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.

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.

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.

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).

Auditory N1 and P2 attenuation in action observation: An event-related potential study considering effects of temporal predictability and individualism.

Tones that are generated by self-performed actions elicit attenuated N1 and P2 amplitudes, as measured by electroencephalography (EEG), compared to identical external tones, which is referred to as neurophysiological sensory attenuation (SA). At the same time, self-generated tones are perceived as less loud compared to external tones (perceptual SA). Action observation led in part to a similar neurophysiological and perceptual SA. The perceptual SA in observers was found in comparison to tones that were temporally predictable, and one study suggested that perceptual SA in observers might depend on the cultural dimension of individualism. In this study, we examined neurophysiological SA for tones elicited by self-performed and observed actions during simultaneous EEG acquisitions in two participants, extending the paradigm with a visual cue condition controlling for effects of temporal predictability. Moreover, we investigated the effect of individualism on neurophysiological SA in action observation. Relative to un-cued external tones, the N1 was only descriptively reduced for tones that were elicited by self-performed or observed actions and significantly attenuated for cued external tones. A P2 attenuation effect relative to un-cued external tones was found in all three conditions, with stronger effects for self- and other-generated tones than for cued external tones. We found no evidence for an effect of individualism. These findings add to previous evidence for neurophysiological SA in action performance and observation with a paradigm well-controlled for the effect of predictability and individualism, showing differential effects of the former on the N1 and P2 components, and no effect of the latter.

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.

Learning new words via feedback-Association between feedback-locked ERPs and recall performance-An exploratory study.

Feedback learning is thought to involve the dopamine system and its projection sites in the basal ganglia and anterior cingulate cortex (ACC), regions associated with procedural learning. Under certain conditions, such as when feedback is delayed, feedback-locked activation is pronounced in the medial temporal lobe (MTL), which is associated with declarative learning. In event-related potential research, the feedback-related negativity (FRN) has been linked to immediate feedback processing, while the N170, possibly reflecting MTL activity, has been related to delayed feedback processing. In the current study, we performed an exploratory investigation on the relation between N170 and FRN amplitude and memory performance in a test for declarative memory (free recall), also exploring the role of feedback delay. To this end, we adapted a paradigm in which participants learned associations between non-objects and non-words with either immediate or delayed feedback, and added a subsequent free recall test. We indeed found that N170, but not FRN amplitudes, depended on later free recall performance, with smaller amplitudes for later remembered non-words. In an additional analysis with memory performance as dependent variable, the N170, but not the FRN amplitude predicted free recall, modulated by feedback timing and valence. This finding shows that the N170 reflects an important process during feedback processing, possibly related to expectations and their violation, but is distinct from the process reflected by the FRN.

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.

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.

Altered performance monitoring in Tourette Syndrome: an MEG investigation.

The error-related negativity (ERN) is an event-related potential component indexing processes of performance monitoring during simple stimulus-response tasks: the ERN is typically enhanced for error processing and conflicting response representations. Investigations in healthy participants and different patient groups have linked the ERN to the dopamine system and to prefrontal information processing. As in patients with Tourette Syndrome (TS) both dopamine release and prefrontal information processing are impaired, we hypothesized that performance monitoring would be altered, which was investigated with magnetencephalography (MEG). We examined performance monitoring in TS patients by assessing the magnetic equivalent of the ERN (mERN). The mERN was investigated in tic-free trials of eight adult, unmedicated TS patients without clinically significant comorbidity and ten matched healthy controls while performing a Go/NoGo task in selected frontocentral channels. The analysis of the response-related amplitudes of the event-related magnetic field showed that TS patients, in contrast to controls, did not show earlier amplitude modulation (between 70 and 105 ms after response onset) depending on response type (errors or correct responses). In both groups significant mERN amplitudes in the time-window between 105 and 160 ms after response onset were detected thus pointing at only later error processing in TS patients. In TS patients, early error-related processing might be affected by an enhanced motor control triggered by a conflict between the targeted high task performance and tic suppression. TS patients seem to tend to initially process all responses as erroneous responses.

Effective connectivity within the neural system for object-directed action representation during aware and unaware tool processing.

Previous brain functional specialization evidence has shown that both aware and unaware visual processing of manipulable objects activate left premotor, parietal, and posterior temporal cortices, which are thought to constitute object-directed action and object-function processing streams. An open question is whether, both under supraliminal and subliminal processing conditions, there is directional spread of activation along these functional streams, leading to causal inter-regional connectivity effects. In this study, we used Dynamic Causal Modelling to estimate the effective connectivity influences within the premotor-parieto-temporal network, as a function of factorial contrasts for Manipulability (manipulable vs non-manipulable objects) and Perceptual Awareness (above vs below perceptual threshold). We modeled forward and backward connections originating from visual area V4, as a region underlying object texture segregation, and spreading through the left premotor-parieto-temporal network. Both above and below perceptual threshold, the visual processing of manipulable objects was associated with a specific increase of reciprocal effective connectivity coupling among left premotor-parieto-temporal regions. Aware and unaware manipulable object processing differed from each other for their distinct patterns of top-down activation enhancement exerted, in the former case, by left premotor-parieto-temporal regions on area V4 and, in the latter case, by left premotor on temporal regions. Although it is only under aware processing conditions that effective connectivity in the action representation system may promote object visual contour segregation in area V4, our results suggest that the encoding of object-action and object-function information can occur through left-hemispheric premotor, parietal, and temporal causal interdependencies, even when the object is not consciously perceived.

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.

Disentangling effects of expectancy, accuracy, and empathy on the processing of observed actions.

A number of studies suggest that event-related potential (ERP) components previously associated with error processing might represent expectation violation instead of valence. When observing others, these processes might further be modulated by trait empathy. We suggest that trait empathy modulates expectancy formation and that these expectancies then influence observed response processing as reflected in a frontocentral negative ERP component resembling the previously described observer error-related negativity. We acquired single trial ERPs of participants who observed another person in a true- or false-belief condition answering correctly or erroneously. Additionally, we prompted participants' expectancy in some trials. Using linear mixed model analyses, we found that for low empathy participants, expectations for the false-belief condition decreased throughout the experiment, so that expectations were more pronounced in participants with higher empathy toward the end of the experiment. We also found that single trial expectancy measures derived from regression models of the measured expectancies predicted the amplitude of the frontocentral negative ERP component, and that neither the addition of empathy nor accuracy or trial type (true- or false-belief) led to the explanation of significantly more variance compared with the model just containing expectancy as predictor. These results suggest that empathy modulates the processing of observed responses indirectly via its effect on expectancy of the response.

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.

Corroborating behavioral evidence for the interplay of representational richness and semantic control in semantic word processing.

This study aimed to replicate and validate concreteness and context effects on semantic word processing. In Experiment 1, we replicated the behavioral findings of Hoffman et al. (Cortex 63,250-266, https://doi.org/10.1016/j.cortex.2014.09.001 , 2015) by applying their cueing paradigm with their original stimuli translated into German. We found concreteness and contextual cues to facilitate word processing in a semantic judgment task with 55 healthy adults. The two factors interacted in their effect on reaction times: abstract word processing profited more strongly from a contextual cue, while the concrete words' processing advantage was reduced but still present. For accuracy, the descriptive pattern of results suggested an interaction, which was, however, not significant. In Experiment 2, we reformulated the contextual cues to avoid repetition of the to-be-processed word. In 83 healthy adults, the same pattern of results emerged, further validating the findings. Our corroborating evidence supports theories integrating representational richness and semantic control mechanisms as complementary mechanisms in semantic word processing.

Illusion of control affects ERP amplitude reductions for auditory outcomes of self-generated actions.

The reduction of neural responses to self-generated stimuli compared to external stimuli is thought to result from the matching of motor-based sensory predictions and sensory reafferences and to serve the identification of changes in the environment as caused by oneself. The amplitude of the auditory event-related potential (ERP) component N1 seems to closely reflect this matching process, while the later positive component (P2/ P3a) has been associated with judgments of agency, which are also sensitive to contextual top-down information. In this study, we examined the effect of perceived control over sound production on the processing of self-generated and external stimuli, as reflected in these components. We used a new version of a classic two-button choice task to induce different degrees of the illusion of control (IoC) and recorded ERPs for the processing of self-generated and external sounds in a subsequent task. N1 amplitudes were reduced for self-generated compared to external sounds, but not significantly affected by IoC. P2/3a amplitudes were affected by IoC: We found reduced P2/3a amplitudes after a high compared to a low IoC induction training, but only for self-generated, not for external sounds. These findings suggest that prior contextual belief information induced by an IoC affects later processing as reflected in the P2/P3a, possibly for the formation of agency judgments, while early processing reflecting motor-based predictions is not affected.

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.

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.

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.

Transition From Sublexical to Lexico-Semantic Stimulus Processing.

Resembling letter-by-letter translation, Morse code can be used to investigate various linguistic components by slowing down the cognitive process of language decoding. Using fMRI and Morse code, we investigated patterns of brain activation associated with decoding three-letter words or non-words and making a lexical decision. Our data suggest that early sublexical processing is associated with activation in brain regions that are involved in sound-patterns to phoneme conversion (inferior parietal lobule), phonological output buffer (inferior frontal cortex: pars opercularis) as well as phonological and semantic top-down predictions (inferior frontal cortex: pars triangularis). In addition, later lexico-semantic processing of meaningful stimuli is associated with activation of the phonological lexicon (angular gyrus) and the semantic system (default mode network). Overall, our data indicate that sublexical and lexico-semantic analyses comprise two cognitive processes that rely on neighboring networks in the left frontal cortex and parietal lobule.