Expectancy and attention bias to spiders: Dissecting anticipation and allocation processes using ERPs.

The current registered report focused on the temporal dynamics of the relationship between expectancy and attention toward threat, to better understand the mechanisms underlying the prioritization of threat detection over expectancy. In the current event-related potentials experiment, a-priori expectancy was manipulated, and attention bias was measured, using a well-validated paradigm. A visual search array was presented, with one of two targets: spiders (threatening) or birds (neutral). A verbal cue stating the likelihood of encountering a target preceded the array, creating congruent and incongruent trials. Following cue presentation, preparatory processes were examined using the contingent negative variation (CNV) component. Following target presentation, two components were measured: early posterior negativity (EPN) and late positive potential (LPP), reflecting early and late stages of natural selective attention toward emotional stimuli, respectively. Behaviorally, spiders were found faster than birds, and congruency effects emerged for both targets. For the CNV, a non-significant trend of more negative amplitudes following spider cues emerged. As expected, EPN and LPP amplitudes were larger for spider targets compared to bird targets. Data-driven, exploratory, topographical analyses revealed different patterns of activation for bird cues compared to spider cues. Furthermore, 400-500 ms post-target, a congruency effect was revealed only for bird targets. Together, these results demonstrate that while expectancy for spider appearance is evident in differential neural preparation, the actual appearance of spider target overrides this expectancy effect and only in later stages of processing does the cueing effect come again into play.

Modulatory effects of goal relevance on emotional attention reveal that fear has a distinct value.

Threat-related stimuli can capture attention. However, it remains debated whether this capture is automatic or not. To address this question, we compared attentional biases to emotional faces using a dot-probe task (DPT) where emotion was never goal-relevant (Experiment 1) or made directly task-relevant by means of induction trials (Experiments 2-3). Moreover, the contingency between the DPT and induction trials was either partial (Experiment 2) or full (Experiment 3). Eye-tracking was used to ascertain that the emotional cue and the subsequent target were processed with peripheral vision. Experiments 1 and 2 both showed that negative faces captured attention, with faster target processing when it appeared on the same side as the preceding fearful face (i.e. fear-valid trials) compared to the opposite side where the neutral face was shown (i.e. fear-invalid trials), but also when it appeared on the side of the preceding neutral face (i.e. happy-invalid trials) compared to the happy face (i.e. happy-valid trials). Importantly, this preferential spatial orienting to negative emotion was not observed in Experiment 3, where the goal relevance of emotion was high. However, in that experiment, fearful faces produced a specific attentional bias during the DPT, which was mostly driven by the induction trials themselves.

Top-Down Modulation of Early Visual Processing in V1: Dissociable Neurophysiological Effects of Spatial Attention, Attentional Load and Task-Relevance.

Until today, there is an ongoing discussion if attention processes interact with the information processing stream already at the level of the C1, the earliest visual electrophysiological response of the cortex. We used two highly powered experiments (each N = 52) and examined the effects of task relevance, spatial attention, and attentional load on individual C1 amplitudes for the upper or lower visual hemifield. Bayesian models revealed evidence for the absence of load effects but substantial modulations by task-relevance and spatial attention. When the C1-eliciting stimulus was a task-irrelevant, interfering distracter, we observed increased C1 amplitudes for spatially unattended stimuli. For spatially attended stimuli, different effects of task-relevance for the two experiments were found. Follow-up exploratory single-trial analyses revealed that subtle but systematic deviations from the eye-gaze position at stimulus onset between conditions substantially influenced the effects of attention and task relevance on C1 amplitudes, especially for the upper visual field. For the subsequent P1 component, attentional modulations were clearly expressed and remained unaffected by these deviations. Collectively, these results suggest that spatial attention, unlike load or task relevance, can exert dissociable top-down modulatory effects at the C1 and P1 levels.

The selective use of punishments on congruent versus incongruent trials in the Stroop task.

Conflict adaptation refers to the dynamic modulation of conflict processing across successive trials and reflects improved cognitive control. Interestingly, aversive motivation can increase conflict adaptation, although it remains unclear through which process this modulation occurs because previous studies presented punishment feedback following suboptimal performance on both congruent and incongruent trials. According to integrative accounts of conflict monitoring and aversive motivation in the dorsal anterior cingulate cortex, punishment feedback following slow or erroneous performance on incongruent trials in particular should lead to improved conflict adaptation. Second, selectively increasing motivation on incongruent trials should reduce the overall congruency effect. The current study sought to test both hypotheses. Specifically, we administered the confound-minimized Stroop task to a large group of participants and manipulated the position of feedback (following either congruent or incongruent trials) and aversive motivation (tied to a monetary loss or not) across different blocks. As expected, the congruency effect was found to be smaller when punishment was coupled with incongruent versus congruent trials. However, results showed that conflict adaptation was increased when punishment feedback was selectively coupled with congruent rather than incongruent trials. Together, these results suggest that aversive motivation does not uniformly improve cognitive control but this gain appears to be context dependent.

Reduced flexibility of cognitive control: reactive, but not proactive control, underpins the congruency sequence effect.

The congruency sequence effect (CSE) refers to facilitated conflict processing following incongruent than congruent trials, and reflects enhanced cognitive control during conflict processing. Although this effect is mostly conceived as being reactive, proactive control can also unlock it under specific circumstances according to previous studies (e.g., when an informative cue is used). However, whether or not humans can flexibly switch between these two complementing control modes remains unclear. To address this question, 55 participants completed the confound-minimized Stroop task in different blocks where the cue about the upcoming trial's congruency was either informative or not, and orthogonally to it, the cue-stimulus interval (CSI) was either short or long. We tested if the size of the CSE could change depending on the specific combination of these two factors, which would indicate that cognitive control depends on the subtle balance between reactive and proactive control, and is therefore flexible. However, results showed that the CSE was significant and comparable across the four combinations of CSI and Cue type, suggesting that it primarily stemmed from reactive control. We discuss our results against the dual mechanism of control (DMC) framework (Braver in Trends Cogn Sci 16:106-113, 2012).

Task Learnability Modulates Surprise but Not Valence Processing for Reinforcement Learning in Probabilistic Choice Tasks.

The goal of temporal difference (TD) reinforcement learning is to maximize outcomes and improve future decision-making. It does so by utilizing a prediction error (PE), which quantifies the difference between the expected and the obtained outcome. In gambling tasks, however, decision-making cannot be improved because of the lack of learnability. On the basis of the idea that TD utilizes two independent bits of information from the PE (valence and surprise), we asked which of these aspects is affected when a task is not learnable. We contrasted behavioral data and ERPs in a learning variant and a gambling variant of a simple two-armed bandit task, in which outcome sequences were matched across tasks. Participants were explicitly informed that feedback could be used to improve performance in the learning task but not in the gambling task, and we predicted a corresponding modulation of the aspects of the PE. We used a model-based analysis of ERP data to extract the neural footprints of the valence and surprise information in the two tasks. Our results revealed that task learnability modulates reinforcement learning via the suppression of surprise processing but leaves the processing of valence unaffected. On the basis of our model and the data, we propose that task learnability can selectively suppress TD learning as well as alter behavioral adaptation based on a flexible cost-benefit arbitration.

Children's automatic evaluation of self-generated actions is different from adults.

Performance monitoring (PM) is central to learning and decision making. It allows individuals to swiftly detect deviations between actions and intentions, such as response errors, and adapt behavior accordingly. Previous research showed that in adult participants, error monitoring is associated with two distinct and robust behavioral effects. First, a systematic slowing down of reaction time speed is typically observed following error commission, which is known as post-error slowing (PES). Second, response errors have been reported to be automatically evaluated as negative events in adults. However, it remains unclear whether (1) children process response errors as adults do (PES), (2) they also evaluate them as negative events, and (3) their responses vary according to the pedagogy experienced. To address these questions, we adapted a simple decision-making task previously validated in adults to measure PES as well as the affective processing of response errors. We recruited 8- to 12-year-old children enrolled in traditional (N = 56) or Montessori (N = 45) schools, and compared them to adults (N = 46) on the exact same task. Results showed that children processed correct actions as positive events, and that adults processed errors as negative events. By contrast, PES was similarly observed in all groups. Moreover, the former effect was observed in traditional schoolchildren, but not in Montessori schoolchildren. These findings suggest that unlike PES, which likely reflects an age-invariant attention orienting toward response errors, their affective processing depends on both age and pedagogy.

Neurophysiological evidence for evaluative feedback processing depending on goal relevance.

Feedback signaling the success or failure of actions is readily exploited to implement goal-directed behavior. Two event-related brain potentials (ERPs) have been identified as reliable markers of evaluative feedback processing: the Feedback-Related Negativity (FRN) and the P3. Recent ERP studies have shown a substantial reduction of these components when the feedback's goal relevance (in terms of goal informativeness) was decreased. However, it remains unclear whether this lowering of evaluative feedback processing at the FRN and P3 levels (i) reflects a common regulation process operating across them or (ii) indirectly and mostly depends on valence processing. To address these questions, 44 participants performed a time estimation task wherein the perceived goal relevance of the feedback following each decision was manipulated via instructions in different blocks. We recorded 64-channel EEG and collected subjective ratings of feedback valence and relevance, separately for goal-relevant and irrelevant conditions. ERP results showed a substantial reduction of the FRN and P3 components for irrelevant than relevant feedback, despite the balanced task relevance between them. Moreover, a Principal Component Analysis (PCA) showed that these two successive ERP effects had dissociable spatiotemporal properties. Crucially, a multivariate multiple regression analysis revealed that goal relevance per se, but not valence, was the unique significant predictor of the amplitude reduction of the FRN and P3 when the feedback was goal irrelevant. Our results suggest that although these ERP components exhibit non-overlapping spatiotemporal properties and performance monitoring effects, they can both be modulated by a common, valence-unspecific process related to goal relevance.

Modulatory effects of positive mood and approach motivation on reward processing: Two sides of the same coin?

In a previous study (Paul & Pourtois, 2017), we found that positive mood substantially influenced the neural processing of reward, mostly by altering expectations and creating an optimistic bias. Under positive mood, the Reward Positivity (RewP) component and fronto-medial theta activity (FMθ) in response to monetary feedback were both changed compared with neutral mood. Nevertheless, whether positive valence per se or motivational intensity drove these neurophysiological effects remained unclear. To address this question, we combined a mindset manipulation with an imagery procedure to create and maintain three different affective states using a between-subjects design: a neutral mood, and positive mood with either high or low motivational intensity. After mood induction, 161 participants performed a simple gambling task while 64-channel EEG was recorded. FMθ activity results showed that irrespective of motivational intensity, positive compared with neutral mood altered reward expectancy. By comparison, RewP was not affected by positive mood nor motivational intensity. These results suggest that positive mood, rather than motivational intensity, is likely driving the change in reward expectation during gambling, which could reflect the presence of an optimistic bias. Moreover, at the methodological level, they confirm that the RewP ERP component and FMθ activity can capture dissociable effects during reward processing.

Abnormal approach-related motivation but spared reinforcement learning in MDD: Evidence from fronto-midline Theta oscillations and frontal Alpha asymmetry.

Major depression is characterized by abnormal reward processing and reinforcement learning (RL). This impairment might stem from deficient motivation processes, in addition to reduced reward sensitivity. In this study, we recorded 64-channel EEG in a large cohort of major depressive disorder (MDD) patients and matched healthy controls (HC) while they performed a standard RL task. Participants were asked to discover, by trial and error, several hidden stimulus-response associations having different reward probabilities, as enforced using evaluative feedback. We extracted induced fronto-midline Theta (FMT) power time-locked to the response and feedback as neurophysiological index of RL. Furthermore, we assessed approach-related motivation by measuring frontal alpha asymmetry concurrently. At the behavioral level, MDD patients and HCs showed comparable RL. At the EEG level, FMT power systematically varied as a function of reward probability, with opposing effects found at the response and feedback levels. Although this global pattern was spared in MDD, at the feedback level these patients showed however a steep FMT power decrease across trials when reward probability was low. Moreover, they showed impaired approach-related motivation during task execution, as reflected by frontal Alpha asymmetry. These results suggest a dissociation between (globally spared) RL and (impaired) approach motivation in MDD.

Behavioral and electrophysiological responses to fairness norm violations in antisocial offenders.

Antisocial personality disorder is characterized by a stable, lifelong pattern of disregard for and violation of others' rights. Disruptions in the representation of fairness norms may represent a key mechanism in the development and maintenance of this disorder. Here, we investigated fairness norm considerations and reactions to their violations. To examine electrophysiological correlates, we assessed the medial frontal negativity (MFN), an event-related potential previously linked to violations of social expectancy and norms. Incarcerated antisocial violent offenders (AVOs, n = 25) and healthy controls (CTLs, n = 24) acted as proposers in the dictator game (DG) and ultimatum game (UG) and received fair vs. unfair UG offers from either another human (social context) or a computer (non-social context). Results showed that AVOs made lower offers in the DG but not the UG, indicating more rational and strategic behavior. Most importantly, when acting as recipients in the UG, acceptance rates were modulated by social context in CTLs, while AVOs generally accepted more offers. Correspondingly, ERP data indicated pronounced MFN amplitudes following human offers in CTLs, whereas MFN amplitudes in AVOs were generally reduced. The current data suggest intact fairness norm representations but altered reactions to their violation in antisocial personality disorder.

Reappraising cognitive control: normal reactive adjustments following conflict processing are abolished by proactive emotion regulation.

The congruency sequence effect (CSE) reflected by the influence of the congruency of the previous trial on the current one translates improved cognitive control (CC). Yet, it remains debated whether reactive or proactive control processes mostly contribute to this effect. To address this question, we administered a Stroop task controlling for effects of feature repetition and contingency learning to a large group of participants, where we manipulated the frequency of incongruent trials in a block-wise fashion to induce either proactive (high-conflict frequency) or reactive (low-conflict frequency) control. Moreover, as the presentation of trial-by-trial evaluative feedback could influence control processes operating at a local level, we compared effect of evaluative vs. neutral feedback on the CSE, for each control mode separately. We tested the prediction that CSE should be influenced by conflict frequency and feedback type concurrently. Results showed that when evaluative feedback was used, the CSE was increased if conflict frequency was low, confirming that the CSE stemmed from reactive control mainly. If conflict frequency was high, a different sequence effect was observed. The use of neutral feedback abolished the modulation of the CSE by conflict frequency. Moreover, correlation results showed that reappraisal, corresponding to a proactive emotion regulation strategy, was negatively related to the CSE in this condition, suggesting that proactive control can alleviate the reactive dominance of the CSE. Altogether, these results suggest that CC is flexible, and its expression depends on the subtle balance between proactive and reactive control processes.

Integration of reward with cost anticipation during performance monitoring revealed by ERPs and EEG spectral perturbations.

Effort expenditure has an aversive connotation and it can lower hedonic feelings. In this study, we explored the electrophysiological correlates of the complex interplay of reward processing with cost anticipation. To this aim, healthy adult participants performed a gambling task where the outcome (monetary reward vs. no-reward) and its expectancy were manipulated on a trial by trial basis while 64-channel EEG was recorded. Crucially, on some trials, the no-reward outcome could be transformed to a rewarding one, pending effort expenditure by means of an orthogonal dot clicking task, enabling us to compare at the electrophysiological level reward processing when cost was anticipated or not. We extracted and compared different markers of reward processing at the feedback level using both classical ERPs and EEG spectral perturbations in specific bands (theta, delta and beta-gamma). At the behavioral level, participants reported enhanced pleasure and relief when the outcome was rewarding but effort expenditure could be avoided, relative to a control condition where the outcome was rewarding but no extra effort was anticipated. In this condition, EEG results showed a larger Reward Positivity ERP component and increased power in the Delta and Beta-gamma bands. By comparison, cost anticipation did not influence the processing of the no-reward outcome at the FRN and frontal midline theta levels. All together, these neurophysiological results suggest that effort avoidance is associated with increased reward processing.

Goals matter: Amplification of the motivational significance of the feedback when goal impact is increased.

Performance monitoring (PM) entails the continuous evaluation of actions and their outcomes. At the electrophysiological level, PM has been consistently related to two event-related brain potentials (ERPs): the Feedback-Related Negativity (FRN) and the P3. In a previous within-subject crossover design study, we showed that feedback's goal impact (i.e., its importance to one's goal) modulated these ERP components, yet in opposing directions. Although high goal impact was associated with a larger P3, the preceding FRN had a lower amplitude than in the low impact condition. We sought to extend these findings here by adopting a between-subjects design for a pure goal impact manipulation. Sixty-eight participants completed a Go/No Go Task while 64-channel electroencephalography was recorded concurrently. They were randomly assigned to either a high or low goal impact condition, manipulated through instructions on the supposed task's diagnosticity, while reward probability was kept similar between conditions. Replicating our previous results, we found that high goal impact yielded a marginally lower FRN, but substantially larger P3 during PM than low goal impact, without arousal or performance differences. Moreover, a principal component analysis confirmed these opposing directions of goal impact modulation. Overall, these results dovetail with the assumption that goal impact influences PM processes.

Cue-target contingencies modulate voluntary orienting of spatial attention: dissociable effects for speed and accuracy.

Voluntary orienting of spatial attention is typically investigated by visually presented directional cues, which are called predictive when they indicate where the target is more likely to appear. In this study, we investigated the nature of the potential link between cue predictivity (the proportion of valid trials) and the strength of the resulting covert orienting of attention. Participants judged the orientation of a unilateral Gabor grating preceded by a centrally presented, non-directional, color cue, arbitrarily prompting a leftwards or rightwards shift of attention. Unknown to them, cue predictivity was manipulated across blocks, whereby the cue was only predictive for either the first or the second half of the experiment. Our results show that the cueing effects were strongly influenced by the change in predictivity. This influence differently emerged in response speed and accuracy. The speed difference between valid and invalid trials was significantly larger when cues were predictive, and the amplitude of this effect was modulated at the single trial level by the recent trial history. Complementary to these findings, accuracy revealed a robust effect of block history and also a different time-course compared with speed, as if it mainly mirrored voluntary processes. These findings, obtained with a new manipulation and using arbitrary non-directional cueing, demonstrate that cue-target contingencies strongly modulate the way attention is deployed in space.

Ensemble representation for multiple facial expressions: Evidence for a capacity limited perceptual process.

We tested the processing capacity of establishing ensemble representation for multiple facial expressions using the simultaneous-sequential paradigm. Each set consisted of 16 faces conveying a variable amount of happy and angry expressions. Participants judged on a continuous scale the perceived average emotion from each face set (Experiment 1). In the simultaneous condition, the 16 faces were presented concurrently; in the sequential condition, two sets, each containing eight faces, were presented successively. Results showed that judgments varied depending on the number of happy versus angry faces contained in the sets and were sensitive at the single trial level to the perceived mean emotion intensity (based on postexperiment ratings), providing evidence of a genuine mean representation rather than the mere use of a single face or enumeration. Experiments 2 and 3 replicated Experiment 1, but implemented a different response format (binary choices) and added masks following each display, respectively. Importantly, in all three experiments, performance was consistently better in the sequential than in the simultaneous condition, revealing a limited-capacity process. A set of control analyses ruled out the use of enumeration or mere subsampling by the participants to perform the task. Collectively, these results indicate that participants could readily extract mean emotion from multiple faces shown concurrently in a set, but this process is best conceived as being capacity limited.

Conflict-driven adaptive control is enhanced by integral negative emotion on a short time scale.

Negative emotion influences cognitive control, and more specifically conflict adaptation. However, discrepant results have often been reported in the literature. In this study, we broke down negative emotion into integral and incidental components using a modern motivation-based framework, and assessed whether the former could change conflict adaptation. In the first experiment, we manipulated the duration of the inter-trial-interval (ITI) to assess the actual time-scale of this effect. Integral negative emotion was induced by using loss-related feedback contingent on task performance, and measured at the subjective and physiological levels. Results showed that conflict-driven adaptive control was enhanced when integral negative emotion was elicited, compared to a control condition without changes in defensive motivation. Importantly, this effect was only found when a short, as opposed to long ITI was used, suggesting that it had a short time scale. In the second experiment, we controlled for effects of feature repetition and contingency learning, and replicated an enhanced conflict adaptation effect when integral negative emotion was elicited and a short ITI was used. We interpret these new results against a standard cognitive control framework assuming that integral negative emotion amplifies specific control signals transiently, and in turn enhances conflict adaptation.

Motivational Salience Modulates Early Visual Cortex Responses across Task Sets.

Motivationally relevant stimuli benefit from strengthened sensory processing. It is unclear, however, if motivational value of positive and negative valence has similar or dissociable effects on early visual processing. Moreover, whether these perceptual effects are task-specific, stimulus-specific, or more generally feature-based is unknown. In this study, we compared the effects of positive and negative motivational value on early sensory processing using ERPs. We tested the extent to which these effects could generalize to new task contexts and to stimuli sharing common features with the motivationally significant ones. At the behavioral level, stimuli paired with positive incentives were learned faster than stimuli paired with neutral or negative outcomes. The ERP results showed that monetary loss elicited higher neural activity in V1 (at the C1 level) compared with reward, whereas the latter influenced postperceptual processing stages (P300). Importantly, the early loss-related effect generalized to new contexts and to new stimuli with common features, whereas the later reward effects did not spill over to the new context. These results suggest that acquired negative motivational salience can influence early sensory processing by means of plastic changes in feature-based processing in V1.

Modulatory effects of happy mood on performance monitoring: Insights from error-related brain potentials.

Goal-adaptive behavior requires the rapid detection of conflicts between actions and intentions or goals. Although many studies have focused in the past on the influence of negative affect on this cognitive control process (and more specifically, on error monitoring), little is known about the possible modulatory effects of positive affect on it. To address this question, we used a standard (positive) mood induction procedure (based on guided imagery) and asked participants to carry out a speeded go/no-go task while high-density electroencephalography was recorded concurrently. As a control condition, we used a group with neutral mood. Event-related potential results showed that the error-related negativity (ERN) component, reflecting early error detection within the dorsal anterior cingulate cortex, was not influenced by happy mood. In contrast, the subsequent error positivity (Pe) component, related to the appraisal of the motivational significance of errors, was reliably smaller in the happy than in the neutral mood group. Complementing source localization analyses showed that this effect was explained by decreased activation within the posterior cingulate and insular cortices. These results were obtained in the absence of group differences regarding behavioral performance and tonic arousal. These findings suggest that happy mood likely decreases and changes the motivational significance of worse-than-expected events (Pe), while leaving their earlier automatic detection (ERN) unaltered. We discuss these new results in terms of dynamic changes in the complex interplay of performance monitoring with motivation.

Spared internal but impaired external reward prediction error signals in major depressive disorder during reinforcement learning.

BACKGROUND: Major depressive disorder (MDD) creates debilitating effects on a wide range of cognitive functions, including reinforcement learning (RL). In this study, we sought to assess whether reward processing as such, or alternatively the complex interplay between motivation and reward might potentially account for the abnormal reward-based learning in MDD. METHODS: A total of 35 treatment resistant MDD patients and 44 age matched healthy controls (HCs) performed a standard probabilistic learning task. RL was titrated using behavioral, computational modeling and event-related brain potentials (ERPs) data. RESULTS: MDD patients showed comparable learning rate compared to HCs. However, they showed decreased lose-shift responses as well as blunted subjective evaluations of the reinforcers used during the task, relative to HCs. Moreover, MDD patients showed normal internal (at the level of error-related negativity, ERN) but abnormal external (at the level of feedback-related negativity, FRN) reward prediction error (RPE) signals during RL, selectively when additional efforts had to be made to establish learning. CONCLUSIONS: Collectively, these results lend support to the assumption that MDD does not impair reward processing per se during RL. Instead, it seems to alter the processing of the emotional value of (external) reinforcers during RL, when additional intrinsic motivational processes have to be engaged.