Error processing in patients with Parkinson's disease: the influence of medication state.

One of the hallmarks of Parkinson's disease (PD) is a depletion of dopamine. Error processing, as reflected in a component of the event-related potential, the so-called error (related) negativity (Ne or ERN) is likely dependent on the midbrain dopaminergic system. In case of an unfavourable event such as an error, this system is assumed to send an error signal to the mediofrontal cortex, which elicits the Ne. Hence, the Ne should be altered in patients with PD. In fact, we earlier found a reduction of the Ne in medicated patients with PD in different tasks while another group found no such reduction in "off-medication" patients in a flanker task. In the present study, we reinvestigated this issue by measuring the Ne in a large group of treated PD patients in the "on"- and "off"-parkinsonian medication state and in matched control subjects in a flanker task. The Ne was found to be the same in the "on-medication" and "off-medication" state, while the motor score in the Unified Parkinson's Disease Rating Scale was different. In both medication states the Ne was smaller in the patients than in the controls. The results show that the Ne reduction found earlier is unaffected by short-term differences in parkinsonian medication. The question remains open whether the long-term medication could have contributed to the Ne reduction.

Effects of stimulus-response compatibility in Parkinson's disease: a psychophysiological analysis.

The present study investigated the mechanisms underlying stimulus-response compatibility effects in Parkinson's disease patients and matched controls. Since basal ganglia are involved in the selection and inhibition of competing responses we examined whether basal ganglia dysfunction in Parkinson's disease leads to greater interference effects compared to the control subjects. Reaction times and lateralized movement-related cortical potentials (lateralized readiness potential: LRP) were recorded in two modified Eriksen flanker tasks. Both groups were influenced by compatibility conditions; interference was seen as enhanced reaction time and error rate, as well as incorrect early LRP and delayed late LRP in incongruent trials. Altogether, behavioral and electrophysiological measures showed the interference to be rather smaller for the patients than for the controls. In contrast, facilitation did not differ among groups. Hence the claim that Parkinson's disease patients are more influenced than controls by interfering directional stimuli appears not always valid.

Speed decrements are seen better than increments through small apertures.

Five subjects observed a random dot pattern that moved horizontally within a rectangular, 40 deg x 4 deg invisible aperture. The long side of this aperture was either parallel or perpendicular to the direction of motion. A two-interval forced-choice procedure was employed for measuring the thresholds for detection of 100-ms speed increments and decrements superimposed on uniform motion of 1.1 s duration and base speeds of 8 deg/s, 16 deg/s, 24 deg/s, and 32 deg/s. The aperture orientation had almost no effect on the detection of decrements. With increasing base speed, the thresholds for increments within the perpendicular aperture rose markedly as compared to those for increments within the parallel aperture and to those for decrements. The results are interpreted in the context of facilitatory interactions between the motion detectors along the trajectory of motion of each individual dot. It is shown how, having rapid motion within the perpendicular aperture, increment detection can severely deteriorate owing to the short lifetime and the small visible path of the dots, whereas decrement detection may be almost unaffected.

Changes of error-related ERPs with age.

Errors in reaction tasks are followed by a negative component of the event-related brain potential (ERP), the error negativity (Ne), which is thought to be a correlate of error detection. In the present study we show that, in tasks that induce different types of errors, the amplitude of the Ne was reduced in elderly (54-65 years old) compared with young subjects (19-25 years old). This reduction was also seen in single trials, as were computed for one of the visual tasks. Moreover, in this data set, the single-trial Ne was also delayed for the elderly compared with the young. These data suggest an alteration of error detection in the elderly, which is only marginally reflected in performance.

Action monitoring, error detection, and the basal ganglia: an ERP study.

The error negativity (Ne or ERN) is an event-related brain potential component, which is assumed to reflect error detection. Recently it has been hypothesized that the basal ganglia are assumed to play a crucial role in error detection. In the present study we ask whether the Ne is altered in patients with Parkinson's disease (PD), who have an impaired function of the basal ganglia. We recorded the Ne in patients and in matched controls, while they performed different tasks that require a relatively high cognitive control, which is supposed to pose particular problems on PD. The Ne was in fact smaller in the patients than in the controls in all tasks. Our results suggest an impairment of error detection in PD for different types of demanding tasks. This supports the hypothesis that the basal ganglia do play an important role for error detection in action monitoring.

Mechanisms of simple and choice reaction to changes in direction of visual motion.

Experiments are presented in which a random dot pattern moved vertically upwards (velocity vector V(1)) and then abruptly changed its direction of motion by the angle alpha (velocity vector V(2)), either to the left or to the right, without changing the speed. Subjects performed simple reactions to the direction change, disregarding its sign. In another experiment choice reactions to the same stimuli were performed: the subjects pushed a left button when the direction change was to the left and a right button when the change was to the right. The simple reaction time decreased monotonically with alpha increasing from 11 degrees to 169 degrees, whereas, within the same range of angles, a U-shaped curve described the function of the choice reaction time versus alpha. Both types of reaction time increased with decreasing the base speed. Difficulties are outlined which occur when the angle of change alpha is considered as 'intensity' of the stimulus. Instead, the parameter mid R:V(2)-V(1)mid R:, the absolute value of the difference between the velocity vectors before and after the change, is shown to be a meaningful 'intensity' parameter for the simple reaction task. The parameter V(2N), the speed of the velocity component normal to the initial velocity vector V(1), is suggested as an 'intensity' parameter for the choice reaction task. It is shown that the simple and choice reactions to changes in direction of visual motion are performed by two distinct mechanisms which seem to work in parallel and may be nearly equally fast for small angles of change, when mid R:V(2)-V(1)mid R: approximately V(2N).

The discrimination of abrupt changes in speed and direction of visual motion.

A random dot pattern that moved within an invisible aperture was used to present two motions contiguously in time. The motions differed slightly either in speed (Experiments 1 and 3) or in direction (Experiments 2 and 4) and the subject had to discriminate the sign of the change (e.g. increment or decrement). The same discrimination task was performed when the two motions were temporally separated by 1 s. In Experiments 1 and 2 discrimination thresholds were measured with motion durations of 0.125, 0.25, 0.5 and 1.0 s and mean speeds of 2, 4, 8, and 16 degrees/s. In Experiments 3 and 4 thresholds were measured with aperture widths of 5 and 20 cm. The discrimination of contiguous motions progressively deteriorated with decreasing duration and mean speed of motion. For the lowest value of duration the Weber fraction for contiguous speeds was more than three times as the Weber fractions for separate speeds. For the same low value of duration the thresholds for discrimination of direction of contiguous motions were only about 50% higher than the thresholds for separate motions. The Weber fraction for contiguous speeds was ca. three times higher with the smaller aperture than with the larger one, provided the ratio 'aperture width mean speed' (i.e. the lifetime of the moving dots) was less than 0.3 s. Aperture width did not affect the discrimination of direction of contiguous motions. The discrimination of contiguous motions is discussed together with the known data for detection of changes in speed and direction. It is suggested that both, detection of changes in speed and discrimination of the sign of speed changes, may be performed by a common visual mechanism.

Early attention effects in human auditory-evoked potentials.

A fundamental question in attention theory concerns the earliest processing stages that can be modulated by selective attention. A series of experiments is reported in which very early attention effects are found under specific conditions in the frequency-following potential (FFP), a brain stem response to low-frequency tone stimuli. In two experiments, stimuli of two different modalities were applied, and attention directed to one of the modalities. In two further experiments, only auditory stimuli were presented. In the first of these last two experiments, a dichotic paradigm with sustained attention to one ear was used, in the second a monotic paired-stimuli paradigm was used, in which the first stimulus served as reference for the second one. Only in the last experiment significant attention effects were found in the latency, but not in the amplitude of the FFP. The results show that a very early attention effect on the latency of the FFP can be demonstrated, but only under highly specific conditions. The size and preconditions of the attention effect suggest that it reflects subtle intramodal tuning mechanisms in the cochlea or in the lower brain stem.

ERP components on reaction errors and their functional significance: a tutorial.

Some years ago we described a negative (Ne) and a later positive (Pe) deflection in the event-related brain potentials (ERPs) of incorrect choice reactions [Falkenstein, M., Hohnsbein, J., Hoormann, J., Blanke, L., 1990. In: Brunia, C.H.M., Gaillard, A.W.K., Kok, A. (Eds.), Psychophysiological Brain Research. Tilburg Univesity Press, Tilburg, pp. 192-195. Falkenstein, M., Hohnsbein, J., Hoormann, J., 1991. Electroencephalography and Clinical Neurophysiology, 78, 447-455]. Originally we assumed the Ne to represent a correlate of error detection in the sense of a mismatch signal when representations of the actual response and the required response are compared. This hypothesis was supported by the results of a variety of experiments from our own laboratory and that of Coles [Gehring, W. J., Goss, B., Coles, M.G.H., Meyer, D.E., Donchin, E., 1993. Psychological Science 4, 385-390. Bernstein, P.S., Scheffers, M.K., Coles, M.G.H., 1995. Journal of Experimental Psychology: Human Perception and Performance 21, 1312-1322. Scheffers, M.K., Coles, M. G.H., Bernstein, P., Gehring, W.J., Donchin, E., 1996. Psychophysiology 33, 42-54]. However, new data from our laboratory and that of Vidal et al. [Vidal, F., Hasbroucq, T., Bonnet, M., 1999. Biological Psychology, 2000] revealed a small negativity similar to the Ne also after correct responses. Since the above mentioned comparison process is also required after correct responses it is conceivable that the Ne reflects this comparison process itself rather than its outcome. As to the Pe, our results suggest that this is a further error-specific component, which is independent of the Ne, and hence associated with a later aspect of error processing or post-error processing. Our new results with different age groups argue against the hypotheses that the Pe reflects conscious error processing or the post-error adjustment of response strategies. Further research is necessary to specify the functional significance of the Pe.

Different error types and error processing in spatial stimulus-response-compatibility tasks: behavioural and electrophysiological data.

We tested the hypothesis that in spatial stimulus-response-compatibility (SRC) tasks two different error types occur: A noise-induced 'general error' independent of SRC and reaction time and a 'position driven error' in incompatible trials with short RT being driven by the irrelevant stimulus position. A second issue was whether error detection is different for these two types of errors, which should be reflected by differences in the error negativity (Ne), since the Ne is seen as a neural correlate of error detection. To study these issues, we used a Simon- and a spatial Stroop-task. In incompatible (vs. compatible) trials we found more errors and a below chance accuracy in fast responses. Neither the amplitude nor the latency of the Ne were significantly affected by the experimental factors. This pattern of behavioural results supports the above hypothesis of two error types in such tasks. The Ne results indicate that error detection is similar for both types of errors.

ERP components in Go/Nogo tasks and their relation to inhibition.

In visual Go/Nogo tasks the ERP usually shows a frontal negativity after Nogo stimuli ("Nogo-N2"), which possibly reflects an inhibition process. However, the Nogo-N2 appears to be very small after auditory stimuli, which is evidence against the inhibition hypothesis. In the present study we tested this hypothesis by evaluating performance differences between subjects. Assuming that for Ss with a high false alarm rate the inhibition process is weakened and/or delayed, they should reveal a smaller and/or later Nogo-N2 than Ss with a low false alarm rate. This prediction was confirmed, which supports the inhibition hypothesis. However, the Nogo-N2 was again much smaller and had a different topography after auditory than after visual stimuli despite similar performance in both modalities. This modality asymmetry was explained by assuming that the inhibitory mechanism reflected in the Nogo-N2 is located at a pre-motor rather than at the motor level. In the second part of the study we compared the Nogo-N2 with a similar phenomenon, the error negativity (Ne), which occurs in trials with commission errors (false alarms). Earlier work suggests that the Ne is a correlate of error detection or inhibition. This raises the possibility that the Ne is a delayed Nogo-N2, i.e., the Ne may reflect a late and hence unsuccessful attempt to inhibit the response after a nontarget. However, the Ne amplitude showed no difference between performance groups and stimulus modalities, as found for the Nogo-N2. Moreover, Ne and Nogo-N2 had different scalp topographies. This suggests that different mechanisms and generators underlie the Ne and the Nogo-N2.

The simple reaction time to changes in direction of visual motion.

Recently Dzhafarov et al. presented a model explaining data on simple reaction time (RT) to unidimensional velocity changes. The authors suggested that having a motion with an initial velocity V0, the velocity change detection system is reinitialized by means of a "subtractive normalization" process. Therefore, any abrupt change from V0 to V1 is detected as if it were the onset of motion with a speed equal to /V1-V0/. They derived that the RT is a function of /V1-V0/(-2/3). We tested this model for the case of two-dimensional velocity changes. Our subjects observed a random dot pattern that moved horizontally, then changed the direction of motion by an angle alpha in the range between 6 degrees and 180 degrees without changing the speed V. Speeds of 4 and 12 deg/s were used. The subjects reacted as quickly as possible to the direction change. The RTs asymptotically decreased with increasing alpha; with 12 deg/s speed the RTs were shorter than those obtained with 4 deg/s. It was shown that the data can be well described as a function of /V1-V0/(-2/3)=(2Vsin(alpha/2))(-2/3). An extension of the "subtractive normalization" hypothesis for the case of two-dimensional velocity changes is proposed. It is based on the assumption that the velocity vector V1 after the change is decomposed into two orthogonal components. Alternative explanations based on the use of position or orientation cues are shown to contradict the data.

Performance differences in reaction tasks are reflected in event-related brain potentials (ERPs).

Event-related potentials (ERPs), which can be extracted from the electroencephalogram (EEG), are assumed to reflect distinct cognitive processes in real time. Hence ERP analysis could be used in cognitive ergonomics as a tool to specify, for example, bottlenecks or sources of individual performance differences. Such specific results may be helpful to change the tasks or train the subjects specifically. In the present exploratory study, the authors investigated whether subjects with large spontaneous differences in performance accuracy, as defined by their error rates in a speeded binary choice reaction task, also differ in the structure of their ERPs. The ten subjects were divided post hoc into two groups with relatively low (about 6%) and high (about 20%) error rates. While the reaction times were not significantly different for both groups, the ERPs revealed clear group differences. First, large differences were seen in the late part of the contingent negative variation (late CNV), which is assumed to reflect preparatory processes. Subjects with few errors ('GOOD') had a large late CNV, while subjects with many errors ('POOR') showed virtually no late CNV. Second, the late P300-subcomponent (which is related to response identification) was smaller and delayed for POOR compared to GOOD subjects. Finally, the ERP shows signs of poor movement control in POOR subjects. The high error rate of POOR subjects can hence be explained by: (1) their insufficient preparation for the next trial (small late CNV), which impaired response identification (small and delayed late P300 subcomponent); and (2) their poor movement control. These interpretations have to be regarded as preliminary and should be validated with larger groups of subjects. In conclusion, the main reasons for the profound performance differences between the groups, namely differential preparation and movement control, could be elucidated by ERP analysis. A potential ergonomics application of these results is that they suggest specific strategies (for example, a preparation and motor control training) to improve the performance of POOR subjects in comparable work conditions.

The time it takes to detect changes in speed and direction of visual motion.

We studied the ability of human observers to detect abrupt changes in velocity of motion of a random dot pattern. The pattern moved horizontally for 0.9 s at velocity V0, then changed to V1 either in speed, or in direction for a time T and returned to the initial motion. The threshold duration for detection of the change was measured for initial speeds of 2, 4, 8 and 16 deg/s. The time to detect a velocity reversal was equal to that for detection of an increase in speed by a factor of three. The time to detect an abrupt cessation of motion was equal to the time for detection of an increase in speed by a factor of two. The time to detect a direction change, the speed being constant, decreased gradually with increasing angle between V0 and V1 from 12 to 180 degrees and with increasing V0; the detection time was a function of (V1-V0) almost independent of the value of V0. This finding supports the hypothesis of Dzhafarov et al. (Percept Psychophys 1993;54:373-750), that the visual system effectively reduces the detection of velocity changes (from V0 to V1) to the presumably more simple detection of a motion onset, from 0 to (V1-V0). The characteristics of the detection process in the cases of uni- and two-dimensional velocity changes are discussed.

A method to improve the latency estimation of the frequency-following potential (FFP).

If the latency of a noisy frequency-following potential (FFP) is estimated by determining the shift of the (periodical) cross-correlation function (CCF) between the stimulus and the FFP, the result may be unambiguous only within +/-1 or +/-2 periods of the CCF, because the absolute maximum and adjacent local maxima may not be significantly different. Here we present a method to amplify this difference by applying amplitude modulated stimuli. Using this method we first illustrate the effect of the method by a simulation and then demonstrate its usefulness by measuring real FFPs and estimating their latencies.

[Event-related potential components related to errors]. / Fehlerbezogene Komponenten im ereigniskorrelierten Potential (EKP).

Event-related potentials (ERPs) of error trials in choice tasks and Go/Nogo tasks are found to be considerably different from the ERPs of the correct trials: In error trial ERPs there is an additional negative (Ne) and an additional positive component (Pe) compared to correct trials. Amplitude and latency variation of both components in different experiments supports the hypothesis that these components reflect different aspects of error processing. The Ne is interpreted as a real-time correlate of error detection, as defined by a mismatch between cognitive representations of the erroneous response and the correct response. The variation of Pe with experimental variables is different from that of the Ne and it also from that of positive components in correct trials, and may therefore reflect an additional aspect of error processing, such as change of response strategies.

What determines the detection of changes in motion velocity? A comment on Dzhafarov, Sekuler, and Allik (1993)

We comment on a recent model aimed at explaining data on speed of reaction to motion onset and to changes in motion velocity. The model is based on calculating the running variance of the stimulus positions passed during the motion. We show that although the model is successful in explaining data on motion onset and suprathreshold velocity changes, it may not be able to explain data on time of reaction to changes in velocity when these are near the detection threshold.

Perception of visual motion with modulated velocity: effects of viewing distance and aperture size.

Subjects observed a random dot pattern that moved horizontally with modulated velocity within an invisible aperture. The velocity contrast, (V2-V1)/V1, was 2/3. Two different percepts occurred while observing this stimulus. At lower modulation frequencies, between 2 and 12 Hz, velocity changes were clearly seen; this percept is called "motion irregularity". At frequencies higher than 20 Hz velocity changes were no longer visible; the moving pattern appeared to be divided into stationary columns of different luminance. We call this percept "pattern irregularity". The critical frequency for detection of motion irregularity was independent of viewing distance; it was an inverted U-shaped function of the linear rather than the angular mean velocity of the pattern. At higher mean velocities the critical frequency increased with increasing aperture size; at lower mean velocities it was not affected by the size of the aperture. It is shown that detection performance is a function of the relative velocity of the pattern, i.e. of the ratio between the mean velocity in deg/sec and the aperture size in deg. Pattern irregularity could be detected at modulation frequencies even above 100 Hz. The critical frequency increased with increasing velocity and with decreasing viewing distance. It is suggested that detection of motion irregularity is determined by two distinct processes that are based on spatial analysis of motion at low relative velocities and temporal analysis at high relative velocities; both processes provide constancy of detection performance regardless of viewing distance. On the other hand, pattern irregularity seems to be detected on the basis of an analysis of the retinal luminance distribution at high modulation frequencies.

Effects of attention and time-pressure on P300 subcomponents and implications for mental workload research.

Our approach to objective measures of mental workload is establishing relationships between components of the event-related brain potential (ERP) and information processing stages. These relationships can be used to infer the influence of specific workload conditions on specific processing stages. We recently showed that the ERP component P300 in choice tasks is composed of two subcomponents, P-SR and P-CR, which are time-related to stimulus-evaluation and response-selection. With these relations we could specify which processing stages were affected when certain workload conditions are varied. When attention was divided between the visual and auditory modalities compared to (unimodal) focused attention, the choice reaction time (RT) was prolonged, primarily in the auditory modality. This delay was mainly reflected in the P-CR latency, which shows that the division of attention mainly impairs the response-selection process in the auditory modality due to a bias of attention towards the visual modality. When the time-pressure was increased, the latency of the P-CR (and not of the P-SR) was shortened, but less than the choice RT. This suggests a (limited) acceleration of response-selection but not of stimulus evaluation. Since the response-selection process was accelerated less than the overt choice RT, an increase of the error rate was consequently observed. In summary we showed that increases of mental workload can induce accelerations or decelerations of specific processing stages which can be monitored by observing latency changes of the affiliated ERP components.

Temporal thresholds and reaction time to changes in velocity of visual motion.

A random dot pattern moved at a velocity V1. The velocity then increased or decreased abruptly to another value V2 for some time and again returned to V1. The temporal threshold, i.e. the duration of V2 that was necessary to detect the change was measured. Thresholds for the detection of the same velocity increment, V2 = 2 x V1, were shorter when the baseline velocity V1 increased from 1 to 8 deg/sec (Expt 1). The temporal threshold decreased as the velocity contrast (V2 - V1)/(V1 + V2) increased from 0.33 to 0.77. The thresholds for the detection of velocity decrements were in general longer than those for the detection of increments (Expt 3). In Expts 2 and 4 the random-dot pattern moved with velocity V1, which abruptly increased or decreased to V2, without returning to V1. The reaction time to the change was measured for the same velocity pairs as those used in the temporal threshold measurements. There was a good correspondence between changes in the reaction times and changes in the thresholds under the various conditions. The data are interpreted on the basis of two hypotheses: higher velocities are detected by mechanisms that respond more rapidly; and integration of velocities occurs when temporally-adjacent motions are presented.