (S1) indicated the most probable response (80% valid) required after a second stimulus (S2). Under time pressure, responses were executed more forcefully while, as could be expected, response times were shorter and errors were more frequent. This pattern of results was not only obtained when time pressure was varied between blocks, but also when varied from trial to trial, by information given by S2. Also invalidly cued responses were executed more forcefully but, as could be expected, in contrast to time pressure, response times were longer and errors were more frequent. The results demonstrate that latency and force of responses may vary in different directions. Ways are outlined on how current hypotheses must be extended in order to account for these results.

Dimensional overlap between arrows as cueing stimuli and responses?. Evidence from contra-ipsilateral differences in EEG potentials.

In the S1-S2 interval, 400 ms after an arrow as S1, an EEG-potential difference occurs between scalp sites contralateral and ipsilateral to arrow direction. Eimer [J. Exp. Psychol. Hum. Percept. Perform. 21 (1995) 837-854] interpreted this difference as a sign of automatic activation of the manual response, due to dimensional overlap of arrows and responses. However, according to Kornblum et al.'s [Psychol. Rev. 97 (1990) 253-270] notion of dimensional overlap, responses can only be automatically primed if they are included in the response set. Therefore, participants of the present study had to respond to S2 in separate blocks either by key-press, as in Eimer [J. Exp. Psychol. Hum. Percept. Perform. 21 (1995) 837-854], or by making saccades. In addition, contra-ipsilateral differences were recorded not only from central positions, overlying the hand-motor area, but across the whole scalp. Contralateral negativity at 400 ms after S1 was indeed found over the hand-motor area in the 'hand blocks'. However, this 'L-400' (=lateralization at 400 ms) was generally as large in the 'eye' blocks as in the 'hand' blocks. Therefore, L-400 does not reflect automatic activation of manual responses in the sense of Kornblum et al. [Psychol. Rev. 97 (1990) 253-270]. Further, its topographical maximum was more anterior than the hand-motor-related negativity that preceded the manual response ('LRP') with its maximum at central sites. Therefore, L-400 probably does not originate in the hand-motor cortex. Rather, it may be related to activity of the lateral premotor cortex found in fMRI studies of spatial orienting. The present EEG study extends these studies by delimiting the time period of this activity, suggesting that it reflects encoding of the spatial properties of the arrow for action.

CNV and temporal uncertainty with 'ageing' and 'non-ageing' S1-S2 intervals.

OBJECTIVES: Uncertainty about the timing of a known external event is an everyday phenomenon but has been rarely investigated with electrophysiological methods. We studied how the amplitude of the contingent negative variation (CNV) is affected by temporal variation of S2 presentation. Competing hypotheses about the development of CNV during the foreperiod until S2 presentation were that CNV either would follow a monotonic trend, be it increasing or decreasing, or alternatively that the time-course of CNV would be affected by the probability with which S2 was presented at each time-point in a given task. METHODS: The interval between cueing stimulus and imperative stimulus was randomly chosen from 3 different values between 1.3 and 2.6 s, using 3 different probability distributions in separate blocks: an 'ageing', a 'non-ageing' and a 'Gaussian' distribution. RESULTS: As previously shown, reaction times were determined by the probability of the imperative stimulus at the given length of the foreperiod. The same was found for CNV amplitude: the effects of temporal uncertainty on CNV mainly depended on the particular distribution of temporal probabilities used in a block. The relevant parameter was the a posteriori probability of event occurrence, very similar to the effects of this parameter on response times. In fact, the major part of the effect of a posteriori probability on CNV was common variation of CNV and response times. CONCLUSIONS: Thus, under temporal uncertainty the amplitude of CNV reflects the subjective expectancies for the occurrence of a given event, with this variation being related to variations in response times.

Lateralized EEG components with direction information for the preparation of saccades versus finger movements.

During preparation of horizontal saccades in humans, several lateralized (relative to saccade direction), event-related EEG components occur that have been interpreted as reflecting activity of frontal and parietal eye fields. We investigated to what degree these components are specific to saccade preparation. EEG lateralization was examined within the interval (1 s) between a first (S1) and a second (S2) stimulus, after which a response had to be made (look left or right, or press a button with the left or right index finger). The visual S1 indicated either the direction (left vs right) and/or the effector (eye vs finger), and S2 (visual/auditory in different blocks) added the information not given by S1. An occipital component (220 ms after S1) was effector-independent, probably reflecting processing of the direction code. The following parietotemporal component (320 ms after S1) was specific for direction information. This component seems more relevant for finger movements than for saccades and may reflect a link between visual perception to action. A later frontal component (480 ms after S1) was specific for direction information and may be related to the planning of a lateral movement. One component was entirely specific for the preparation of a finger movement (the lateralized readiness potential before S2). Thus, several different lateralized processes in the S1-S2 interval could be delineated, reflecting hand-specific preparation, processing of the direction code, and the coordination of perception and action, but no components were observed as being specific for saccade preparation.

Consequences of altered cerebellar input for the cortical regulation of motor coordination, as reflected in EEG potentials.

The cerebellum is certainly involved in fine coordination of movements, but has no efferences of its own to the muscles. Thus, it can exert its influence only via other cerebral areas that have those efferences. This study investigated in patients with cerebellar atrophy how cortical motor areas are affected by dysfunction of the cerebellum. The main question was whether the patients' slow cortical electroencephalogram (EEG) potentials during key-press preparation and execution would be generally altered or would be specifically altered when fine coordination was needed. In the coordination task, right- and left-hand keys had to be pressed simultaneously with different forces, under visual feedback. Control tasks were to press with both hands equally or with one hand only. The patients indeed had a performance deficit in the coordination task. Their cortical EEG potentials were already drastically reduced in the simple tasks, but were enhanced by the same amount as in healthy subjects when more coordination was needed. These results suggest that the cerebellum is not exclusively active in fine coordination, but is generally involved in any kind of preparatory and executive activity, whereas the motor cortex becomes more active with fine coordination. The role of the cerebellum might be to provide the motor cortex with information needed for coordinating movements. In cerebellar atrophy, this altered input may be sufficient for the motor cortex in controlling simple tasks, but not for complex ones.

Slow EEG potentials (contingent negative variation and post-imperative negative variation) in schizophrenia: their association to the present state and to Parkinsonian medication effects.

Between warning signal (S1) and imperative signal (S2), the EEG shifts negatively (contingent negative variation, CNV) reflecting preparation and expectancy. Reduced CNV and continued negativity after S2 (post-imperative negative variation, PINV) have been repeatedly found in schizophrenic patients and have been interpreted as a deficit in attentional processes (CNV) and as uncertainty about the correctness of one's own response to the S2 (PINV). Recent studies obtained a CNV reduction specifically at central sites but not at frontal ones. The present study investigated whether these alterations of slow negative potentials depend on present state of symptoms, on the particular task used, and on neuroleptic medication. Therefore, out-patients and in-patients were studied, two different S1-S2 tasks were used, and the control groups were healthy subjects and patients with Parkinson's disease. The central CNV reduction was stable across tasks and across in-patients and outpatients. Frontal CNV was reduced in in-patients but in only one of the two tasks in outpatients. The schizophrenic patients' enhanced PINV was larger contralaterally than ipsilaterally to the responding hand, correlated with medication, and occurred in similar way in patients with Parkinson's disease. Thus, the PINV increase might reflect the Parkinsonian side effects of the anti-psychotic medication. In contrast, the central CNV reduction appears as a stable marker of schizophrenia, the frontal CNV reduction as a state-dependent effect. The central CNV reduction might reflect impairment in forming stable stimulus-response associations, the relative frontal enhancement might reflect the out-patients' attempt at compensating that impairment.

Spatial S-R compatibility with centrally presented stimuli. An event-related asymmetry study on dimensional overlap.

Lateral presentation of relevant information facilitates manual responses if the side of relevant information corresponds to the side of the response. Recently, temporally overlapping EEG asymmetries over the central motor cortex and posterior sites were reported as a possible correlate of the sensory-motor integration of spatial information. The present study investigated whether sensory-motor integration of spatial information can occur with symbolic spatial information the same way as with laterally presented stimuli. The task required participants to respond to arrows (target stimuli), which were "flanked" (from above and below) by neutral stimuli or by other arrows (compatible or not). In Experiment 1, this task was compared to the same task with letters as stimuli and to an incompatible task where participants had to respond "against" the arrow direction. The effect of the flankers on response times was largest if subjects had to respond to the arrows in the common way. This was also the only task of Experiment 1 for which marked EEG asymmetries related to the direction of the flankers were observed. In Experiment 2, the onsets of target stimulus and flankers differed in time. Event-related lateralizations of the EEG over sensory and primary motor areas--as a lateralized readiness potential--were always, apparently automatically, evoked by flanking arrows, indicating automatic response activation evoked by symbolic spatial information. In accordance to recent theories of temporally decaying response activation, manual responses were affected only if the target was either shortly preceded by or appeared simultaneously with the flankers. The temporal overlap of EEG asymmetries related to direction encoding, automatic response activation, and to response preparation indicated that a widespread cortical network is activated by a salient directional information that enables subjects to respond quickly if the directional code of the stimulus overlaps with the directional code of the response.

Lateralized human cortical activity for shifting visuospatial attention and initiating saccades.

Lateralized human cortical activity for shifting visuospatial attention and initiating saccades. J. Neurophysiol. 80: 2900-2910, 1998. The relation between shifts of visual attention and saccade preparation was investigated by studying their electrophysiological correlates in human scalp-recorded electroencephalogram (EEG). Participants had to make saccades either to a saliently colored or to a gray circle, simultaneously presented in opposite visual hemifields, under different task instructions. EEG was measured within the short interval between stimulus onset and saccade, focusing on lateralized activity, contralateral either to the side of the relevant stimulus or to the direction of the saccade. Three components of lateralization were found: 1) activity contralateral to the relevant stimulus irrespective of saccade direction, peaking 250 ms after stimulus onset, largest above lateral parietal sites, 2) activity contralateral to the relevant stimulus if the stimulus was also the target of the saccade, largest 330-480 ms after stimulus onset, widespread over the scalp but with a focus again above lateral parietal sites, and 3) activity contralateral to saccade direction, beginning about 100 ms before the saccade, largest above mesial parietal sites, with some task-dependent fronto-central contribution. Because of their sensitivity to task variables, component 1 is interpreted as the shifting of attention to the relevant stimulus, component 2 is interpreted as reflecting the enhancement of the attentional shift if the relevant stimulus is also the saccade target, and component 3 is interpreted as the triggering signal for saccade execution. Thus human neurophysiological data provided evidence both for independent and interdependent processes of saccade preparation and shifts of visual attention.

Responses to cued signals in Parkinson's disease. Distinguishing between disorders of cognition and of activation.

Impairment of movement execution in Parkinson's disease could be due to disorders of cognition and/or of activation. These two factors are hard to separate by measuring response times only. Therefore, in this study response force and event-related EEG potentials were measured continuously during tasks in which subjects had to respond to cued signals. Fifteen patients with Parkinson's disease and 15 healthy subjects were studied during two tasks: (i) the 'clock task', in which the signal's identity was fully precued but its presentation time was uncertain and (ii) the 'validity task' in which the cue did not always predict the response validly. Thus, the clock task required more sustained attention, and the validity task sometimes required fast switching. The patients generally responded slower than control subjects. In the clock task, the response times of both groups changed to the same extent with presentation time, whereas in the validity task the patients were additionally slower than the control group with invalidly cued signals. The patients generally had a weaker response force and a lower rate of force production. In the clock task, both force measures changed with presentation time in the control group only, whereas in the validity task, the two measures increased in both groups to the same extent with invalidly cued signals. The contingent negative variation amplitudes in the patients' event-related EEG potentials were reduced, reflecting reduced activation of movement preparation, whereas lateralization of the motor cortices (i.e. the lateralized readiness potential) did not differ significantly between groups, reflecting unimpaired response selection. Force and contingent negative variation were generally reduced in the patients showing that their general slowing is at least partially due to impaired activation. Task-specific problems added to the general activation deficit; the lack of modulation of response force by presentation time revealed pronounced deficits of activation in the monotonous clock task. The specific delay of responses with invalidly cued signals, unparalleled by activation measures, might suggest a problem of cognition. The task-specific deficits may reflect a basic dilemma for patients with Parkinson's disease: cognitive problems may arise in complex tasks but disorders of activation may become pronounced in more simple, monotonous tasks.

Shifting attention between global features and small details: an event-related potential study.

In two experiments, large letters H or Z composed of small letters (also H or Z) were presented. Subjects had to make a two-choice motor response (e.g. H--left key, Z--right key). A cue presented 500 ms before the letter indicated which level (global or local) was relevant. In Experiment I, a third letter (T) sometimes appeared either at the cued or the non-cued level; in the former case, subjects had to shift their attention and to respond to stimulus features located at the non-cued level. The interference effect (RT delay in response to incongruent stimuli as compared to congruent ones) was larger when the local, rather than global, level was cued. A slow anterior negativity preceding globally-cued stimuli and shorter N1 and P2 ERP component latencies to these stimuli indicated better preparation for processing of global, as compared to local, stimulus features. The shift from local to global focus yielded a larger increase of RT, error rate, and of the P600 latency than the global-to-local shift. The P600 latency changes were parallel to those of RT. In Experiment II, the attentional shift was provoked by stimulus color red-colored letters meant that the cue was invalid, and thus, subjects had to respond to the non-cued level. Neither the interference nor the attentional shift demonstrated any asymmetry between the global and local levels. ERPs also did not differ substantially after local and global cues. In the condition demanding a shift of focus (invalid cue, incongruent letter), a positive deflection of the lateralized readiness potential indicated the activation of the wrong response channel. The large RT increment in this condition was not accompanied by an increase of the P600 latency. Two possible mechanisms of attentional shift may be proposed, the first related to perceptual processes (e.g. an additional visual search), and the second, to the competition between two response intentions.

Lateralised cortical activity due to preparation of saccades and finger movements: a comparative study.

The topography and time course of event-related asymmetries of the EEG associated with horizontal saccadic eye movements and finger movements was compared in a 4-choice response task, where the subjects had to respond to the imperative stimulus (S2) by moving the right or left index finger or by making a saccade to the right or the left. The cue stimulus (S1) contained full, partial, or no information about the direction and the effector. In case of finger movements 3 distinct lateralisations were found: (1) increased negativity over the motor cortex contralateral to the future movement direction, (2) increased contralateral negativity at temporoparietal sites beginning 200 ms after delivery of the directional information, and (3) increased ipsilateral negativity at temporo-parietal sites beginning 350-500 ms after delivered direction and effector information. The early temporo-parietal lateralisation was also visible in case of saccadic eye movements and in case of effector-unspecific directional information. Before saccadic eye movements no other distinct lateralisation could be observed at any recording site. In sum, lateralised cortical activities due to preparation processes for finger movements and due to effector-unspecific processing of directional information for motor preparation by the posterior parietal cortex could be demonstrated, whereas no distinct lateralisation due to preparation for saccadic eye movements was visible.

[EEG correlates of different methods of information processing in response selection]. / EEG-Korrelate unterschiedlicher Informationsverarbeitung zur Reaktionsauswahl.

The lateralized readiness potential as a measure of response tendencies has become a relevant method in experimental psychology within the last decade. This measure delivers information about response preparation long before the overt response. Applying the method of measuring event-related lateralizations (ERL) of the EEG to posterior sites, correlates of processing of spatial information and of response selection have been found. The present study investigated the topographical distribution of asymmetries at the moment of response selection. Response selection was assumed to be reflected in a discrete peak of parietal asymmetries. At this time point two distinct topographies were found that might reflect two different types of information processing. If response selection was based on spatial information a parieto-occipital maximum was found. In contrast, if centrally presented symbolic stimuli were used for response selection, lateralizations were largest at central (motor areas) and at parietal sites.

On-line brain potential correlates of right parietal patients' attentional deficit.

EEG potentials evoked by cues and targets were recorded in Posner's visual cueing task from 10 patients with lesions of the right parietal cortex and from age-matched healthy subjects. The patients' N1 component evoked by left-side cues was reduced at the right-parietal recording site, suggesting a general impairment in processing left-side visual input. As usual, patients' keypress responses were delayed when left targets were preceded by right cues. There were two correlates of this delay in the patients' EEG potentials evoked by the critical combination of right cue/left target: their mean amplitude 160-280 ms after target onset ('Nd') was less negative than with other combinations of cue and target, and the following frontal P300 was enhanced. The Nd reduction seems to be an on-line measure of patients' momentary decrease of attention for the left hemifield, while the frontal P300 might reflect the patients' attempts at reorienting. In conclusion, different components were sensitive to different aspects of the patients' disorder, suggesting the utility of this approach for developing detailed hypotheses on the mechanisms of attentional deficits involved in visual extinction and neglect.

The interaction of stimulus- and response-related processes measured by event-related lateralizations of the EEG.

The present study focused on the relationship between movement- and stimulus-related asymmetries of the electroencephalogram (EEG). In seven tasks the same bilateral stimuli containing asymmetric information were presented but response requirements differed. Three functionally distinct asymmetries were found: (1) an asymmetry over the motor cortex prior to unimanual movements, (2) an asymmetry over the posterior cortex beginning about 20 ms after the start of the movement, and (3) an early increase of negativity contralateral to a relevant stimulus (200-300 ms after stimulus onset) that was maximal at temporo-parietal sites but was also visible at central sites. Although related to stimulus side, this asymmetry was modulated by response requirements: it was largely abolished with simple responses, smaller with nogo than with Go stimuli and occurred twice when a sequence of simple and choice responses was required. Therefore, the early temporo-parietal asymmetry most probably reflects an interface between sensory and movement-related processes.

Preparation for action: an ERP study about two tasks provoking variability in response speed.

This study focused on the covariation of response speed and event-related potentials during response preparation and on whether these variations can be brought under experimental control. Two S1-S2 choice response tasks with temporal uncertainty were conducted. In Experiment 1, S1 was 100% informative. Fast subjects showed larger P3s with S1 than slow subjects. The terminal CNV (tCNV) increased intraindividually with response speed. In Experiment 2, 50% of S1s were uninformative and the visual display was designed to attract more attention. Effects of information were found on P3 amplitude, on the topography of tCNV, and on the temporal distribution of response times. Interindividual differences disappeared in Experiment 2. The results suggest that group differences in Experiment 1 were due to different strategies of allocating visual attention. Interindividual variations of strategy showed a pattern of effects different from intraindividual variations of efficiency.

Auditory selective attention is impaired in Parkinson's disease--event-related evidence from EEG potentials.

Selective attention refers to the ability to focus on one channel of information in the presence of distracting other channels. For the visual modality, results on impairments of selective attention have been conflicting in patients with idiopathic Parkinson's disease (PD). Independent of possible interferences from visual or movement disturbances selective attention can be measured as the so-called 'processing negativity' (PN) using auditory evoked potentials. Therefore, auditory selective attention with the PN was measured in 14 patients with PD and 16 control subjects. Subjects had to attend to tones presented to one ear (i.e. to press a button to occasionally presented longer tones) and ignore tones presented to the other ear. Tones were presented at a rate of 1/s ('slow') or 2/s ('fast'). PN was measured as the difference of the potentials evoked by attended minus ignored standard tones. PN was significantly smaller in the PD patients than in the controls with slow presentation. There was no difference between both groups with fast presentation. PN remained unchanged when patients had a 12-h withdrawal of their usual anti-Parkinsonian drug therapy. PD patients and controls did not differ in their P3 component evoked in the usual 'oddball' task nor in the mismatch negativity evoked by the occasionally longer tones in the PN task. The results provide evidence for an impairment of auditory selective attention that is specific for patients with PD (i.e. independent of the P3 component).

Response force and reaction time in a simple reaction task under time pressure.

The aim of this study was to examine the effect of effort on timing and force of simple reactions. To do this, we performed an experiment, arranged like a video-game, in which these variables were measured under different time-pressure conditions. In accordance with our expectations reaction time was shorter and force amplitude larger when the time for responses was limited. These findings are discussed within the framework of Sanders' (1983) model of stress.

Differences in P3 amplitudes between schizophrenics and healthy controls vary between the different events presented in a guessing task.

P3 amplitudes were often found to be reduced in schizophrenics, but in varying degree. We studied in a guessing task whether variations of group differences could even be obtained within the same task, by measuring P3 in the potentials evoked by the three qualitatively different events that occurred in each trial. We hypothesized that such variations were due to variations of subjective task relevance associated with each event. In each trial, a light or a sound was presented. Subjects had to confirm this 'primary event' by a delayed response, and then the current amount of money earned by correct responses was displayed. In the certain condition, the primary event was preceded by the appropriate cue on the screen ('light' or 'sound') whereas in the uncertain condition, the word 'guess' appeared. The task-relevance hypothesis predicted that schizophrenics would have smaller P3s than the control group with the primary event in the uncertain condition, but that the groups would not differ for the P3s evoked by the other events (certain primary event, cue and earnings display in uncertain and certain conditions). Essentially, this predicted pattern of results was obtained, but additional assumptions are needed for the task-relevance hypothesis to account for the precise pattern of results. Analysis of subjects' guessing behavior showed that control subjects' guesses were affected by the outcome of their preceding guesses while schizophrenics' guesses were not. This result provides some additional support for the hypothesis that the guessing task is less relevant to schizophrenics than to control subjects.