Time for Action: Neural Basis of the Costs and Benefits of Temporal Predictability for Competing Response Choices.

The brain can anticipate the time of imminent events to optimize sensorimotor processing. Yet, there can be behavioral costs of temporal predictability under situations of response conflict. Here, we sought to identify the neural basis of these costs and benefits by examining motor control processes in a combined EEG-EMG study. We recorded electrophysiological markers of response activation and inhibition over motor cortex when the onset-time of visual targets could be predicted, or not, and when responses necessitated conflict resolution, or not. If stimuli were temporally predictable but evoked conflicting responses, we observed increased intertrial consistency in the delta range over the motor cortex involved in response implementation, perhaps reflecting increased response difficulty. More importantly, temporal predictability differentially modulated motor cortex activity as a function of response conflict before the response was even initiated. This effect occurred in the hemisphere ipsilateral to the response, which is involved in inhibiting unwanted actions. If target features all triggered the same response, temporal predictability increased cortical inhibition of the incorrect response hand. Conversely, if different target features triggered two conflicting responses, temporal predictability decreased inhibition of the incorrect, yet prepotent, response. This dissociation reconciles the well-established behavioral benefits of temporal predictability for nonconflicting responses as well as its costs for conflicting ones by providing an elegant mechanism that operates selectively over the motor cortex involved in suppressing inappropriate actions just before response initiation. Taken together, our results demonstrate that temporal information differentially guides motor activity depending on response choice complexity.

Don't Stop Me Now: Neural Underpinnings of Increased Impulsivity to Temporally Predictable Events.

Although the benefit of temporal predictability for behavior is long-established, recent studies provide evidence that knowing when an important event will occur comes at the cost of greater impulsivity. Here, we investigated the neural basis of inhibiting actions to temporally predictable targets using an EEG-EMG method. In our temporally cued version of the stop-signal paradigm (two-choice task), participants used temporal information delivered by a symbolic cue to speed their responses to the target. In a quarter of the trials, an auditory signal indicated that participants had to inhibit their actions. Behavioral results showed that although temporal cues speeded RTs, they also impaired the ability to stop actions as indexed by longer stop-signal reaction time. In line with behavioral benefits of temporal predictability, EEG data demonstrated that acting at temporally predictable moments facilitated response selection at the cortical level (reduced frontocentral negativity just before the response). Likewise, activity of the motor cortex involved in suppression of incorrect response hand was stronger for temporally predictable events. Thus, by keeping an incorrect response in check, temporal predictability likely enabled faster implementation of the correct response. Importantly, there was no effect of temporal cues on the EMG-derived index of online, within-trial inhibition of subthreshold impulses. This result shows that although participants were more prone to execute a fast response to temporally predictable targets, their inhibitory control was, in fact, unaffected by temporal cues. Altogether, our results demonstrate that greater impulsivity when responding to temporally predictable events is paralleled by enhanced neural motor processes involved in response selection and implementation rather than impaired inhibitory control.

Developmental changes in impulse control: Trial-by-trial EMG dissociates the evolution of impulse strength from its subsequent suppression.

Goal-oriented behavior can be disrupted by irrelevant information that automatically activates incorrect responses. While behavioral errors reveal response capture in such situations, they are only the tip of the iceberg. Additional subliminal activations of the incorrect responses (partial errors) can be revealed on correctly responded trials thanks to electromyography (EMG). In the current study, for the first time, EMG recorded in children was combined with distributional analyses. This allowed to investigate the properties of incorrect response activations and to highlight developmental changes in impulse control. A sample of 114 children aged 6-14 years was studied. Children performed a Simon task in which the irrelevant stimulus-position automatically activates a response that might be compatible or incompatible with the correct one. On incompatible trials, the automatic response activation must be overcome by controlled response selection. As previously observed in adults, our approach revealed the presence of an automatic EMG activation of the incorrect response elicited by the irrelevant stimulus dimension. Further, it revealed another independent source at the origin of incorrect response activations: the tendency to guess for response alternation. Both sources increased the frequency of early incorrect EMG activations, indicating impulsive responding. In addition, the influence of both sources decreased with increasing age. Thus, development is marked by improved ability to manage distractibility on the one hand and decreased tendency to rely on a guessing strategy on the other.

Conflict processing in kindergarten children: New evidence from distribution analyses reveals the dynamics of incorrect response activation and suppression.

The development of cognitive control is known to follow a long and protracted development. However, whether the interference effect in conflict tasks in children would entail the same core processes as in adults, namely an automatic activation of incorrect response and its subsequent suppression, remains an open question. We applied distributional analyses to reaction times and accuracy of 5- and 6-year-old children performing three conflict tasks (flanker, Simon, and Stroop) in a within-participants design. This revealed both strong commonalities and differences between children and adults. As in adults, fast responses were more error prone than slow ones on incompatible trials, indicating a fast "automatic" activation of the incorrect response. In addition, the strength of this activation differed across tasks, following a pattern similar to that of adults. Moreover, modeling the data with a drift diffusion model adapted for conflict tasks allowed one to better assess the origin of the typical slowing down observed in children. Besides showing that advanced distribution analyses can be successfully applied to children, the current results support the notion that interference effects in 5- and 6-year-olds are driven by mechanisms very similar to the ones at play in adults but with different time courses.

Using Covert Response Activation to Test Latent Assumptions of Formal Decision-Making Models in Humans.

Most decisions that we make build upon multiple streams of sensory evidence and control mechanisms are needed to filter out irrelevant information. Sequential sampling models of perceptual decision making have recently been enriched by attentional mechanisms that weight sensory evidence in a dynamic and goal-directed way. However, the framework retains the longstanding hypothesis that motor activity is engaged only once a decision threshold is reached. To probe latent assumptions of these models, neurophysiological indices are needed. Therefore, we collected behavioral and EMG data in the flanker task, a standard paradigm to investigate decisions about relevance. Although the models captured response time distributions and accuracy data, EMG analyses of response agonist muscles challenged the assumption of independence between decision and motor processes. Those analyses revealed covert incorrect EMG activity ("partial error") in a fraction of trials in which the correct response was finally given, providing intermediate states of evidence accumulation and response activation at the single-trial level. We extended the models by allowing motor activity to occur before a commitment to a choice and demonstrated that the proposed framework captured the rate, latency, and EMG surface of partial errors, along with the speed of the correction process. In return, EMG data provided strong constraints to discriminate between competing models that made similar behavioral predictions. Our study opens new theoretical and methodological avenues for understanding the links among decision making, cognitive control, and motor execution in humans. SIGNIFICANCE STATEMENT: Sequential sampling models of perceptual decision making assume that sensory information is accumulated until a criterion quantity of evidence is obtained, from where the decision terminates in a choice and motor activity is engaged. The very existence of covert incorrect EMG activity ("partial error") during the evidence accumulation process challenges this longstanding assumption. In the present work, we use partial errors to better constrain sequential sampling models at the single-trial level.

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.

Linking Theoretical Decision-making Mechanisms in the Simon Task with Electrophysiological Data: A Model-based Neuroscience Study in Humans.

A current challenge for decision-making research is in extending models of simple decisions to more complex and ecological choice situations. Conflict tasks (e.g., Simon, Stroop, Eriksen flanker) have been the focus of much interest, because they provide a decision-making context representative of everyday life experiences. Modeling efforts have led to an elaborated drift diffusion model for conflict tasks (DMC), which implements a superimposition of automatic and controlled decision activations. The DMC has proven to capture the diversity of behavioral conflict effects across various task contexts. This study combined DMC predictions with EEG and EMG measurements to test a set of linking propositions that specify the relationship between theoretical decision-making mechanisms involved in the Simon task and brain activity. Our results are consistent with a representation of the superimposed decision variable in the primary motor cortices. The decision variable was also observed in the EMG activity of response agonist muscles. These findings provide new insight into the neurophysiology of human decision-making. In return, they provide support for the DMC model framework.

Single subject analyses reveal consistent recruitment of frontal operculum in performance monitoring.

There are continuing uncertainties regarding whether performance monitoring recruits the anterior insula (aI) and/or the frontal operculum (fO). The proximity and morphological complexity of these two regions make proper identification and isolation of the loci of activation extremely difficult. The use of group averaging methods in human neuroimaging might contribute to this problem. The result has been heterogeneous labeling of this region as aI, fO, or aI/fO, and a discussion of results oriented towards either cognitive or interoceptive functions depending on labeling. In the present article, we adapted the spatial preprocessing of functional magnetic resonance imaging data to account for group averaging artifacts and performed a subject-by-subject analysis in three performance monitoring tasks. Results show that functional activity related to feedback or action monitoring consistently follows local morphology in this region and demonstrate that the activity is located predominantly in the fO rather than in the aI. From these results, we propose that a full understanding of the respective role of aI and fO would benefit from increased spatial resolution and subject-by-subject analysis.

Online extraction and single trial analysis of regions contributing to erroneous feedback detection.

Understanding how the brain processes errors is an essential and active field of neuroscience. Real time extraction and analysis of error signals provide an innovative method of assessing how individuals perceive ongoing interactions without recourse to overt behaviour. This area of research is critical in modern Brain-Computer Interface (BCI) design, but may also open fruitful perspectives in cognitive neuroscience research. In this context, we sought to determine whether we can extract discriminatory error-related activity in the source space, online, and on a trial by trial basis from electroencephalography data recorded during motor imagery. Using a data driven approach, based on interpretable inverse solution algorithms, we assessed the extent to which automatically extracted error-related activity was physiologically and functionally interpretable according to performance monitoring literature. The applicability of inverse solution based methods for automatically extracting error signals, in the presence of noise generated by motor imagery, was validated by simulation. Representative regions of interest, outlining the primary generators contributing to classification, were found to correspond closely to networks involved in error detection and performance monitoring. We observed discriminative activity in non-frontal areas, demonstrating that areas outside of the medial frontal cortex can contribute to the classification of error feedback activity.

Conflict tasks and the diffusion framework: Insight in model constraints based on psychological laws.

Formal models of decision-making have traditionally focused on simple, two-choice perceptual decisions. To date, one of the most influential account of this process is Ratcliff's drift diffusion model (DDM). However, the extension of the model to more complex decisions is not straightforward. In particular, conflicting situations, such as the Eriksen, Stroop, or Simon tasks, require control mechanisms that shield the cognitive system against distracting information. We adopted a novel strategy to constrain response time (RT) models by concurrently investigating two well-known empirical laws in conflict tasks, both at experimental and modeling levels. The two laws, predicted by the DDM, describe the relationship between mean RT and (i) target intensity (Piéron's law), (ii) standard deviation of RT (Wagenmakers-Brown's law). Pioneering work has shown that Piéron's law holds in the Stroop task, and has highlighted an additive relationship between target intensity and compatibility. We found similar results in both Eriksen and Simon tasks. Compatibility also violated Wagenmakers-Brown's law in a very similar and particular fashion in the two tasks, suggesting a common model framework. To investigate the nature of this commonality, predictions of two recent extensions of the DDM that incorporate selective attention mechanisms were simulated and compared to the experimental results. Both models predict Piéron's law and the violation of Wagenmakers-Brown's law by compatibility. Fits of the models to the RT distributions and accuracy data allowed us to further reveal their relative strengths and deficiencies. Combining experimental and computational results, this study sets the groundwork for a unified model of decision-making in conflicting environments.

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.

General-purpose monitoring during speech production.

The concept of "monitoring" refers to our ability to control our actions on-line. Monitoring involved in speech production is often described in psycholinguistic models as an inherent part of the language system. We probed the specificity of speech monitoring in two psycholinguistic experiments where electroencephalographic activities were recorded. Our focus was on a component previously reported in nonlinguistic manual tasks and interpreted as a marker of monitoring processes. The error negativity (Ne, or error-related negativity), thought to originate in medial frontal areas, peaks shortly after erroneous responses. A component of seemingly comparable properties has been reported, after errors, in tasks requiring access to linguistic knowledge (e.g., speech production), compatible with a generic error-detection process. However, in contrast to its original name, advanced processing methods later revealed that this component is also present after correct responses in visuomotor tasks. Here, we reported the observation of the same negativity after correct responses across output modalities (manual and vocal responses). This indicates that, in language production too, the Ne reflects on-line response monitoring rather than error detection specifically. Furthermore, the temporal properties of the Ne suggest that this monitoring mechanism is engaged before any auditory feedback. The convergence of our findings with those obtained with nonlinguistic tasks suggests that at least part of the monitoring involved in speech production is subtended by a general-purpose mechanism.

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.

Preparing to React: A Behavioral Study on the Interplay between Proactive and Reactive Action Inhibition.

Motor preparation, based on one's goals and expectations, allows for prompt reactions to stimulations from the environment. Proactive and reactive inhibitory mechanisms modulate this preparation and interact to allow a flexible control of responses. In this study, we investigate these two control mechanisms with an ad hoc cued Go/NoGo Simon paradigm in a within-subjects design, and by measuring subliminal motor activities through electromyographic recordings. Go cues instructed participants to prepare a response and wait for target onset to execute it (Go target) or inhibit it (NoGo target). Proactive inhibition keeps the prepared response in check, hence preventing false alarms. Preparing the cue-coherent effector in advance speeded up responses, even when it turned out to be the incorrect effector and reactive inhibition was needed to perform the action with the contralateral one. These results suggest that informative cues allow for the investigation of the interaction between proactive and reactive action inhibition. Partial errors' analysis suggests that their appearance in compatible conflict-free trials depends on cue type and prior preparatory motor activity. Motor preparation plays a key role in determining whether proactive inhibition is needed to flexibly control behavior, and it should be considered when investigating proactive/reactive inhibition.

Speeding-up while growing-up: Synchronous functional development of motor and non-motor processes across childhood and adolescence.

Describing the maturation of information processing in children is fundamental for developmental science. Although non-linear changes in reaction times have been well-documented, direct measurement of the development of the different processing components is lacking. In this study, electromyography was used to quantify the maturation of premotor and motor processes on a sample of 114 children (6-14 years-old) and 15 adults. Using a model-based approach, we show that the development of these two components is well-described by an exponential decrease in duration, with the decay rate being equal for the two components. These findings provide the first unbiased evidence in favour of the common developmental rate of nonmotor and motor processes by directly confronting rates of development of different processing components within the same task. This common developmental rate contrasts with the differential physical maturation of region-specific cerebral gray and white matter. Tentative paths of interpretation are proposed in the discussion.

Assessing model-based inferences in decision making with single-trial response time decomposition.

The latent psychological mechanisms involved in decision-making are often studied with quantitative models based on evidence accumulation processes. The most prolific example is arguably the drift-diffusion model (DDM). This framework has frequently shown good to very good quantitative fits, which has prompted its wide endorsement. However, fit quality alone does not establish the validity of a model's interpretation. Here, we formally assess the model's validity with a novel cross-validation approach based on the recording of muscular activities, which directly relate to the standard interpretation of various model parameters. Specifically, we recorded electromyographic activity along with response times (RTs), and used it to decompose every RT into 2 components: a premotor time (PMT) and motor time (MT). The latter interval, MT, can be directly linked to motor processes and hence to the nondecision parameter of DDM. In two canonical perceptual decision tasks, we manipulated stimulus strength, speed-accuracy trade-off, and response force and quantified their effects on PMT, MT, and RT. All 3 factors consistently affected MT. The DDM parameter for nondecision processes recovered the MT effects in most situations, with the exception of the fastest responses. The extent of the good fits and the scope of the mis-estimations that we observed allow drawing new limits of the interpretability of model parameters. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Stronger impulse capture and impaired inhibition of prepotent action in children with ADHD performing a Simon task: An electromyographic study.

OBJECTIVE: A deficit in interference control is commonly reported in children with attention deficit hyperactivity disorder (ADHD). This has mainly been interpreted as a difficulty in inhibiting inappropriate responses. However, it could be due to at least two distinct and independent processes, which are often confounded: The activation or suppression of impulsive responses. The aim of the present study was to separate the contribution of these two processes. METHOD: We compared performance of 26 children with ADHD to that of 26 nonADHD children using a novel approach based on electromyographic activity (EMG) analysis. EMG allows two distinct indices to be computed: Incorrect activation rate, which is an index of the intensity of impulse capture and correction rate, which provides a direct measure of the ability to suppress automatic responses. RESULTS: Children with ADHD were slower, committed more errors, and had a larger interference effect than nonADHD children. Moreover, we observed a greater incorrect activation rate and a lower correction rate in the ADHD group. CONCLUSIONS: Our data suggest that the difficulties in interference control found in children with ADHD are explained by both impaired inhibitory processes and a greater propensity to activate automatic responses. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

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.