Fifty Percent of the Time, Tones Come Every Time: Stronger Prediction Error Effects on Neurophysiological Sensory Attenuation for Self-generated Tones.

The N1/P2 amplitude reduction for self-generated tones in comparison to external tones in EEG, which has recently also been described for action observation, is an example of the so-called sensory attenuation. Whether this effect is dependent on motor-based or general predictive mechanisms is unclear. Using a paradigm, in which actions (button presses) elicited tones in only half the trials, this study examined how the processing of the tones is modulated by the prediction error in each trial in a self-performed action compared with action observation. In addition, we considered the effect of temporal predictability by adding a third condition, in which visual cues were followed by external tones in half the trials. The attenuation result patterns differed for N1 and P2 amplitudes, but neither showed an attenuation effect beyond temporal predictability. Interestingly, we found that both N1 and P2 amplitudes reflected prediction errors derived from a reinforcement learning model, in that larger errors coincided with larger amplitudes. This effect was stronger for tones following button presses compared with cued external tones, but only for self-performed and not for observed actions. Taken together, our results suggest that attenuation effects are partially driven by general predictive mechanisms irrespective of self-performed actions. However, the stronger prediction-error effects for self-generated tones suggest that distinct motor-related factors beyond temporal predictability, potentially linked to reinforcement learning, play a role in the underlying mechanisms. Further research is needed to validate these initial findings as the calculation of the prediction errors was limited by the design of the experiment.

The Role of the Human Cerebellum for Learning from and Processing of External Feedback in Non-Motor Learning: A Systematic Review.

This review aimed to systematically identify and comprehensively review the role of the cerebellum in performance monitoring, focusing on learning from and on processing of external feedback in non-motor learning. While 1078 articles were screened for eligibility, ultimately 36 studies were included in which external feedback was delivered in cognitive tasks and which referenced the cerebellum. These included studies in patient populations with cerebellar damage and studies in healthy subjects applying neuroimaging. Learning performance in patients with different cerebellar diseases was heterogeneous, with only about half of all patients showing alterations. One patient study using EEG demonstrated that damage to the cerebellum was associated with altered neural processing of external feedback. Studies assessing brain activity with task-based fMRI or PET and one resting-state functional imaging study that investigated connectivity changes following feedback-based learning in healthy participants revealed involvement particularly of lateral and posterior cerebellar regions in processing of and learning from external feedback. Cerebellar involvement was found at different stages, e.g., during feedback anticipation and following the onset of the feedback stimuli, substantiating the cerebellum's relevance for different aspects of performance monitoring such as feedback prediction. Future research will need to further elucidate precisely how, where, and when the cerebellum modulates the prediction and processing of external feedback information, which cerebellar subregions are particularly relevant, and to what extent cerebellar diseases alter these processes.

Emotional cues reduce Pavlovian interference in feedback-based go and nogo learning.

It is easier to execute a response in the promise of a reward and withhold a response in the promise of a punishment than vice versa, due to a conflict between cue-related Pavlovian and outcome-related instrumental action tendencies in the reverse conditions. This robust learning asymmetry in go and nogo learning is referred to as the Pavlovian bias. Interestingly, it is similar to motivational tendencies reported for affective facial expressions, i.e., facilitation of approach to a smile and withdrawal from a frown. The present study investigated whether and how learning from emotional faces instead of abstract stimuli modulates the Pavlovian bias in reinforcement learning. To this end, 137 healthy adult participants performed an orthogonalized Go/Nogo task that fully decoupled action (go/nogo) and outcome valence (win points/avoid losing points). Three groups of participants were tested with either emotional facial cues whose affective valence was either congruent (CON) or incongruent (INC) to the required instrumental response, or with neutral facial cues (NEU). Relative to NEU, the Pavlovian bias was reduced in both CON and INC, though still present under all learning conditions. Importantly, only for CON, the reduction of the Pavlovian bias effect was adaptive by improving learning performance in one of the conflict conditions. In contrast, the reduction of the Pavlovian bias in INC was completely driven by decreased learning performance in non-conflict conditions. These results suggest a potential role of arousal/salience in Pavlovian-instrumental regulation and cue-action congruency in the adaptability of goal-directed behavior. Implications for clinical application are discussed.

When a Sunny Day Gives You Butterflies: An Electrophysiological Investigation of Concreteness and Context Effects in Semantic Word Processing.

Theories on controlled semantic cognition assume that word concreteness and linguistic context interact during semantic word processing. Methodological approaches and findings on how this interaction manifests at the electrophysiological and behavioral levels are heterogeneous. We measured ERPs and RTs applying a validated cueing paradigm with 19 healthy participants, who performed similarity judgments on concrete or abstract words (e.g., "butterfly" or "tolerance") after reading contextual and irrelevant sentential cues. Data-driven analyses showed that concreteness increased and context decreased negative-going deflections in broadly distributed bilateral clusters covering the N400 and N700/late positive component time range, whereas both reduced RTs. Crucially, within a frontotemporal cluster in the N400 time range, contextual (vs. irrelevant) information reduced negative-going amplitudes in response to concrete but not abstract words, whereas a contextual cue reduced RTs only in response to abstract but not concrete words. The N400 amplitudes did not explain additional variance in the RT data, which showed a stronger contextual facilitation for abstract than concrete words. Our results support separate but interacting effects of concreteness and context on automatic and controlled stages of contextual semantic processing and suggest that effects on the electrophysiological versus behavioral level obtained with this paradigm are dissociated.

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.

My view on your actions: Dynamic changes in viewpoint-dependent auditory ERP attenuation during action observation.

It has been suggested that during action observation, a sensory representation of the observed action is mapped onto one's own motor system. However, it is largely unexplored what this may imply for the early processing of the action's sensory consequences, whether the observational viewpoint exerts influence on this and how such a modulatory effect might change over time. We tested whether the event-related potential of auditory effects of actions observed from a first- versus third-person perspective show amplitude reductions compared with externally generated sounds, as revealed for self-generated sounds. Multilevel modeling on trial-level data showed distinct dynamic patterns for the two viewpoints on reductions of the N1, P2, and N2 components. For both viewpoints, an N1 reduction for sounds generated by observed actions versus externally generated sounds was observed. However, only during first-person observation, we found a temporal dynamic within experimental runs (i.e., the N1 reduction only emerged with increasing trial number), indicating time-variant, viewpoint-dependent processes involved in sensorimotor prediction during action observation. For the P2, only a viewpoint-independent reduction was found for sounds elicited by observed actions, which disappeared in the second half of the experiment. The opposite pattern was found in an exploratory analysis concerning the N2, revealing a reduction that increased in the second half of the experiment, and, moreover, a temporal dynamic within experimental runs for the first-person perspective, possibly reflecting an agency-related process. Overall, these results suggested that the processing of auditory outcomes of observed actions is dynamically modulated by the viewpoint over time.

The impact of social anxiety on feedback-based go and nogo learning.

The term "Pavlovian" bias describes the phenomenon that learning to execute a response to obtain a reward or to inhibit a response to avoid punishment is much easier than learning the reverse. The present study investigated the interplay between this learning bias and individual levels of social anxiety. Since avoidance behavior is a hallmark feature of social anxiety and high levels of social anxiety have been associated with better learning from negative feedback, it is conceivable that the Pavlovian bias is altered in individuals with high social anxiety, with a strong tendency to avoid negative feedback, especially (but not only) in a nogo context. In addition, learning may be modulated by the individual propensity to learn from positive or negative feedback, which can be assessed as a trait-like feature. A sample of 84 healthy university students completed an orthogonalized go/nogo task that decoupled action type (go/nogo) and outcome valence (win/avoid) and a probabilistic selection task based upon which the individual propensity to learn from positive and negative feedback was determined. Self-reported social anxiety and learning propensity were used as predictors in linear mixed-effect model analysis of performance accuracy in the go/nogo task. Results revealed that high socially anxious subjects with a propensity to learn better from negative feedback showed particularly pronounced learning for nogo to avoid while lacking significant learning for nogo to win as well as go to avoid. This result pattern suggests that high levels of social anxiety in concert with negative learning propensity hamper the overcoming of Pavlovian bias in a win context while facilitating response inhibition in an avoidance context. The present data confirm the robust Pavlovian bias in feedback-based learning and add to a growing body of evidence for modulation of feedback learning by individual factors, such as personality traits. Specifically, results show that social anxiety is associated with altered Pavlovian bias, and might suggest that this effect could be driven by altered basal ganglia function primarily affecting the nogo pathway.

Similarities and Differences between Performers and Observers in Processing Auditory Action Consequences: Evidence from Simultaneous EEG Acquisition.

In our social environment, we easily distinguish stimuli caused by our own actions (e.g., water splashing when I fill my glass) from stimuli that have an external source (e.g., water splashing in a fountain). Accumulating evidence suggests that processing the auditory consequences of self-performed actions elicits N1 and P2 ERPs of reduced amplitude compared to physically identical but externally generated sounds, with such reductions being ascribed to neural predictive mechanisms. It is unexplored, however, whether the sensory processing of action outcomes is similarly modulated by action observation (e.g., water splashing when I observe you filling my glass). We tested 40 healthy participants by applying a methodological approach for the simultaneous EEG recording of two persons: An observer observed button presses executed by a performer in real time. For the performers, we replicated previous findings of a reduced N1 amplitude for self- versus externally generated sounds. This pattern differed significantly from the one in observers, whose N1 for sounds generated by observed button presses was not attenuated. In turn, the P2 amplitude was reduced for processing action- versus externally generated sounds for both performers and observers. These findings show that both action performance and observation affect the processing of action-generated sounds. There are, however, important differences between the two in the timing of the effects, probably related to differences in the predictability of the actions and thus also the associated stimuli. We discuss how these differences might contribute to recognizing the stimulus as caused by self versus others.

Effects of trait empathy and expectation on the processing of observed actions.

Recent evidence suggests that the processing of observed actions may reflect an action prediction error, with more pronounced mediofrontal negative event-related potentials (ERPs) for unexpected actions. This evidence comes from an application of a false-belief task, where unexpected correct responses elicited high ERP amplitudes. An alternative interpretation is that the ERP component reflects vicarious error processing, as objectively correct responses were errors from the observed person's perspective. In this study, we aimed to disentangle the two possibilities by adding the factor task difficulty, which varied expectations without affecting the definition of (vicarious) errors, and to explore the role of empathy in action observation. We found that the relationship between empathy and event-related potentials (ERPs) mirrored the relationship between empathy and behavioral expectancy measures. Only in the easy task condition did higher empathy lead to stronger expectancy of correct responses in the true-belief and of errors in the false-belief condition. A compatible pattern was found for an early ERP component (150-200 ms) after the observed response, with a larger negativity for error than correct responses in the true-belief and the reverse pattern in the false-belief condition, but only in highly empathic participants. We conclude that empathy facilitates the formation of expectations regarding the actions of others. These expectations then modulate the processing of observed actions, as indicated by the ERPs in the present study.

Selective Devaluation Affects the Processing of Preferred Rewards.

The present study investigated whether the representation of subjective preferences in the event-related potential is manipulable through selective devaluation, i.e., the consumption of a specific food item until satiety. Thirty-four participants completed a gambling task in which they chose between virtual doors to find one of three snack items, representing a high, medium, or low preference outcome as defined by individual desire-to-eat ratings. In one of two test sessions, they underwent selective devaluation of the high preference outcome. In the other, they completed the task on an empty stomach. Consistent with previous findings, averaged across sessions, amplitudes were increased for more preferred rewards in the time windows of P2, late FRN, and P300. As hypothesised, we also found a selective devaluation effect for the high preference outcome in the P300 time window, reflected in a decrease in amplitude. The present results provide evidence for modulations of reward processing not only by individual factors, such as subjective preferences, but also by the current motivational state. Importantly, the present data suggest that selective devaluation effects in the P300 may be a promising tool to further characterise altered valuation of food rewards in the context of eating disorders and obesity.

Asymmetric coupling of action and outcome valence in active and observational feedback learning.

Learning to execute a response to obtain a reward or to inhibit a response to avoid punishment is much easier than learning the reverse, which has been referred to as "Pavlovian" biases. Despite a growing body of research into similarities and differences between active and observational learning, it is as yet unclear if Pavlovian learning biases are specific for active task performance, i.e., learning from feedback provided for one's own actions, or if they persist also when learning by observing another person's actions and subsequent outcomes. The present study, therefore, investigated the influence of action and outcome valence in active and observational feedback learning. Healthy adult volunteers completed a go/nogo task that decoupled outcome valence (win/loss) and action (execution/inhibition) either actively or by observing a virtual co-player's responses and subsequent feedback. Moreover, in a more naturalistic follow-up experiment, pairs of subjects were tested with the same task, with one subject as active learner and the other as observational learner. The results revealed Pavlovian learning biases both in active and in observational learning, with learning of go responses facilitated in the context of reward obtainment, and learning of nogo responses facilitated in the context of loss avoidance. Although the neural correlates of active and observational feedback learning have been shown to differ to some extent, these findings suggest similar mechanisms to underlie both types of learning with respect to the influence of Pavlovian biases. Moreover, performance levels and result patterns were similar in those observational learners who had observed a virtual co-player and those who had completed the task together with an active learner, suggesting that inclusion of a virtual co-player in a computerized task provides an effective manipulation of agency.

How words get meaning: The neural processing of novel object names after sensorimotor training.

The hypothesis that individual experience affects the formation and processing of conceptual representations is controversially debated. Previous training studies with novel tool-like objects have found experience effects on conceptual representations as measured in tasks requiring the processing of object pictures. This study instead explored the neural processing of training-induced word meaning of novel object names. We asked whether the type of experience gained during object concept formation specifically modulates object name processing. In three training sessions with novel tool-like objects, two groups of healthy participants gained either active or observational manipulation experience as well as purely visual experience, while learning pseudowords serving as object names. In an fMRI session after training, participants were presented with the learned novel object names in a lexical decision task. Results revealed that processing novel object names in comparison to meaningless pseudowords elicits a word-like activation pattern in frontal, parietal and temporal regions known to underlie lexical-semantic processing, thus suggesting word meaning formation. Experience-specific modulations did not emerge as regional activation effects. However, a post-hoc analysis revealed that the type of experience (manipulation versus visual) as well as the way, in which the manipulation was learned (active versus observational) led to specific functional connectivity increases between semantic regions and neuronal assemblies in brain areas coding for object manipulation and related visuospatial information. These results suggest that the emergence of conceptual processing for novel object names might be grounded in functional brain networks specifically coding for the experience with their referents.

The Interaction between Semantic Representation and Episodic Memory.

The experimental evidence on the interrelation between episodic memory and semantic memory is inconclusive. Are they independent systems, different aspects of a single system, or separate but strongly interacting systems? Here, we propose a computational role for the interaction between the semantic and episodic systems that might help resolve this debate. We hypothesize that episodic memories are represented as sequences of activation patterns. These patterns are the output of a semantic representational network that compresses the high-dimensional sensory input. We show quantitatively that the accuracy of episodic memory crucially depends on the quality of the semantic representation. We compare two types of semantic representations: appropriate representations, which means that the representation is used to store input sequences that are of the same type as those that it was trained on, and inappropriate representations, which means that stored inputs differ from the training data. Retrieval accuracy is higher for appropriate representations because the encoded sequences are less divergent than those encoded with inappropriate representations. Consistent with our model prediction, we found that human subjects remember some aspects of episodes significantly more accurately if they had previously been familiarized with the objects occurring in the episode, as compared to episodes involving unfamiliar objects. We thus conclude that the interaction with the semantic system plays an important role for episodic memory.

Processing of Positive and Negative Feedback in Patients with Cerebellar Lesions.

It is well accepted that the cerebellum plays a crucial role in the prediction of the sensory consequences of movements. Recent findings of altered error processing in patients with selective cerebellar lesions led to the hypothesis that feedback processing and feedback-based learning might be affected by cerebellar damage as well. Thus, the present study investigated learning from and processing of positive and negative feedback in 12 patients with selective cerebellar lesions and healthy control subjects. Participants performed a monetary feedback learning task. The processing of positive and negative feedback was assessed by means of event-related potentials (ERPs) during the learning task and during a separate task in which the frequencies of positive and negative feedback were balanced. Patients did not show a general learning deficit compared to controls. Relative to the control group, however, patients with cerebellar lesions showed significantly higher ERP difference wave amplitudes (rewards-losses) in a time window between 250 and 450 ms after feedback presentation, possibly indicating impaired outcome prediction. The analysis of the original waveforms suggested that patients and controls primarily differed in their pattern of feedback-related negativity and P300 amplitudes. Our results add to recent findings on altered performance monitoring associated with cerebellar damage and demonstrate, for the first time, alterations of feedback processing in patients with cerebellar damage. Unaffected learning performance appears to suggest that chronic cerebellar lesions can be compensated in behaviour.

From perceptual to lexico-semantic analysis--cortical plasticity enabling new levels of processing.

Certain kinds of stimuli can be processed on multiple levels. While the neural correlates of different levels of processing (LOPs) have been investigated to some extent, most of the studies involve skills and/or knowledge already present when performing the task. In this study we specifically sought to identify neural correlates of an evolving skill that allows the transition from perceptual to a lexico-semantic stimulus analysis. Eighteen participants were trained to decode 12 letters of Morse code that were presented acoustically inside and outside of the scanner environment. Morse code was presented in trains of three letters while brain activity was assessed with fMRI. Participants either attended to the stimulus length (perceptual analysis), or evaluated its meaning distinguishing words from nonwords (lexico-semantic analysis). Perceptual and lexico-semantic analyses shared a mutual network comprising the left premotor cortex, the supplementary motor area (SMA) and the inferior parietal lobule (IPL). Perceptual analysis was associated with a strong brain activation in the SMA and the superior temporal gyrus bilaterally (STG), which remained unaltered from pre and post training. In the lexico-semantic analysis post learning, study participants showed additional activation in the left inferior frontal cortex (IFC) and in the left occipitotemporal cortex (OTC), regions known to be critically involved in lexical processing. Our data provide evidence for cortical plasticity evolving with a learning process enabling the transition from perceptual to lexico-semantic stimulus analysis. Importantly, the activation pattern remains task-related LOP and is thus the result of a decision process as to which LOP to engage in.

From feedback- to response-based performance monitoring in active and observational learning.

Humans can adapt their behavior by learning from the consequences of their own actions or by observing others. Gradual active learning of action-outcome contingencies is accompanied by a shift from feedback- to response-based performance monitoring. This shift is reflected by complementary learning-related changes of two ACC-driven ERP components, the feedback-related negativity (FRN) and the error-related negativity (ERN), which have both been suggested to signal events "worse than expected," that is, a negative prediction error. Although recent research has identified comparable components for observed behavior and outcomes (observational ERN and FRN), it is as yet unknown, whether these components are similarly modulated by prediction errors and thus also reflect behavioral adaptation. In this study, two groups of 15 participants learned action-outcome contingencies either actively or by observation. In active learners, FRN amplitude for negative feedback decreased and ERN amplitude in response to erroneous actions increased with learning, whereas observational ERN and FRN in observational learners did not exhibit learning-related changes. Learning performance, assessed in test trials without feedback, was comparable between groups, as was the ERN following actively performed errors during test trials. In summary, the results show that action-outcome associations can be learned similarly well actively and by observation. The mechanisms involved appear to differ, with the FRN in active learning reflecting the integration of information about own actions and the accompanying outcomes.

Impairments in learning by monetary rewards and alcohol-associated rewards in detoxified alcoholic patients.

BACKGROUND: Excessive alcohol consumption has been linked to structural and functional brain changes associated with cognitive, emotional, and behavioral impairments. It has been suggested that neural processing in the reward system is also affected by alcoholism. The present study aimed at further investigating reward-based associative learning and reversal learning in detoxified alcohol-dependent patients. METHODS: Twenty-one detoxified alcohol-dependent patients and 26 healthy control subjects participated in a probabilistic learning task using monetary and alcohol-associated rewards as feedback stimuli indicating correct responses. Performance during acquisition and reversal learning in the different feedback conditions was analyzed. RESULTS: Alcohol-dependent patients and healthy control subjects showed an increase in learning performance over learning blocks during acquisition, with learning performance being significantly lower in alcohol-dependent patients. After changing the contingencies, alcohol-dependent patients exhibited impaired reversal learning and showed, in contrast to healthy controls, different learning curves for different types of rewards with no increase in performance for high monetary and alcohol-associated feedback. CONCLUSIONS: The present findings provide evidence that dysfunctional processing in the reward system in alcohol-dependent patients leads to alterations in reward-based learning resulting in a generally reduced performance. In addition, the results suggest that alcohol-dependent patients are, in particular, more impaired in changing an established behavior originally reinforced by high rewards.

Active and observational reward learning in adults with autism spectrum disorder: relationship with empathy in an atypical sample.

INTRODUCTION: Autism spectrum disorders (ASDs) are characterised by disturbances in social behaviour. A prevailing hypothesis suggests that these problems are related to deficits in assigning rewarding value to social stimuli. The present study aimed to examine monetary reward processing in adults with ASDs by means of event-related potentials (ERPs). METHODS: Ten individuals with mild ASDs (Asperger's syndrome and high-functioning autism) and 12 healthy control subjects performed an active and an observational probabilistic reward-learning task. RESULTS: Both groups showed similar overall learning performance. With respect to reward processing, subjects with ASDs exhibited a general reduction in feedback-related negativity (FRN) amplitude, irrespective of feedback valence and type of learning (active or observational). Individuals with ASDs showed lower scores for cognitive empathy, while affective empathy did not differ between groups. Correlation analyses revealed that higher empathy (both cognitive and affective) negatively affected performance in observational learning in controls and in active learning in ASDs (only cognitive empathy). No relationships were seen between empathy and ERPs. CONCLUSIONS: Reduced FRN amplitudes are discussed in terms of a deficit in fast reward processing in ASDs, which may indicate altered reward system functioning.

Prediction-error-dependent processing of immediate and delayed positive feedback.

Learning often involves trial-and-error, i.e. repeating behaviours that lead to desired outcomes, and adjusting behaviour when outcomes do not meet our expectations and thus lead to prediction errors (PEs). PEs have been shown to be reflected in the reward positivity (RewP), an event-related potential (ERP) component between 200 and 350 ms after performance feedback which is linked to striatal processing and assessed via electroencephalography (EEG). Here we show that this is also true for delayed feedback processing, for which a critical role of the hippocampus has been suggested. We found a general reduction of the RewP for delayed feedback, but the PE was similarly reflected in the RewP and the later P300 for immediate and delayed positive feedback, while no effect was found for negative feedback. Our results suggest that, despite processing differences between immediate and delayed feedback, positive PEs drive feedback processing and learning irrespective of delay.

That means something to me: How linguistic and emotional experience affect the acquisition, representation, and processing of novel abstract concepts.

We used a novel linguistic training paradigm to investigate the experience-dependent acquisition, representation, and processing of novel emotional and neutral abstract concepts. Participants engaged in mental imagery (n = 32) or lexico-semantic rephrasing (n = 34) of linguistic material during five training sessions and successfully learned the novel abstract concepts. Feature production after training showed that specifically emotion features enriched the emotional concepts' representations. Unexpectedly, for participants engaging in vivid mental imagery during training a higher semantic richness of the acquired emotional concepts slowed down lexical decisions. Rephrasing, in turn, promoted a better learning and processing performance than imagery, probably due to stronger established lexical associations. Our results confirm the importance of emotional and linguistic experience and additional deep lexico-semantic processing for the acquisition, representation, and processing of abstract concepts. (PsycInfo Database Record (c) 2024 APA, all rights reserved).