Executive control and response expectancy: A Laplacian ERP study

The event‐related potential called “Error Negativity” (Ne, ERN), which appears when subjects commit errors in choice reaction time tasks, is a marker of response monitoring. By introducing a response probability bias, we show that the Ne is sensitive to response expectancy. We further show that the small negativity evoked by correct responses (Ne‐like, CRN) is also sensitive to response expectancy: On unexpected responses, the former decreases while the latter increases to such an extent that the amplitudes of the two components are in the same range of magnitude. Although the sensitivity of the Ne to response expectancy is compatible with the current models accounting for the Ne, the common sensitivity of the Ne and the Ne‐like supports the idea that they reflect functionally similar monitoring processes.     

Motor inhibition and response expectancy: A Laplacian ERP study

In between-hand choice reaction time tasks, the motor cortex involved in the required response (contralateral) has been shown to be activated while the motor cortex involved in the non-required response (ipsilateral) has been shown to be inhibited. The aim of the present study was to test the hypothesis that ipsilateral inhibition reflects an active mechanism aimed at preventing errors. To this end, the risk of committing errors in between-hand choice reaction time tasks was manipulated by introducing a response probability bias. The surface Laplacian transforms of electroencephalographic waves recorded over the motor cortices, contralateral and ipsilateral to the responding hand were compared. Results showed that contralateral activation was not modulated by the risk of committing errors while ipsilateral inhibition was sensitive to this risk in a gradual manner: the higher the risk, the stronger the inhibition.     

Neural inhibition and interhemispheric connections in two-choice reaction time: A Laplacian ERP study

In between-hand choice reaction time tasks, the motor cortex involved in the required response is activated while the motor cortex involved in the non-required response is inhibited. Such an inhibition could be implemented actively between the responses defined as possible alternatives by the task instructions or, alternatively, could passively result from some kind of "reciprocal inhibition" between the two motor cortices. The present study addressed this issue. To this end, we compared the surface Laplacian transforms of electroencephalographic (EEG) waves recorded over the contralateral and ipsilateral motor cortices in between-hand and within-hand choice conditions. The dynamics of the recorded EEG activities suggest that inhibition is implemented in a feed-forward manner between the cortical zones controlling the different response alternatives rather than between homologous motor cortical structures.     

Neural activation in cognitive motor processes: comparing motor imagery and observation of gymnastic movements

The simulation concept suggested by Jeannerod (Neuroimage 14:S103-S109, 2001) defines the S-states of action observation and mental simulation of action as action-related mental states lacking overt execution. Within this framework, similarities and neural overlap between S-states and overt execution are interpreted as providing the common basis for the motor representations implemented within the motor system. The present brain imaging study compared activation overlap and differential activation during mental simulation (motor imagery) with that while observing gymnastic movements. The fMRI conjunction analysis revealed overlapping activation for both S-states in primary motor cortex, premotor cortex, and the supplementary motor area as well as in the intraparietal sulcus, cerebellar hemispheres, and parts of the basal ganglia. A direct contrast between the motor imagery and observation conditions revealed stronger activation for imagery in the posterior insula and the anterior cingulate gyrus. The hippocampus, the superior parietal lobe, and the cerebellar areas were differentially activated in the observation condition. In general, these data corroborate the concept of action-related S-states because of the high overlap in core motor as well as in motor-related areas. We argue that differential activity between S-states relates to task-specific and modal information processing.     

Functional brain changes in early Parkinson's disease during motor response and motor inhibition

Motor impairment represents the main clinical feature of Parkinson's disease (PD). Cognitive deficits are also frequently observed in patients with PD, with a prominent involvement of executive functions and visuo-spatial abilities. We used event-related functional MRI (fMRI) and a paradigm based on visual attention and motor inhibition (Go/NoGO-task) to investigate brain activations in 13 patients with early PD in comparison with 11 healthy controls. The two groups did not report behavioural differences in task performance. During motor inhibition (NoGO-effect), PD patients compared to controls showed an increased activation in the prefrontal cortex and in the basal ganglia. They also showed a reduced and less coherent hemodynamic response in the occipital cortex. These results indicate that specific cortico-subcortical functional changes, involving not only the fronto-striatal network but also the temporal-occipital cortex, are already present in patients with early PD and no clinical evidence of cognitive impairment. We discuss our findings in terms of compensatory mechanisms (fronto-striatal changes) and preclinical signs of visuo-perceptual deficits and visual hallucinations.     

Response inhibition in adolescent earthquake survivors with and without posttraumatic stress disorder: A combined behavioral and ERP study

The aim of this study was to investigate whether adolescent patients with posttraumatic stress disorder (PTSD) show an impairment of executive control in a response inhibition task and to investigate its neurophysiological correlates using event-related potentials (ERPs). We analyzed data from 25 Wenchuan earthquake survivors between 15 and 19 years of age (16 diagnosed with PTSD) using a Go/NoGo task. The PTSD group made more commission errors than the non-PTSD group, indicating impairment in response inhibition. The PTSD group responded faster to Go trials and there was a significant negative correlation between their reaction time and commission/omission errors, reflecting a speed-accuracy tradeoff for the PTSD group. The PTSD group exhibited a shorter NoGo-N2 latency than the non-PTSD group, suggesting faster monitoring or detection of the response conflict. These results suggest that the impairment of response inhibition in adolescent participants with PTSD is related to their impulsive cognitive functioning.     

Influence of acute stress on response inhibition in healthy men: An ERP study

The current study investigated the influence of acute stress and the resulting cortisol increase on response inhibition and its underlying cortical processes, using EEG. Before and after an acute stressor or a control condition, 39 healthy men performed a go/no‐go task while ERPs (N2, P3), reaction times, errors, and salivary cortisol were measured. Acute stress impaired neither accuracy nor reaction times, but differentially affected the neural correlates of response inhibition; namely, stress led to enhanced amplitudes of the N2 difference waves (N2d, no‐go minus go), indicating enhanced response inhibition and conflict monitoring. Moreover, participants responding to the stressor with an acute substantial rise in cortisol (high cortisol responders) showed reduced amplitudes of the P3 of the difference waves (P3d, no‐go minus go) after the stressor, indicating an impaired evaluation and finalization of the inhibitory process. Our findings indicate that stress leads to a reallocation of cognitive resources to the neural subprocesses of inhibitory control, strengthening premotor response inhibition and the detection of response conflict, while concurrently diminishing the subsequent finalization process within the stream of processing.     

Individual differences in decisiveness: ERP correlates of response inhibition and error monitoring

The aim of the current study was to examine whether and how self‐reported decisiveness is associated with response inhibition and performance monitoring. We hypothesized that these two cognitive control mechanisms, both of which are often associated with decision making, would differ in individuals varying in decisiveness. We focused on ERP correlates and behavioral measures of inhibition and error processing in the stop‐signal task. We expected a negative relationship between decisiveness and behavioral measures of inhibitory control. We also hypothesized that stop‐signal‐locked N1 and P3 components and response‐locked error‐related negativity (ERN) would be less pronounced when participants self‐reported higher levels of decisiveness. Correlation analysis identified an association between high decisiveness, long stop‐signal reaction time, and low inhibition rate. Analysis with mixed‐effects linear models revealed that stop signals evoked less pronounced N1 and P3 in individuals scoring higher on decisiveness in both successfully and unsuccessfully inhibited trials. Additionally, high decisiveness was linked to reduced error monitoring, as indicated by decreased ERNs. Importantly, we also found positive association between P3 onset latency and decisiveness, suggesting that individuals scoring higher on this measure have relatively less ability to rapidly engage the stopping process. Thus, our findings primarily indicate that decisiveness is negatively associated with the efficiency of both response inhibition and error monitoring. They also suggest that highly decisive people may share some characteristics of diminished cognitive control with impulsive individuals.     

Programming the duration of a motor sequence: role of the primary and supplementary motor areas in man

Event-related potentials were recorded in a reaction time (RT) paradigm, where the duration of a learned interval (either 0.7 s or 2.5 s) delimited by two brief button-presses was to be accurately controlled. A preparatory signal (PS) either did not give or gave prior information concerning the duration of the following response (neutral condition or primed conditions, respectively). In the latter case, the information was either validated (valid condition) or invalidated (invalid condition) by the response signal (RS). When duration was not known in advance (invalid and neutral conditions), RTs were longer before a response of short than long duration. This difference was not found under the valid condition. During the preparatory period (PP), the amplitude of the contingent negative variation (CNV) was larger when the duration was primed than when it was not. A larger CNV appeared when the PS primed a short rather than a long duration. This effect occurred in the early part of the PP over the supplementary motor area (SMA) and in its latest part over the primary motor area (MI). The RT and the electrophysiological pattern were interpreted as revealing the occurrence of programming operations regarding the temporal dimension of the response. The time course of the CNV over the SMA and MI suggested that these two areas were hierarchically organized. Between the RS and the onset of the response, differences probably related to programming effects were still found over MI: the activities were larger under the valid than under the neutral condition. However, no sign of deprogramming (expected in the invalid condition) was observed: similar amplitudes were found under the neutral and invalid conditions. Deprogramming operations seemed to be postponed during response execution where the invalid condition evoked larger activities than the two other conditions over the SMA. Finally, MI but not the SMA yielded a Bereitschaftpotential before the second press ending the response (i.e., during response execution). These results suggest that the duration of a motor response can be a part of the motor program and that the SMA plays a major role in programming processes but not in response execution, contrary to MI.     

Stop-signal response inhibition in schizophrenia: behavioural, event-related potential and functional neuroimaging data

Inhibitory control deficits are well documented in schizophrenia, supported by impairment in an established measure of response inhibition, the stop-signal reaction time (SSRT). We investigated the neural basis of this impairment by comparing schizophrenia patients and controls matched for age, sex and education on behavioural, functional magnetic resonance imaging (fMRI) and event-related potential (ERP) indices of stop-signal task performance. Compared to controls, patients exhibited slower SSRT and reduced right inferior frontal gyrus (rIFG) activation, but rIFG activation correlated with SSRT in both groups. Go stimulus and stop-signal ERP components (N1/P3) were smaller in patients, but the peak latencies of stop-signal N1 and P3 were also delayed in patients, indicating impairment early in stop-signal processing. Additionally, response-locked lateralised readiness potentials indicated response preparation was prolonged in patients. An inability to engage rIFG may predicate slowed inhibition in patients, however multiple spatiotemporal irregularities in the networks underpinning stop-signal task performance may contribute to this deficit.     

Premotor and supplementary motor cortex in rhesus monkeys: neuronal activity during externally- and internally-instructed motor tasks

Summary We compared neuronal activity in the premotor (PM) and supplementary motor cortex (SM) of two rhesus monkeys as they performed two tasks. In an externally-instructed task, a visuospatial instruction stimulus indicated which of two touch pads should be the target of a forelimb movement. In an internally-instructed task, the visuospatial stimulus was either irrelevant or not presented, but in either case the target alternated between the two touch pads in blocks of 20 trials each. In both tasks, the monkey withheld movement for a self-timed delay period. Neuronal activity modulation during the delay period (set-related activity) and immediately before movement (movement-related activity) was comparable in PM and SM, both in terms of the proportion of cells with both of those activity patterns and their depth of modulation. Thus, our findings do not provide strong support for a clear-cut functional division between PM and SM regarding the control of externally- and internally-instructed limb movements. Within PM, 57 out of 96 cells with set-related activity showed similar modulation during the two tasks, supporting the proposition that such activity contributes to the preparation for a limb movement. In 32 of the 39 PM set-related neurons that showed a significant activity difference between the two tasks, activity was greater in the externally-instructed task. This finding supports the hypothesis that set-related activity in PM contributes more to sensorially-instructed than to other movements.     

Corticostriatal projections from the somatic motor areas of the frontal cortex in the macaque monkey: segregation versus overlap of input zones from the primary motor cortex, the supplementary motor area, and the premotor cortex

It is an important issue to address the mode of information processing in the somatic motor circuit linking the frontal cortex and the basal ganglia. In the present study, we investigated the extent to which corticostriatal input zones from the primary motor cortex (MI), the supplementary motor area (SMA), and the premotor cortex (PM) of the macaque monkey might overlap in the putamen. Intracortical microstimulation was performed to map the MI, SMA, and dorsal (PMd) and ventral (PMv) divisions of the PM. Then, two different anterograde tracers were injected separately into somatotopically corresponding regions of two given areas of the MI, SMA, PMd, and PMv. With respect to the PMd and PMv, tracer injections were centered on their forelimb representations. Corticostriatal input zones from hindlimb, forelimb, and orofacial representations of the MI and SMA were, in this order, arranged from dorsal to ventral within the putamen. Dense input zones from the MI were located predominantly in the lateral aspect of the putamen, whereas those from the SMA were in the medial aspect of the putamen. On the other hand, corticostriatal inputs from forelimb representations of the PMd and PMv were distributed mainly in the dorsomedial sector of the putamen. Thus, the corticostriatal input zones from the MI and SMA were considerably segregated though partly overlapped in the mediolateral central aspect of the putamen, while the corticostriatal input zone from the PM largely overlapped that from the SMA, but not from the MI. Received: 30 June 1997 / Accepted: 2 October 1997     

Response- and Stimulus-Related ERP Asymmetries in a Tonal Oddball Task: A Laplacian Analysis

Previous studies have found greater P3 amplitude over right than left hemisphere sites in a tonal oddball task with a reaction time (RT) response. This asymmetry had a central topography, and interacted with response hand. Identification of the processes underlying these asymmetries requires the use of additional methods for separating response- and stimulus-related contributions. We applied local Hjorth and spherical spline algorithms to compute surface Laplacian topographies of ERP data recorded from 30 scalp electrodes in a pooled sample of 46 right-handed healthy adults. For both methods, the current sources underlying the late positive complex were largest at medial parietal regions, but were asymmetric at central and frontocentral sites. Although a frontocentral sink contralateral to the response hand contributed to the asymmetry of the classic P3 peak, the source asymmetry was most robust after the sink had resolved. The late source was largest at electrode C4 for right hand responses, and was further enhanced in subjects showing a dichotic left ear advantage, but was unrelated to response speed. We conclude that the right hemisphere source reflects an interaction of response-related asymmetries with right hemisphere processes responsible for pitch discrimination.     

Electrophysiological correlates of short-latency afferent inhibition: a combined EEG and TMS study

Cutaneous stimulation produces short-latency afferent inhibition (SAI) of motor-evoked potentials (MEPs) elicited by transcranial magnetic stimulation (TMS). Since the demonstration of SAI is primarily based on the attenuation of MEPs, its cortical origin is not yet fully understood. In the present study we combined TMS with concurrent electroencephalography (EEG) in order to obtain direct cortical correlates of SAI. TMS-evoked EEG responses and MEPs were analysed with and without preceding electrical stimulation of the index finger cutaneous afferents in ten healthy volunteers. We show that the attenuation of MEPs by cutaneous stimulation has its counterpart in the attenuation of the N100 EEG response. Moreover, the attenuation of the cortical N100 component correlated positively with the strength of SAI, indicating that the transient changes in cortical excitability can be reflected in the amplitude dynamics of MEPs. We hypothesize that the hyperpolarization of the pyramidal cells due to SAI lowers the capacity of TMS to induce the inhibitory current needed to elicit N100, thus leading to its attenuation. We suggest that the observed interaction of two inhibitory processes, SAI and N100, provides further evidence for the cortical origin of SAI. R. Bikmullina and D. Kičić equally contributed to the study.     

Inhibitory motor control in apneic and insomniac patients: a stop task study

The aim of this study was to assess with a stop task the inhibitory motor control efficiency--a major component of executive control functions--in patients suffering from sleep disorders. Twenty-two patients with untreated obstructive sleep apnea syndrome (OSAS) (mean age 46 +/- 9 years; mean apnea-hypopnea index, AHI = 30 +/- 20) and 13 patients with psychophysiological insomnia (mean age 47 +/- 12 years) were compared with individually matched healthy controls. Sleep disturbances in the patient populations were clinically and polysomnographically diagnosed. The stop task has a frequent visual 'Go' stimulus to set up a response tendency and a less frequent auditory 'Stop' signal to withhold the planned or prepotent response. The stop signal reaction time (SSRT) reflects the time to internally suppress the ongoing response. SSRT was slower for the apneic patients than for their respective controls (248 +/- 107 versus 171 +/- 115 ms, anova, P < 0.05) but not for the insomniac patients compared with their controls (235 +/- 112 versus 194 +/- 109 ms, NS). Moreover, in apneic patients, slower SSRT was associated with lower nocturnal oxygen saturation (r = -0.477, P < 0.05). By contrast, neither apneics nor insomniacs differed from their matched controls for reaction times on Go trials. To conclude, unlike insomniacs, OSAS patients present an impaired inhibitory motor control, an executive function which is required in many common everyday life situations. Inhibitory motor control relies on the integrity of the inferior prefrontal cortex, which could be affected by nocturnal oxyhemoglobin desaturation in apneic patients.     

Further evidence for excitability changes in human primary motor cortex during ipsilateral voluntary contractions

The present study aimed to further investigate whether the intracortical neural circuits within the primary motor cortex (M1) are modulated during ipsilateral voluntary finger movements. Single- and paired-pulse (interstimulus intervals, ISIs; 3 ms and 12 ms) transcranial magnetic stimulations of the left M1 were applied to elicit motor evoked potential (MEP) in the right first dorsal interosseous (Rt-FDI) muscle during voluntary contractions (10% and 30% maximum voluntary contraction) of the left FDI (Lt-FDI) muscle. F-waves of Rt-FDI muscle were recorded under these left index-finger conditions for ensuring that the excitability changes occur at the supraspinal level. MEPs were also recorded during motor imagery of the left index-finger abduction instead of overt movement. The results showed that, in single-pulse transcranial magnetic stimulation (TMS) paradigm, MEPs in Rt-FDI muscle were markedly enhanced during voluntary contractions of Lt-FDI muscle compared with the complete resting state. In paired-pulse TMS paradigm, the short intracortical inhibition was significantly reduced in proportion to increments of the ipsilateral muscle contraction, whereas the intracortical facilitation had no change. F-wave of Rt-FDI muscle was unchanged under these conditions, while MEP in Rt-FDI muscle was also enhanced during motor imagery of the left index-finger abduction. Based on the present results, it is suggested that the intracortical inhibitory neural circuits may be modulated in the transition from rest to activity of the ipsilateral homonymous muscle. The excitability changes in M1 might be induced by overflows of voluntary drive given to the ipsilateral limb, probably via the transcallosal pathway.     

Contralateral and ipsilateral motor activation in visual simple reaction time: a test of the hemispheric coactivation model

Motor potentials contralateral versus ipsilateral to the responding hand were examined in a visual simple reaction time (RT) experiment in order to test the hemispheric coactivation model of Miller (Cogn Psychol 49:118-154, 2004). Visual stimuli were presented on the left side of fixation, on the right side, or on both sides, and in the RT task participants had to respond as quickly as possible to the onset of any stimulus. The same stimulus displays were also presented in a counting task, for which participants had merely to count the stimuli. Hemisphere-specific movement-related potentials contralateral and ipsilateral to the responding hand were isolated by subtracting count-task ERPs from RT-task ERPs. Consistent with the hemispheric coactivation model, there was evidence of movement-related ipsilateral positivity as well as contralateral negativity, suggesting that the motor areas of both hemispheres contribute to response initiation in simple RT. The distinction between contralateral and ipsilateral motor activation appears useful in clarifying the roles of the two hemispheres in response initiation.     

Excitatory and inhibitory processes in primary motor cortex during the foreperiod of a warned reaction time task are unrelated to response expectancy

Reaction time (RT) is shortened when a warning signal precedes the response signal, a finding attributed to response preparation during the foreperiod between the warning and response signals. In a previous experiment, we delivered transcranial magnetic stimulation (TMS) during the short constant foreperiod of a warned RT task and found simultaneous suppression of motor evoked potential (MEP) amplitude and reduction of short-interval intracortical inhibition (SICI) on warned trials (Sinclair and Hammond in Exp Brain Res 186:385–392, 2008). To investigate the extent to which these phenomena are associated with response preparation we measured MEP amplitude and SICI during the foreperiod of a warned RT task in which three different warning signals specified the probability (0, 0.5, or 0.83) of response signal presentation. MEP amplitude was suppressed (Experiment 1) and SICI reduced (Experiment 2) equally in all of the warned conditions relative to when TMS was delivered in the inter-trial interval (ITI) suggesting that the modulation of primary motor cortex excitability during the foreperiod does not depend on momentary response expectancy induced by the warning signal. The reduction of SICI and suppression of MEP amplitude can be explained by assuming that a warning signal induces automatic motor cortical activation which is balanced by a competing inhibition to prevent premature response. A composite measure which weighted both speed and accuracy of response was positively correlated with the MEP amplitude during both the foreperiod and the ITI, suggesting that high motor cortical excitability is associated with optimized preparatory strategies for fast and accurate response.     

Organization of the forelimb area in squirrel monkey motor cortex: representation of digit,wrist, and elbow muscles

Summary The EMG in 8 to 14 hand, forearm, and arm muscles evoked by intracortical electrical stimulation was recorded at 433 sites in layer V in the region of the forelimb area of the primary motor cortex (MI) of three squirrel monkeys during ketamine anesthesia. At each site, the EMG was recorded at movement threshold (T) and at 1.5T and 2T at each site (but 60 A), and the threshold movement was noted. In the animals examined, the total MI forelimb area identified by movements or EMG occupied about 25 to 35 mm². At most sites from which a forelimb movement was evoked, EMG activity was evoked in one or more of the recorded muscles. One group of sites located rostrolaterally to the main forelimb area was separated by an intervening zone largely related to the face. The average area from which digit, wrist, elbow, or shoulder movement was evoked at threshold was nearly the same, and their movement thresholds were not significantly different. Average movement thresholds across the anterior-posterior extent of MI were also similar. All muscles recorded could be activated by cortical stimulation. Most commonly more than one muscle was activated from a single site. The highest individual EMG levels were produced at sites from which more than one muscle was activated. These results suggest that small regions of MI influence multiple muscles. Individual muscles were typically activated at multiple, spatially separated locations. For many muscles, increasing the stimulation intensity revealed additional separate areas of activation. Spatial locations of different muscles showed considerable interanimal variation. The size of most muscle representations was relatively large. The smallest representations always included the intrinsic hand muscles and the largest included the proximal muscles. Orderly topographic relationships among forelimb joints or muscles within the MI forelimb area were not apparent. Although distal muscle activation tended to be found posteriorly in the forelimb area and proximal muscles tended to be activated from anterior sites, both could be activated from broadly distributed and overlapping areas. The broad, overlapping nature of the muscle representation supports the concept that a small region of cortex is involved in controlling functional groups of muscles. The intermingling of muscle representations may provide a substrate for local cortical interactions among territories representing various muscle synergies or for changing associations of muscle groups. The representation plan derived from these mappings contains elements of all previously described summaries of MI organization. We argue that none of these adequately summarizes the internal organization of the MI forelimb area. Instead, patterns of organization that include functional combinations of muscles must be considered.     

Identification of cortical activation and white matter architecture according to short-term motor learning in the human brain: functional MRI and diffusion tensor tractography study

Objectives

The purpose of this study was to examine whether two weeks of short-term motor training led to changes of cortical activation patterns and white matter integrity in cortical and subcortical structures according to motor skill acquisition, using functional MRI (fMRI) and diffusion tensor image (DTI).

Methods

We enrolled twenty healthy volunteers, who were randomly assigned to training and control groups. The training group was trained with a serial reaction time (SRT) task, one hour a day for 10 days within two weeks, whereas the control group had no training. Movement accuracy (MA) and movement time (MT) were tested twice before and after training, while fMRI was scanned during SRT task. Immediately after these tests, DTI was conducted.

Results

The training group showed significant differences in the increase of MA and the reduction of MT, compared with control group. The activated volume of cortices related to motor function was gradually decreased in the training group, according to motor skill acquisition. DTI analysis showed no significant differences between pre- and post-tests in both groups.

Conclusions

Our findings indicated that short-term motor training led to cortical activation patterns of the cerebral cortex according to implicit motor learning. However, changes of white matter integrity were not observed. It seems that short-term motor training may not be enough to change white matter architectures, due to lack of the training period.