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.

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.

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.

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.

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.

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.

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.

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.

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.

Linking EEG signals, brain functions and mental operations: Advantages of the Laplacian transformation.

Electroencephalography (EEG) is a very popular technique for investigating brain functions and/or mental processes. To this aim, EEG activities must be interpreted in terms of brain and/or mental processes. EEG signals being a direct manifestation of neuronal activity it is often assumed that such interpretations are quite obvious or, at least, straightforward. However, they often rely on (explicit or even implicit) assumptions regarding the structures supposed to generate the EEG activities of interest. For these assumptions to be used appropriately, reliable links between EEG activities and the underlying brain structures must be established. Because of volume conduction effects and the mixture of activities they induce, these links are difficult to establish with scalp potential recordings. We present different examples showing how the Laplacian transformation, acting as an efficient source separation method, allowed to establish more reliable links between EEG activities and brain generators and, ultimately, with mental operations. The nature of those links depends on the depth of inferences that can vary from weak to strong. Along this continuum, we show that 1) while the effects of experimental manipulation can appear widely distributed with scalp potentials, Laplacian transformation allows to reveal several generators contributing (in different manners) to these modulations, 2) amplitude variations within the same set of generators can generate spurious differences in scalp potential topographies, often interpreted as reflecting different source configurations. In such a case, Laplacian transformation provides much more similar topographies, evidencing the same generator(s) set, and 3) using the LRP as an index of response activation most often produces ambiguous results, Laplacian-transformed response-locked ERPs obtained over motor areas allow resolving these ambiguities.

Spatial and temporal resolutions of EEG: Is it really black and white? A scalp current density view.

Among the different brain imaging techniques, electroencephalography (EEG) is classically considered as having an excellent temporal resolution, but a poor spatial one. Here, we argue that the actual temporal resolution of conventional (scalp potentials) EEG is overestimated, and that volume conduction, the main cause of the poor spatial resolution of EEG, also distorts the recovered time course of the underlying sources at scalp level, and hence degrades the actual temporal resolution of EEG. While Current Source Density (CSD) estimates, through the Surface Laplacian (SL) computation, are well known to dramatically reduce volume conduction effects and hence improve EEG spatial resolution, its positive impact on EEG temporal resolution is much less recognized. In two simulation studies, we first show how volume conduction and reference electrodes distort the scalp potential time course, and how SL transform provides a much better spatio-temporal description. We then exemplify similar effects on two empirical datasets. We show how the time courses of the scalp potentials mis-estimate the latencies of the relevant brain events and that CSD provides a much richer, and much more accurate, view of the spatio-temporal dynamics of brain activity.

Basics for sensorimotor information processing: some implications for learning.

In sensorimotor activities, learning requires efficient information processing, whether in car driving, sport activities or human-machine interactions. Several factors may affect the efficiency of such processing: they may be extrinsic (i.e., task-related) or intrinsic (i.e., subjects-related). The effects of these factors are intimately related to the structure of human information processing. In the present article we will focus on some of them, which are poorly taken into account, even when minimizing errors or their consequences is an essential issue at stake. Among the extrinsic factors, we will discuss, first, the effects of the quantity and quality of information, secondly, the effects of instruction and thirdly motor program learning. Among the intrinsic factors, we will discuss first the influence of prior information, secondly how individual strategies affect performance and, thirdly, we will stress the fact that although the human brain is not structured to function errorless (which is not new) humans are able to detect their errors very quickly and (in most of the cases), fast enough to correct them before they result in an overt failure. Extrinsic and intrinsic factors are important to take into account for learning because (1) they strongly affect performance, either in terms of speed or accuracy, which facilitates or impairs learning, (2) the effect of certain extrinsic factors may be strongly modified by learning and (3) certain intrinsic factors might be exploited for learning strategies.

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.

Choking under monitoring pressure: being watched by the experimenter reduces executive attention.

Performing more poorly given one's skill level ("choking") is likely in situations that offer an incentive if a certain outcome is achieved (outcome pressure) or when one is being watched by others-especially when one's performance is being evaluated (monitoring pressure). According to the choking literature, outcome pressure is associated with reduced executive control of attention, whereas monitoring pressure is associated with increased, yet counterproductive, attention to skill processes. Here, we show the first evidence that monitoring pressure-being watched by the experimenter-may lead individuals with higher working memory to choke on a classic measure of executive control-just the task effect thought to result from outcome pressure. Not only does this finding help refine our understanding of the processes underlying choking under monitoring pressure, but it also leads to a new look at classic audience effects, with an important implication for experimental psychology.

Dopamine precursors depletion impairs impulse control in healthy volunteers.

The aim of the present study was to decipher the role of the dopamine system in impulse control. Impulsive actions entail (i) activation of the motor system by an impulse, which is an urge to act and (ii) a failure to suppress that impulse, when inappropriate, in order to prevent an error. These two aspects of action impulsivity can be experimentally disentangled in conflict reaction time tasks such as the Simon task, which measures susceptibility to acting on spontaneous impulses (as well as the proficiency of suppressing these impulses). In 12 healthy volunteers performing a Simon task, dopamine availability was reduced with an amino acid drink deficient in the dopamine precursors, phenylalanine and tyrosine. Classic behavioral measures were augmented with an analysis of the electromyographic activity of the response effectors. Electromyography allows one to detect covert activations undetectable with strictly behavioral measures and further reveals the participants' ability to quickly suppress covert activations before they result in an overt movement. Following dopamine depletion, compared with a placebo condition, participants displayed comparable impulse activation but were less proficient at suppressing the interference from this activation. These results provide evidence that the dopamine system is directly involved in the suppression of maladaptive response impulses.

Dopa therapy and action impulsivity: subthreshold error activation and suppression in Parkinson's disease.

RATIONALE: Impulsive actions entail (1) capture of the motor system by an action impulse, which is an urge to act and (2) failed suppression of that impulse in order to prevent a response error. Several studies indicate that dopaminergic treatment can induce action impulsivity in patients diagnosed with Parkinson's disease (PD). Whether this effect is due to increased impulse expression or to decreased impulse suppression remains to be deciphered. METHOD: We used a novel approach based on electromyographic (EMG) analyses to decipher the effects of the patient's usual dopaminergic therapy on the expression and suppression of subliminal erroneous impulses. To this end, we used a within-subject design and took advantage of the Simon task, that elicits prepotent response tendencies. The patients (N = 15) performed the task on their usual dopaminergic medication and after complete medication withdrawal (for at least 12 h). RESULTS: The correction rate that measures the ability to suppress subthreshold impulsive muscle activity was lower when the patients were on medication as compared to their off medication state (p < 0.05). The incorrect activation rate that measures the capture of the motor system by action impulses was unaffected by medication. CONCLUSIONS: Dopa therapy affected action impulsivity. Although medication did not influence the incidence of fast action impulses, it significantly reduced patients' ability to abort and suppress muscle activation related to the incorrect response alternative.

Tactile stimulations and wheel rotation responses: toward augmented lane departure warning systems.

When an on-board system detects a drift of a vehicle to the left or to the right, in what way should the information be delivered to the driver? Car manufacturers have so far neglected relevant results from Experimental Psychology and Cognitive Neuroscience. Here we show that this situation possibly led to the sub-optimal design of a lane departure warning system (AFIL, PSA Peugeot Citroën) implemented in commercially available automobile vehicles. Twenty participants performed a two-choice reaction time task in which they were to respond by clockwise or counter-clockwise wheel-rotations to tactile stimulations of their left or right wrist. They performed poorer when responding counter-clockwise to the right vibration and clockwise to the left vibration (incompatible mapping) than when responding according to the reverse (compatible) mapping. This suggests that AFIL implements the worse (incompatible) mapping for the operators. This effect depended on initial practice with the interface. The present research illustrates how basic approaches in Cognitive Science may benefit to Human Factors Engineering and ultimately improve man-machine interfaces and show how initial learning can affect interference effects.

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.

Distributional reaction time properties in the Eriksen task: marked differences or hidden similarities with the Simon task?

In conflict tasks, the irrelevant stimulus attribute needs to be suppressed for the correct response to be produced. In the Simon task, earlier researchers have proposed that this suppression is the reason that, after an initial increase, the interference effect decreases for longer RTs, as reflected by late, negative-going delta plots. This view has been challenged by observations of positive-going delta plots, even for long RTs, in other conflict tasks, despite a similar necessity for suppression. For late negative-going delta plots to be interpreted as reflecting suppression, a necessary, although maybe not sufficient, condition is that similar patterns should be observed for other conflict tasks. We reasoned that a similar suppression could be present, but hidden, in the Eriksen flanker task. By recording and analyzing electromyograms of the muscles involved in response execution, we could compute delta plots separately for trials that elicited a subthreshold incorrect response activation (partial error). Late negative-going delta plots were observable on partial-error trials, although they were weaker than for the Simon task, reducing the impact of this inversion on the overall distribution. We further showed that this pattern is modulated by time pressure. Those results indicate that mechanisms leading to negative-going delta plots, similar to those observed in the Simon task, are also at play in the Eriksen task. The link between negative-going delta plots and executive online control is discussed.