A measure of the interference effect distribution.

We elaborated an index, the Interference Distribution Index, which allows quantifying the relation between response times and the size of the interference effect. This index is associated with an intuitive graphical representation, the Lorenz-interference plot. We show that this index has some convenient properties in terms of sensitivity to changes in the distribution of the interference effect and to aggregation of individual data. Moreover, it turns out that this index is the only one (up to an arbitrary increasing transformation) possessing these properties. The relevance of this index is illustrated through simulations of a cognitive model of interference effects and reanalysis of experimental data.

Controlling your impulses: electrical stimulation of the human supplementary motor complex prevents impulsive errors.

To err is human. However, an inappropriate urge does not always result in error. Impulsive errors thus entail both a motor system capture by an urge to act and a failed inhibition of that impulse. Here we show that neuromodulatory electrical stimulation of the supplementary motor complex in healthy humans leaves action urges unchanged but prevents them from turning into overt errors. Subjects performed a choice reaction-time task known to trigger impulsive responses, leading to fast errors that can be revealed by analyzing accuracy as a function of poststimulus time. Yet, such fast errors are only the tip of the iceberg: electromyography (EMG) revealed fast subthreshold muscle activation in the incorrect response hand in an even larger proportion of overtly correct trials, revealing covert response impulses not discernible in overt behavior. Analyzing both overt and covert response tendencies enables to gauge the ability to prevent these incorrect impulses from turning into overt action errors. Hyperpolarizing the supplementary motor complex using transcranial direct current stimulation (tDCS) preserves action impulses but prevents their behavioral expression. This new combination of detailed behavioral, EMG, and tDCS techniques clarifies the neurophysiology of impulse control, and may point to avenues for improving impulse control deficits in various neurologic and psychiatric disorders.

Effects of hyperbaric nitrogen-induced narcosis on response-selection processes.

Certain underwater circumstances carry risk of inert gas narcosis. Impairment of sensorimotor information processing due to narcosis, induced by normobaric nitrous oxide or high partial nitrogen pressure, has been broadly evidenced, by a lengthening of the reaction time (RT). However, the locus of this effect remains a matter of debate. We examined whether inert gas narcosis affects the response-selection stage of sensorimotor information processing. We compared an air normobaric condition with a hyperbaric condition in which 10 subjects were subjected to 6 absolute atmospheres of 8.33% O2 Nitrox. In both conditions, subjects performed a between-hand choice-RT task in which we explicitly manipulated the stimulus-response association rule. The effect of this manipulation (which is supposed to affect response-selection processes) was modified by inert gas narcosis. It is concluded, therefore, that response selection processes are among the loci involved in the effect of inert gas narcosis on information processing.

Why is there an error negativity on correct trials? A reappraisal.

In healthy subjects, the Error Negativity (Ne) was initially reported on errors and on partial errors, only. Later on, application of the Laplacian transformation to EEG data unmasked a Ne-like wave (Nc) that shares a main generator with the Ne, suggesting that the Nc is just a small Ne. However, the reason why a small Ne would persist on correct responses remains unclear. Now, sometimes, subthreshold EMG activations in the muscles corresponding to correct responses (not strong enough to reach the response threshold) can precede full-blown correct responses. These "partially correct" activities seem to correspond to (force) execution errors, as they evoke a sizeable Ne. Within the frames of the Reward Value and Prediction Model or of the Predicted Response-Outcome model we propose that the action monitoring system evokes a Ne/Nc on correct responses because, even when a correct choice has been made, the accuracy of response (force) execution cannot be fully predicted. If this interpretation is correct, it can be assumed that, once these execution errors have been corrected, the correctness of the (full-blown) correcting response is highly predictable. Consequently, they should evoke a smaller Nc/Ne than "pure" correct responses. We show, that for the response thresholds set in the present experiment, the correcting response of the trials containing a partially correct activation evoke no identifiable Nc at all. Therefore it seems that there usually is an Error Negativity on correct trials because the correctness of response (force) execution cannot be fully predicted.

Dynamics of executive control and motor deficits in parkinsonian rats.

While there is general agreement that in Parkinson's disease (PD), striatal dopamine (DA) depletion causes motor deficits, the origin of the associated cognitive impairments remains a matter of debate. The present study aimed to decipher the influence of a partial 6-hydroxydopamine (6-OHDA) lesion of striatal DA nerve terminals in rats performing a reaction time task previously used to assess cognitive deficits in PD patients. The effects of two behavioral manipulations-foreperiod duration and stimulus-response congruence-known to affect motor processes and executive control, respectively, were studied over 8 weeks postsurgery in control and lesion animals. Two weeks after surgery, the lesion abolished the effect of foreperiod, confirming the direct involvement of striatal DA in motor processes, but failed to alter the effect of congruence. During the following weeks, the effect of foreperiod was reinstated, indicating a recovery of lesion-induced motor symptoms. This recovery was accompanied by a progressive increase of the congruence effect, signaling an executive control deficit in lesion animals. This result provides the first evidence that 6-OHDA lesioned rats exhibit the same cognitive impairment as PD patients in this task. The deficit, however, built up progressively after the lesion and may result from adaptations mitigating lesion-induced motor deficits.

On the Comparison Between the Nc/CRN and the Ne/ERN.

After the Error Negativity (Ne or ERN) has been described on full-blown errors and on partial error, a smaller Error Negativity-like wave (CRN or Nc) has also been evidenced on correct trials, first in patients with schizophrenia and, later on, in healthy subjects. The functional significance of the Nc as compared to the Ne is of critical importance since most models accounting for the genesis of the Ne on errors and partial errors cannot account for the existence of the Nc if this Nc simply corresponds to a small Ne. On the contrary, if the Nc and the Ne are two completely distinct components, then the existence of a Nc poses no constraint to the existing models. To this end, we examine in the present review the similarities and the differences existing between the Ne and the Nc regarding their functional properties and their anatomical origin.

Mechanisms and dynamics of cortical motor inhibition in the stop-signal paradigm: a TMS study.

The ability to stop ongoing motor responses in a split-second is a vital element of human cognitive control and flexibility that relies in large part on prefrontal cortex. We used the stop-signal paradigm to elucidate the engagement of primary motor cortex (M1) in inhibiting an ongoing voluntary motor response. The stop-signal paradigm taps the ability to flexibly countermand ongoing voluntary behavior upon presentation of a stop signal. We applied single-pulse TMS to M1 at several intervals following the stop signal to track the time course of excitability of the motor system related to generating and stopping a manual response. Electromyography recorded from the flexor pollicis brevis allowed quantification of the excitability of the corticospinal tract and the involvement of intracortical GABA(B)ergic circuits within M1, indexed respectively by the amplitude of the motor-evoked potential and the duration of the late part of the cortical silent period (SP). The results extend our knowledge of the neural basis of inhibitory control in three ways. First, the results revealed a dynamic interplay between response activation and stopping processes at M1 level during stop-signal inhibition of an ongoing response. Second, increased excitability of inhibitory interneurons that drives SP prolongation was evident as early as 134 msec following the instruction to stop. Third, this pattern was followed by a stop-related reduction of corticospinal excitability implemented around 180 after the stop signal. These findings point to the recruitment of GABA(B)ergic intracortical inhibitory circuits within M1 in stop-signal inhibition and support the notion of stopping as an active act of control.

Rostral Cingulate Zone and correct response monitoring: ICA and source localization evidences for the unicity of correct- and error-negativities.

Falkenstein et al. (1991) first described a negative wave occurring just after an erroneous response in choice Reaction time tasks ("Error Negativity"-Ne or "Error Related Negativity"-ERN). Thanks to Laplacian transform of the data, Vidal et al. (2000, 2003a) described a wave on correct trials with similar topography and latency, although of smaller amplitude compared to the errors. A critical question is whether the Ne observed on errors and the negativity reported on correct trials reflect the same (modulated) activity, or whether they reflect completely different mechanisms. These two alternative possibilities were tested thanks to Independent Component Analysis (ICA) and source localization. ICA results showed that the waves recorded on errors and correct trials can be accounted for by the same independent component, corresponding to a dipolar source located within the Rostral Cingulate Zone. Source localization on the raw data also confirmed a common generator for correct and error trials. These data suggest that the waves on errors and correct trials reflect the same brain activity, whose amplitude varies as a function of the correctness of the response. The implications of this result for cognitive control are discussed.

Direct evidence for cortical suppression of somatosensory afferents during visuomotor adaptation.

Upon exposure to novel visuomotor relationships, the information carried by visual and proprioceptive signals becomes discrepant, often disrupting motor execution. It has been shown that degradation of the proprioceptive sense (arising either from disease or experimental manipulation) enhances performance when drawing with mirror-reversed vision. Given that the central nervous system can exert a dynamic control over the transmission of afferent signals, reducing proprioceptive inflow to cortical areas could be part of the normal adaptive mechanisms deployed in healthy humans upon exposure to novel visuomotor environments. Here we address this issue by probing the transmission of somatosensory afferents throughout the course of adaptation to a visuomotor conflict, by recording median nerve somatosensory evoked potentials. We show that early exposure to tracing with mirror-reversed vision is accompanied by substantial proprioceptive suppression occurring in the primary somatosensory cortex (S1). This proprioceptive gating is gradually alleviated as performance increases with adaptation, returning to baseline levels. Peripheral and spinal evoked potentials were not modulated throughout, suggesting that the gating acted to reduce cortico-cortico excitability directly within S1. These modulations provide neurophysiological evidence for flexibility in sensory integration during visuomotor adaptation, which may functionally serve to reduce the sensory conflict until the visuo-proprioceptive mapping is updated.

Spatio-temporal dynamics of reach-related neural activity for visual and somatosensory targets.

The parieto-frontal network plays a crucial role in the transformations that convert visual information into motor commands for hand reaching movements. Here we use electroencephalography to determine whether the planning of reaching movements to visual and somatosensory targets involves a similar spatio-temporal pattern of neural activity. Subjects performed reaching movements toward spatial locations defined either by visual (light-emitting diode) or somatosensory (vibration of a fingertip of the contralateral hand) stimuli. To identify the activations associated with sensorimotor transformations, we subtracted the event-related potentials recorded in a "static" task (the stimuli were presented but no movement was initiated) from those recorded in a "reach" task (a reach had to be initiated toward the spatial location of the stimuli). In the visual condition, reach-related activities were observed over parietal, premotor and sensorimotor areas contralateral to the reaching hand. Activation was first observed over parietal areas 140 ms after stimulus onset and progressed to frontal areas. The proprioceptive condition recruited a similar set of structures as for visual targets. However, the temporal pattern of activity within these cortical areas differed greatly. Activity was sustained over premotor and sensorimotor areas throughout the reaction time interval, occurring simultaneously with the parietal activation. These results suggest that a common cortical network serves to transform visual and somatosensory signals into motor commands, but that the interactions between the structures of this network differ. This raises the possibility that different coordinate frames are used to encode the motor error for the two target modalities.

Sequential adjustments before and after partial errors.

In choice reaction time tasks, subjects speed up before making an error, but slow down afterward to prevent the occurrence of a new error. In some trials, the correct response is preceded by an incorrect electromyographic (EMG) activation too small to reach the response threshold. In this article, we show that these incorrect EMG activations give rise to the same sequential effects as overt errors: Before a trial containing an incorrect EMG activation, subjects speed up, whereas after that trial, they slow down. These activations reflect errors that have been detected, inhibited, and corrected in time. As such, they index the involvement of online executive control.

Errors and Action Monitoring: Errare Humanum Est Sed Corrigere Possibile.

It was recognized long ago by Seneca through his famous "errare humanum est." that the human information processing system is intrinsically fallible. What is newer is the fact that, at least in sensorimotor information processing realized under time pressure, errors are largely dealt with by several (psycho)physiological-specific mechanisms: prevention, detection, inhibition, correction, and, if these mechanisms finally fail, strategic behavioral adjustments following errors. In this article, we review several datasets from laboratory experiments, showing that the human information processing system is well equipped not only to detect and correct errors when they occur but also to detect, inhibit, and correct them even before they fully develop. We argue that these (psycho)physiological mechanisms are important to consider when the brain works in everyday settings in order to render work systems more resilient to human errors and, thus, safer.

A Simon-like effect in Go/No-Go tasks performed in isolation.

The present study was conducted to decipher whether a spatial correspondence effect can emerge in Go/No-Go tasks (cSE, in reference to Donders' type c task) performed in isolation (participant alone in the cubicle). To this aim, a single participant was centrally positioned in front of a device and was required to respond by a hand key-press to the color of the stimulus. Half the participants were seated in front of a table equipped with only one response key and the other half in front of a table equipped with two response keys (one active and the other one useless). Using a substantial number of subjects (48) and trials (960), the present study revealed a numerically small but statistically reliable cSE. This result contrasts with referential coding predictions and suggests that the representation of a concurrently active response is not a prerequisite for the cSE to emerge. Moreover, the presence of a second response button in the participant's peripersonal space exerted no measurable influence on the cSE. The lack of statistical power of numerous previous studies may explain why the cSE has often been considered to be nil.

[Psychomotor retardation associated to depression differs from that of normal aging]. / Mise en évidence d'un ralentissement psychomoteur spécifique à la dépression chez le sujet âgé

Psychomotor retardation (PMR) is a main symptom of depressive illness. In the elderly, it is associated with the severity of depression and poor prognosis. However, PMR is also commonly associated with normal aging, Therefore, depressive PMR is frequently misinterpreted as the age-related slowing in the elderly, which contributes to poor recognition of depression. Moreover, neurobiological and neuroanatomical studies on PMR hardly allowed to discriminate geriatric depression from normal aging. Reaction time experiments have rarely been tested in elderly population although they are particularly suited for examining PMR. We performed two reaction time experiments using an additive factor analysis in healthy and depressed old individuals with the hypothesis that PMR associated to depression differed from that of normal aging. Results showed that age-related PMR affected all stages of central nervous system information processing, while PMR associated with depression is limited to the components of response-selection and motor-adjustment. These results clearly show that PMR in geriatric depression differed from the age related slowing. Depression also spares stimulus preprocessing in old individuals as it did in younger adults.

The Way We Do the Things We Do: How Cognitive Contexts Shape the Neural Dynamics of Motor Areas in Humans.

In spontaneously triggered movements the nature of the executed response has a prominent effect on the intensity and the dynamics of motor areas recruitment. Under time pressure, the time course of motor areas recruitment is necessarily shorter than that of spontaneously triggered movements because RTs may be extremely short. Moreover, different classes of RT tasks allow examining the nature and the dynamics of motor areas activation in different cognitive contexts. In the present article, we review experimental results obtained from high temporal resolution methods (mainly, but not exclusively EEG ones), during voluntary movements; these results indicate that the activity of motor areas not only depends on the nature of the executed movement but also on the cognitive context in which these movements have to be executed.

Dopamine and response selection: an Acute Phenylalanine/Tyrosine Depletion study.

The role of dopaminergic system in decision-making is well documented, and evidence suggests that it could play a significant role in response selection processes. The N-40 is a fronto-central event-related potential, generated by the supplementary motor areas (SMAs) and a physiological index of response selection processes. The aim of the present study was to determine whether infraclinical effects of dopamine depletion on response selection processes could be evidenced via alterations of the N-40. We obtained a dopamine depletion in healthy volunteers with the acute phenylalanine and tyrosine depletion (APTD) method which consists in decreasing the availability of dopamine precursors. Subjects realized a Simon task in the APTD condition and in the control condition. When the stimulus was presented on the same side as the required response, the stimulus-response association was congruent and when the stimulus was presented on the opposite side of the required response, the stimulus-response association was incongruent. The N-40 was smaller for congruent associations than for incongruent associations. Moreover, the N-40 was sensitive to the level of dopaminergic activity with a decrease in APTD condition compared to control condition. This modulation of the N-40 by dopaminergic level could not be explained by a global decrease of cerebral electrogenesis, since negativities and positivities indexing the recruitment of the primary motor cortex (anatomically adjacent to the SMA) were unaffected by APTD. The specific sensitivity of N-40 to ATPD supports the model of Keeler et al. (Neuroscience 282:156-175, 2014) according to which the dopaminergic system is involved in response selection.

Subthalamic nucleus stimulation, dopaminergic treatment and impulsivity in Parkinson's disease.

BACKGROUND: Deep brain stimulation of the subthalamic nucleus (STN DBS) is known to increase response speed and lower response accuracy in Parkinson's disease (PD) patients. It has been proposed that this speed-accuracy tradeoff is due to enhanced sensitivity of the motor system to sensory information. An alternative possibility is that this effect is due to weakened suppressive processes. The two alternative interpretations can be tested by analyzing the electromyographic activity (EMG) of the response agonists when the patients perform conflict reaction time tasks. In those tasks, fast subthreshold muscle impulses often occur in the agonist of the incorrect response. These impulses are partial errors that are suppressed before being behaviourally committed. MATERIAL AND METHODS: Here we analyzed the EMG of the response agonists recorded while sixteen PD patients performed a Simon task that elicits prepotent response tendencies so as to decipher (i) whether STN DBS affects the expression and/or suppression of subthreshold muscle impulses that are critical for action control and (ii) the interaction between dopaminergic treatment and STN DBS. The patients were tested On and Off STN DBS and On and Off dopaminergic medication in a full factorial design. RESULTS: STN DBS not only impaired the proficiency to suppress subliminal action impulses (p = 0.01) but also favoured the muscular expression of fast incorrect impulses (p < 0.001). Dopaminergic treatment only affected the action impulses suppression (p = 0.02) and did not change the effect of STN DBS on impulsive action control. CONCLUSION: Contrary to a recent proposal, STN DBS impaired rather than improved action control by weakening erroneous impulse suppression, whether the patients were On or Off their usual medication. These findings are discussed in light of a recent proposal (Servant M, White C, Montagnini A, Burle B, 2015) that reconciles partial errors with accumulation-to-bound models of decision making. Our results suggest that medication specifically lowers the mechanical threshold while STN DBS lowers the mechanical threshold and to a lesser extent the EMG-threshold.

Beyond decision! Motor contribution to speed-accuracy trade-off in decision-making.

Both in real life and experimental settings, increasing response speed typically leads to more error-prone actions. Processes underlying such a "speed-accuracy trade-off" (SAT) are usually assumed to be purely decisional: cautiousness would be determined only by the amount of sensory evidence required to select a response. The present data challenges this largely accepted view, by directly showing that motor processes are speeded up under time pressure. In a choice reaction time task where emphasis was put either on response speed or accuracy, motor processes were investigated through the analysis of muscular activity related to response execution. When response speed was emphasized, the time between electromyographic onset and behavioral response (motor time) was also speeded up (contributing to more than 20 % of the total effect on global reaction time). This speeded execution (likely due to a more efficient motor command) may also explain why participants are less able to interrupt incorrect response execution once started (Burle et al., Psychonomic Bulletin & Review, 21(4), 1003-1010, 2014), leading to more overt errors. Pointing to a speed-accuracy exchange within motor processes themselves, the present results call for a re-evaluation of widely accepted assumptions about SAT, and more generally, decision-making processes. They are discussed in the context of recent extensions of the drift diffusion model framework, questioning the strict separation between decisional and motor processes.

On-line action monitoring of response execution: An electrophysiological study.

In between-hand choice-RT-tasks, small incorrect EMG activations occurring before the correct response ("partial errors") are assumed to reflect the detection, inhibition and correction of erroneous hand selection, revealing the existence of an action monitoring system, acting "on-line". Now, EMG activations of the correctly selected hand muscles, too small to reach the response threshold, may also occur before these hand muscles produce an overt correct response ("partial corrects"). We hypothesized that partial corrects reflect incorrect execution of correctly selected responses. We found 1) that response force was smaller on trials preceding a partial correct trial and 2) that the Error Negativity, a performance sensitive ERP, assumed to reveal "on-line" action monitoring, was larger for partial corrects than for correct trials. This also suggests that the competence of the action monitoring system is not restricted to selection errors but also extends to execution errors.

Knowing when to respond and the efficiency of the cortical motor command: a Laplacian ERP study.

The objective was to test whether motor preparation can modulate the efficiency of the cortical motor command. The electroencephalogram (EEG) was recorded from electrodes located over the primary sensorimotor cortices during the performance of a between-hand choice reaction time task in which foreperiod duration (the interval between the warning and the imperative signals, 800 vs. 2800 ms) was varied across blocks of trials. In order to increase the spatial resolution of the EEG traces, surface Laplacian was estimated. The amplitude of the negative wave developing over the hemisphere contralateral to the response was smaller for the short foreperiod associated with the best performance level. These results indicate that the activation of the primary sensorimotor cortex involved in the response is less pronounced for the short foreperiod, suggesting that temporal advance information increases the efficiency of the cortical motor command.