Sensorimotor prediction is used to direct gaze toward task-relevant locations in a goal-directed throwing task.

Previous research has shown that action effects of self-generated movements are internally predicted before outcome feedback becomes available. To test whether these sensorimotor predictions are used to facilitate visual information uptake for feedback processing, we measured eye movements during the execution of a goal-directed throwing task. Participants could fully observe the effects of their throwing actions (ball trajectory, and either hitting or missing a target) in most of the trials. In a portion of the trials, the ball trajectory was not visible, and participants only received static information about the outcome. We observed a large proportion of predictive saccades, shifting gaze toward the goal region before the ball arrived and outcome feedback became available. Fixation locations after predictive saccades systematically covaried with future ball positions in trials with continuous ball flight information, but notably also in trials with static outcome feedback and only efferent and proprioceptive information about the movement that could be used for predictions. Fixation durations at the chosen positions after feedback onset were modulated by action outcome (longer durations for misses than for hits) and outcome uncertainty (longer durations for narrow vs. clear outcomes). Combining both effects, durations were longest for narrow errors and shortest for clear hits, indicating that the chosen locations offer informational value for feedback processing. Thus, humans are able to use sensorimotor predictions to direct their gaze toward task-relevant feedback locations. Outcome-dependent saccade latency differences (miss vs. hit) indicate that also predictive valuation processes are involved in planning predictive saccades.

Midfrontal theta dynamics index the monitoring of postural stability.

Stepping is a common strategy to recover postural stability and maintain upright balance. Postural perturbations have been linked to neuroelectrical markers such as the N1 potential and theta frequency dynamics. Here, we investigated the role of cortical midfrontal theta dynamics of balance monitoring, driven by balance perturbations at different initial standing postures. We recorded electroencephalography, electromyography, and motion tracking of human participants while they stood on a platform that delivered a range of forward and backward whole-body balance perturbations. The participants' postural threat was manipulated prior to the balance perturbation by instructing them to lean forward or backward while keeping their feet-in-place in response to the perturbation. We hypothesized that midfrontal theta dynamics index the engagement of a behavioral monitoring system and, therefore, that perturbation-induced theta power would be modulated by the initial leaning posture and perturbation intensity. Targeted spatial filtering in combination with mixed-effects modeling confirmed our hypothesis and revealed distinct modulations of theta power according to postural threat. Our results provide novel evidence that midfrontal theta dynamics subserve action monitoring of human postural balance. Understanding of cortical mechanisms of balance control is crucial for studying balance impairments related to aging and neurological conditions (e.g. stroke).

Slip or fallacy? Effects of error severity on own and observed pitch error processing in pianists.

Errors elicit a negative, mediofrontal, event-related potential (ERP), for both own errors (error-related negativity; ERN) and observed errors (here referred to as observer mediofrontal negativity; oMN). It is unclear, however, if the action-monitoring system codes action valence as an all-or-nothing phenomenon or if the system differentiates between errors of different severity. We investigated this question by recording electroencephalography (EEG) data of pianists playing themselves (Experiment 1) or watching others playing (Experiment 2). Piano pieces designed to elicit large errors were used. While active participants' ERN amplitudes differed between small and large errors, observers' oMN amplitudes did not. The different pattern in the two groups of participants was confirmed in an exploratory analysis comparing ERN and oMN directly. We suspect that both prediction and action mismatches can be coded in action monitoring systems, depending on the task, and a need-to-adapt signal is sent whenever mismatches happen to indicate the magnitude of the needed adaptation.

More implicit and more explicit motor imagery tasks for exploring the mental representation of hands and feet in action.

The mental representation of the body in action can be explored using motor imagery (MI) tasks. MI tasks can be allocated along a continuum going from more implicit to more explicit tasks, where the discriminant is the degree of action monitoring required to solve the tasks (which is the awareness of using the mental representation of our own body to monitor our motor imagery). Tasks based on laterality judgments, such as the Hand Laterality Task (HLT) and the Foot Laterality Task (FLT), provide an example of more implicit tasks (i.e., less action monitoring is required). While, an example of a more explicit task is the Mental Motor Chronometry task (MMC) for hands and feet, where individuals are asked to perform or imagine performing movements with their limbs (i.e., more action monitoring is required). In our study, we directly compared hands and feet at all these tasks for the first time, as these body districts have different physical features as well as functions. Fifty-five participants were asked to complete an online version of the HLT and FLT (more implicit measure), and an online version of the MMC task for hands and feet (more explicit measure). The mental representation of hands and feet in action differed only when the degree of action monitoring decreased (HLT ≠ FLT); we observed the presence of biomechanical constraints only for hands. Differently, when the degree of action monitoring increased hands and feet did not show any difference (MMC hands = MMC feet). Our results show the presence of a difference in the mental representation of hands and feet in action that specifically depends on the degree of action monitoring.

Cortical responses to whole-body balance perturbations index perturbation magnitude and predict reactive stepping behavior.

The goal of this study was to determine whether the cortical responses elicited by whole-body balance perturbations were similar to established cortical markers of action monitoring. Postural changes imposed by balance perturbations elicit a robust negative potential (N1) and a brisk increase of theta activity in the electroencephalogram recorded over midfrontal scalp areas. Because action monitoring is a cognitive function proposed to detect errors and initiate corrective adjustments, we hypothesized that the possible cortical markers of action monitoring during balance control (N1 potential and theta rhythm) scale with perturbation intensity and the eventual execution of reactive stepping responses (as opposed to feet-in-place responses). We recorded high-density electroencephalogram from eleven young individuals, who participated in an experimental balance assessment. The participants were asked to recover balance following anteroposterior translations of the support surface at various intensities, while attempting to maintain both feet in place. We estimated source-resolved cortical activity using independent component analysis. Combining time-frequency decomposition and group-level general linear modeling of single-trial responses, we found a significant relation of the interaction between perturbation intensity and stepping responses with multiple cortical features from the midfrontal cortex, including the N1 potential, and theta, alpha, and beta rhythms. Our findings suggest that the cortical responses to balance perturbations index the magnitude of a deviation from a stable postural state to predict the need for reactive stepping responses. We propose that the cortical control of balance may involve cognitive control mechanisms (i.e., action monitoring) that facilitate postural adjustments to maintain postural stability.

Freedom to act enhances the sense of agency, while movement and goal-related prediction errors reduce it.

The Sense of Agency (SoA) is the experience of controlling one's movements and their external consequences. Accumulating evidence suggests that freedom to act enhances SoA, while prediction errors are known to reduce it. Here, we investigated if prediction errors related to movement or to the achievement of the goal of the action exert the same influence on SoA during free and cued actions. Participants pressed a freely chosen or cued-colored button, while observing a virtual hand moving in the same or in the opposite direction-i.e., movement-related prediction error-and pressing the selected or a different color-i.e., goal-related prediction error. To investigate implicit and explicit components of SoA, we collected indirect (i.e., Synchrony Judgments) and direct (i.e., Judgments of Causation) measures. We found that participants judged virtual actions as more synchronous when they were free to act. Additionally, movement-related prediction errors reduced both perceived synchrony and judgments of causation, while goal-related prediction errors impaired exclusively the latter. Our results suggest that freedom to act enhances SoA and that movement and goal-related prediction errors lead to an equivalent reduction of SoA in free and cued actions. Our results also show that the influence of freedom to act and goal achievement may be limited, respectively, to implicit and explicit SoA, while movement information may affect both components. These findings provide support to recent theories that view SoA as a multifaceted construct, by showing that different action cues may uniquely influence the feeling of control.

Contrasting time and frequency domains: ERN and induced theta oscillations differentially predict post-error behavior.

The present study investigated the neural dynamics of error processing in both the time and frequency domains, as well as associated behavioral phenomena, at the single-trial level. We used a technique that enabled us to separately investigate the evoked and induced aspects of the EEG signal (Cohen & Donner, 2013, Journal of Neurophysiology, 110[12], 2752-2763). We found that at the single-trial level, while the (evoked) error-related negativity (ERN) predicted only post-error slowing (PES)-and only when errors occurred on incongruent trials-induced frontal midline theta power served as a robust predictor of both PES and post-error accuracy (PEA) regardless of stimulus congruency. Mediation models of both electrophysiological indices demonstrated that although the relationship between theta and PEA was mediated by PES, there was not a relationship between the ERN and PEA. Our data suggest that although the ERN and frontal midline theta index functionally related underlying cognitive processes, they are not simply the same process manifested in different domains. In addition, our findings are consistent with the adaptive theory of post-error slowing, as PES was positively associated with post-error accuracy at the single-trial level. More generally, our study provides additional support for the inclusion of a time-frequency approach to better understand the role of medial frontal cortex in action monitoring.

Violation of expectations about movement and goal achievement leads to Sense of Agency reduction.

The control of one's own movements and of their impact on the external world generates a feeling of control referred to as Sense of Agency (SoA). SoA is experienced when actions match predictions and is reduced by unpredicted events. The present study investigated the contribution of monitoring two fundamental components of action-movement execution and goal achievement-that have been most often explored separately in previous research. We have devised a new paradigm in which participants performed goal-directed actions while viewing an avatar's hand in a mixed-reality scenario. The hand performed either the same action or a different one, simultaneously or after various delays. Movement of the virtual finger and goal attainment were manipulated, so that they could match or conflict with the participants' expectations. We collected judgments of correspondence (an explicit index of SoA that overcomes the tendency to over-attribute actions to oneself) by asking participants if the observed action was synchronous or not with their action. In keeping with previous studies, we found that monitoring both movement execution and goal attainment is relevant for SoA. Moreover, we expanded previous findings by showing that movement information may be a more constant source of SoA modulation than goal information. Indeed, an incongruent movement impaired SoA irrespective of delay duration, while a missed goal did so only when delays were short. Our novel paradigm allowed us to simultaneously manipulate multiple action features, a characteristic that makes it suitable for investigating the contribution of different sub-components of action in modulating SoA in healthy and clinical populations.

How action selection influences the sense of agency: An ERP study.

Sense of agency (SoA) refers to the feeling that we are in control of our actions and, through them, of events in the outside world. One influential view claims that the SoA depends on retrospectively matching the expected and actual outcomes of action. However, recent studies have revealed an additional, prospective component to SoA, driven by action selection processes. We used event-related potentials (ERPs) to clarify the neural mechanisms underlying prospective agency. Subliminal priming was used to manipulate the fluency of selecting a left or right hand action in response to a supraliminal target. These actions were followed by one of several coloured circles, after a variable delay. Participants then rated their degree of control over this visual outcome. Incompatible priming impaired action selection, and reduced sense of agency over action outcomes, relative to compatible priming. More negative ERPs immediately after the action, linked to post-decisional action monitoring, were associated with reduced agency ratings over action outcomes. Additionally, feedback-related negativity evoked by the outcome was also associated with reduced agency ratings. These ERP components may reflect brain processes underlying prospective and retrospective components of sense of agency respectively.

Cultural influences on neural basis of inhibitory control.

Research on neural basis of inhibitory control has been extensively conducted in various parts of the world. It is often implicitly assumed that neural basis of inhibitory control is universally similar across cultures. Here, we investigated the extent to which culture modulated inhibitory-control brain activity at both cultural-group and cultural-value levels of analysis. During fMRI scanning, participants from different cultural groups (including Caucasian-Americans and Japanese-Americans living in the United States and native Japanese living in Japan) performed a Go/No-Go task. They also completed behavioral surveys assessing cultural values of behavioral consistency, or the extent to which one's behaviors in daily life are consistent across situations. Across participants, the Go/No-Go task elicited stronger neural activity in several inhibitory-control areas, such as the inferior frontal gyrus (IFG) and anterior cingulate cortex (ACC). Importantly, at the cultural-group level, we found variation in left IFG (L-IFG) activity that was explained by a cultural region where participants lived in (as opposed to race). Specifically, L-IFG activity was stronger for native Japanese compared to Caucasian- and Japanese-Americans, while there was no systematic difference in L-IFG activity between Japanese- and Caucasian-Americans. At the cultural-value level, we found that participants who valued being "themselves" across situations (i.e., having high endorsement of behavioral consistency) elicited stronger rostral ACC activity during the Go/No-Go task. Altogether, our findings provide novel insight into how culture modulates the neural basis of inhibitory control.

Midfrontal theta tracks action monitoring over multiple interactive time scales.

Quickly detecting and correcting mistakes is a crucial brain function. EEG studies have identified an idiosyncratic electrophysiological signature of online error correction, termed midfrontal theta. Midfrontal theta has so far been investigated over the fast time-scale of a few hundred milliseconds. But several aspects of behavior and brain activity unfold over multiple time scales, displaying "scale-free" dynamics that have been linked to criticality and optimal flexibility when responding to changing environmental demands. Here we used a novel line-tracking task to demonstrate that midfrontal theta is a transient yet non-phase-locked response that is modulated by task performance over at least three time scales: a few hundred milliseconds at the onset of a mistake, task performance over a fixed window of the previous 5s, and scale-free-like fluctuations over many tens of seconds. These findings provide novel evidence for a role of midfrontal theta in online behavioral adaptation, and suggest new approaches for linking EEG signatures of human executive functioning to its neurobiological underpinnings.

Individual differences in error tolerance in humans: Neurophysiological evidences.

When interacting in error-prone environments, humans display different tolerances to changing their decisions when faced with erroneous feedback information. Here, we investigated whether these individual differences in error tolerance (ET) were reflected in neurophysiological mechanisms indexing specific motivational states related to feedback monitoring. To explore differences in ET, we examined the performance of 80 participants in a probabilistic reversal-learning task. We then compared event-related brain responses (ERPs) of two extreme groups of participants (High ET and Low ET), which showed radical differences in their propensity to maintain newly learned rules after receiving spurious negative feedback. We observed that High ET participants showed reduced anticipatory activity prior to the presentation of incoming feedback, informing them of the correctness of their performance. This was evidenced by measuring the amplitude of the stimulus-preceding negativity (SPN), an ERP component indexing attention and motivational engagement of incoming informative feedback. Postfeedback processing ERP components (the so-called Feedback-Related Negativity and the P300) also showed reduced amplitude in this group (High ET). The general decreased responsiveness of the High ET group to external feedback suggests a higher proneness to favor internal(rule)-based strategies, reducing attention to external cues and the consequent impact of negative evaluations on decision making. We believe that the present findings support the existence of specific cognitive and motivational processes underlying individual differences on error-tolerance among humans, contributing to the ongoing research focused on understanding the mental processes behind human fallibility in error-prone scenarios.

Potentiated processing of negative feedback in depression is attenuated by anhedonia.

BACKGROUND: Although cognitive theories of depression have postulated enhanced processing of negatively valenced information, previous EEG studies have shown both increased and reduced sensitivity for negative performance feedback in MDD. To reconcile these paradoxical findings, it has been speculated that sensitivity for negative feedback is potentiated in moderate MDD, but reduced in highly anhedonic subjects. The goal of this study was to test this hypothesis by analyzing the feedback-related negativity (FRN), frontomedial theta power (FMT), and source-localized anterior midcingulate cortex (aMCC) activity after negative feedback. METHODS: Fourteen unmedicated participants with Major Depressive Disorder (MDD) and 15 control participants performed a reinforcement learning task while 128-channel Electroencephalogram (EEG) was recorded. FRN, FMT, and LORETA source-localized aMCC activity after negative and positive feedback were compared between groups. RESULTS: The MDD group showed higher FRN amplitudes and aMCC activation to negative feedback than controls. Moreover, aMCC activation to negative feedback was inversely related to self-reported anhedonia. In contrast, self-reported anxiety correlated with feedback-evoked frontomedial theta (FMT) within the depression group. CONCLUSIONS: The present findings suggest that, among depressed and anxious individuals, enhanced processing of negative feedback occurs relatively early in the information processing stream. These results extend prior work and indicate that although moderate depression is associated with elevated sensitivity for negative feedback, high levels of anhedonia may attenuate this effect.

Metacognition of agency and theory of mind in adults with high functioning autism.

We investigated metacognition of agency in adults with high functioning autism or Asperger Syndrome (HFA/AS) using a computer task in which participants moved the mouse to get the cursor to touch the downward moving X's and avoid the O's. They were then asked to make judgments of performance and judgments of agency. Objective control was either undistorted, or distorted by adding turbulence (i.e., random noise) or a time Lag between the mouse and cursor movements. Participants with HFA/AS used sensorimotor cues available in the turbulence and lag conditions to a lesser extent than control participants in making their judgments of agency. Furthermore, the failure to use these internal diagnostic cues to their own agency was correlated with decrements in a theory of mind task. These findings suggest that a reduced sensitivity to veridical internal cues about the sense of agency is related to mentalizing impairments in autism.

Temporal unpredictability increases error monitoring as revealed by EEG-EMG investigation.

Reacting in an unpredictable context increases error monitoring as evidenced by greater error-related negativity (ERN), an electrophysiological marker linked to an evaluation of response outcomes. We investigated whether ERN also increased when participants evaluated their responses to events that appeared in unpredictable versus predictable moments in time. We complemented electroencephalographic (EEG) analysis of cortical activity by measuring performance monitoring processes at the peripheral level using electromyography (EMG). Specifically, we used EMG data to quantify how temporal unpredictability would affect motor time (MT), the interval between the onset of muscle activity, and the mechanical response. MT increases following errors, indexing online error detection, and an attempt to stop incorrect actions. In our temporally cued version of the stop-signal task, symbolic cues predicted (temporally predictable condition) or not (temporally unpredictable condition) the onset of a target. In 25% of trials, an auditory signal occurred shortly after the target presentation, informing participants that they should inhibit their response completely. Response times were slower, and fewer inhibitory errors were made during temporally unpredictable than predictable trials, indicating enhanced control of unwanted actions when target onset time was unknown. Importantly, the ERN to inhibitory errors was greater in temporally unpredictable relative to temporally predictable conditions. Similarly, EMG data revealed prolonged MT when reactions to temporally unpredictable targets had not been stopped. Taken together, our results show that a temporally unpredictable environment increases the control of unwanted actions, both at cortical and peripheral levels, suggesting a higher subjective cost of maladaptive responses to temporally uncertain events.

Making precise movements increases confidence in perceptual decisions.

Perceptual decision-making often lacks explicit feedback, making confidence in our choices pivotal for guiding subsequent actions. Recent studies have highlighted the role of motor responses in modulating decision confidence. Two competing mechanisms have been proposed to elucidate this phenomenon. The "fluency hypothesis" posits that the ease and smoothness of executing a motor response can serve as a cue to enhance retrospective confidence. Conversely, the "monitoring hypothesis" suggests that the extent of action monitoring during response selection may boost retrospective confidence, with heightened monitoring potentially offsetting response fluency. We conducted a pre-registered experiment to directly test these hypotheses. Participants engaged in a perceptual task involving the discrimination of Gabor patch orientation. Perceptual responses required high or low motor precision, manipulated by the size of target circles that participants had to reach with the computer mouse to provide a response. Contrary to the "fluency hypothesis", our results showed that, in trials requiring higher precision (utilizing small circles), participants reported higher confidence levels compared to trials with less demanding responses (involving larger circles). Importantly, this increase in confidence did not coincide with any change in perceptual accuracy. These findings align with the "monitoring hypothesis," suggesting that the degree of action monitoring during response execution can indeed influence retrospective decision confidence.

The cerebellum monitors errors and entrains executive networks.

Frontal midline θ (Fmθ) activity occurs in medial prefrontal cortices (mPFC), when expected and actual outcomes conflict. Cerebellar forward models could inform the mPFC about this mismatch. To verify this hypothesis we correlated the mPFC activation during a visuomotor tracking task (VM) with performance accuracy, in control and cerebellum-lesioned participants. Additionally, purely visual (V), motor (M) and a motor plus visual tasks (V + M) were performed. An Independent Component, with a mid-frontal topography scalp map and equivalent dipole location in the dorsal anterior cingulate cortex accounted for Fmθ. In control participants Fmθ power increased during VM, when the error level crossed a threshold, but not during V + M, M and V. This increase scaled with tracking error. Fmθ power failed to increase during VM in cerebellar participants, even at highest tracking errors. Thus, in control participants, activation of mPFC is induced when a continuous monitoring effort for online error detection is required. The presence of a threshold error for enhancing Fmθ, suggests the switch from an automatic to an executive tracking control, which recruits the mPFC. Given that the cerebellum stores forward models, the absence of Fmθ increases during tracking errors in cerebellar participants indicates that cerebellum is necessary for supplying the mPFC with prediction error-related information. This occurs when automatic control falters, and a deliberate correction mechanism needs to be triggered. Further studies are needed to verify if this alerting function also occurs in the context of the other cognitive and non-cognitive functions in which the cerebellum is involved.

Visual sensitivity at the service of action control in posterior parietal cortex.

The posterior parietal cortex (PPC) serves as a crucial hub for the integration of sensory with motor cues related to voluntary actions. Visual input is used in different ways along the dorsomedial and the dorsolateral visual pathways. Here we focus on the dorsomedial pathway and recognize a visual representation at the service of action control. Employing different experimental paradigms applied to behaving monkeys while single neural activity is recorded from the medial PPC (area V6A), we show how plastic visual representation can be, matching the different contexts in which the same object is proposed. We also present data on the exchange between vision and arm actions and highlight how this rich interplay can be used to weight different sensory inputs in order to monitor and correct arm actions online. Indeed, neural activity during reaching or reach-to-grasp actions can be excited or inhibited by visual information, suggesting that the visual perception of action, rather than object recognition, is the most effective factor for area V6A. Also, three-dimensional object shape is encoded dynamically by the neural population, according to the behavioral context of the monkey. Along this line, mirror neuron discharges in V6A indicate the plasticity of visual representation of the graspable objects, that changes according to the context and peaks when the object is the target of one's own action. In other words, object encoding in V6A is a visual encoding for action.

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.

Perfectionism-related variations in error processing in a task with increased response selection complexity.

Perfectionists strive for a flawless performance because they are intrinsically motivated to set and achieve high goals (personal standards perfectionism; PSP) and/or because they are afraid to be negatively evaluated by others (evaluative concern perfectionism; ECP). We investigated the differential relationships of these perfectionism dimensions with performance, post-response adaptation, error processing (reflected by two components of the event-related potential: error/correct negativity - Ne/c; error/correct positivity - Pe/c) and error detection. In contrast to previous studies, we employed a task with increased response selection complexity providing more room for perfectionistic dispositions to manifest themselves. Although ECP was related to indicators of increased preoccupation with errors, high-EC perfectionists made more errors than low-EC perfectionists. This observation may be explained by insufficient early error processing as indicated by a reduced Ne/c effect and a lack of post-response adaptation. PSP had a moderating effect on the relationship between ECP and early error processing. Our results provide evidence that pure-EC perfectionists may spend many of their cognitive resources on error-related contents and worrying, leaving less capacity for cognitive control and thus producing a structural lack of error processing.