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.

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.

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.

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.

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.

Decoding the neural representation of fine-grained conceptual categories.

Neuroscientific research on conceptual knowledge based on the grounded cognition framework has shed light on the organization of concrete concepts into semantic categories that rely on different types of experiential information. Abstract concepts have traditionally been investigated as an undifferentiated whole, and have only recently been addressed in a grounded cognition perspective. The present fMRI study investigated the involvement of brain systems coding for experiential information in the conceptual processing of fine-grained semantic categories along the abstract-concrete continuum. These categories consisted of mental state-, emotion-, mathematics-, mouth action-, hand action-, and leg action-related meanings. Thirty-five sentences for each category were used as stimuli in a 1-back task performed by 36 healthy participants. A univariate analysis failed to reveal category-specific activations. Multivariate pattern analyses, in turn, revealed that fMRI data contained sufficient information to disentangle all six fine-grained semantic categories across participants. However, the category-specific activity patterns showed no overlap with the regions coding for experiential information. These findings demonstrate the possibility of detecting specific patterns of neural representation associated with the processing of fine-grained conceptual categories, crucially including abstract ones, though bearing no anatomical correspondence with regions coding for experiential information as predicted by the grounded cognition hypothesis.

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.

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 prenatal brain readiness for speech processing: A review on foetal development of auditory and primordial language networks.

Despite consolidated evidence for the prenatal ability to elaborate and respond to sounds and speech stimuli, the ontogenetic functional brain maturation of language responsiveness in the foetus is still poorly understood. Recent advances in in-vivo foetal neuroimaging have contributed to a finely detailed picture of the anatomo-functional hallmarks that define the prenatal neurodevelopment of auditory and language-related networks. Here, we first outline available evidence for the prenatal development of auditory and language-related brain structures and of their anatomical connections. Second, we focus on functional connectivity data showing the emergence of auditory and primordial language networks in the foetal brain. Third, we recapitulate functional neuroimaging studies assessing the prenatal readiness for sound processing, as a crucial prerequisite for the foetus to experientially respond to spoken language. In conclusion, we suggest that the state of the art has reached sufficient maturity to directly assess the neural mechanisms underlying the prenatal readiness for speech processing and to evaluate whether foetal neuromarkers can predict the postnatal development of language acquisition abilities and disabilities.

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.

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.

The role of trait empathy in the processing of observed actions in a false-belief task.

Empathic brain responses are characterized by overlapping activations between active experience and observation of an emotion in another person, with the pattern for observation being modulated by trait empathy. Also for self-performed and observed errors, similar brain activity has been described, but findings concerning the role of empathy are mixed. We hypothesized that trait empathy modulates the processing of observed responses if expectations concerning the response are based on the beliefs of the observed person. In the present study, we utilized a false-belief task in which observed person's and observer's task-related knowledge were dissociated and errors and correct responses could be expected or unexpected. While theta power was generally modulated by the expectancy of the observed response, a negative mediofrontal event-related potential (ERP) component was more pronounced for unexpected observed actions only in participants with higher trait empathy (assessed by the Empathy Quotient), as revealed by linear mixed effects analyses. Cognitive and affective empathy, assessed by the Interpersonal Reactivity Index, were not significantly related to the ERP component. The results suggest that trait empathy can facilitate the generation of predictions and thereby modulate specific aspects of the processing of observed actions, while the contributions of specific empathy components remain unclear.

Stress effects on learning and feedback-related neural activity depend on feedback delay.

Depending on feedback timing, the neural structures involved in learning differ, with the dopamine system including the dorsal striatum and anterior cingulate cortex (ACC) being more important for learning from immediate than delayed feedback. As stress has been shown to promote striatum-dependent learning, the current study aimed to explore if stress differentially affects learning from and processing of immediate and delayed feedback. One group of male participants was stressed using the socially evaluated cold pressor test, and another group underwent a control condition. Subsequently, participants performed a reward learning task with immediate (500 ms) and delayed (6,500 ms) feedback while brain activity was assessed with electroencephalography (EEG). While stress enhanced the accuracy for delayed relative to immediate feedback, it reduced the feedback-related negativity (FRN) valence effect, which is the amplitude difference between negative and positive feedback. For the P300, a reduced valence effect was found in the stress group only for delayed feedback. Frontal theta power was most pronounced for immediate negative feedback and was generally reduced under stress. Moreover, stress reduced associations of FRN and theta power with trial-by-trial accuracy. Associations between stress-induced cortisol increases and EEG components were examined using linear mixed effects analyses, which showed that the described stress effects were accompanied by associations between the stress-induced cortisol increases and feedback processing. The results indicate that stress and cortisol affect different aspects of feedback processing. Instead of an increased recruitment of the dopamine system and the ACC, the results may suggest enhanced salience processing and reduced cognitive control under stress.

Separate and overlapping functional roles for efference copies in the human thalamus.

How the perception of space is generated from the multiple maps in the brain is still an unsolved mystery in neuroscience. A neural pathway ascending from the superior colliculus through the medio-dorsal (MD) nucleus of thalamus to the frontal eye field has been identified in monkeys that conveys efference copy information about the metrics of upcoming eye movements. Information sent through this pathway stabilizes vision across saccades. We investigated whether this motor plan information might also shape spatial perception even when no saccades are performed. We studied patients with medial or lateral thalamic lesions (likely involving either the MD or the ventrolateral (VL) nuclei). Patients performed a double-step task testing motor updating, a trans-saccadic localization task testing visual updating, and a localization task during fixation testing a general role of motor signals for visual space in the absence of eye movements. Single patients with medial or lateral thalamic lesions showed deficits in the double-step task, reflecting insufficient transfer of efference copy. However, only a patient with a medial lesion showed impaired performance in the trans-saccadic localization task, suggesting that different types of efference copies contribute to motor and visual updating. During fixation, the MD patient localized stationary stimuli more accurately than healthy controls, suggesting that patients compensate the deficit in visual prediction of saccades - induced by the thalamic lesion - by relying on stationary visual references. We conclude that partially separable efference copy signals contribute to motor and visual stability in company of purely visual signals that are equally effective in supporting trans-saccadic perception.

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.

Moral decision making under modafinil: a randomized placebo-controlled double-blind crossover fMRI study.

RATIONALE: Modafinil is increasingly used by healthy humans as a neuroenhancer in order to improve cognitive functioning. Research on the effects of modafinil on cognition yielded most consistent findings for complex tasks relying on the prefrontal cortex (PFC). OBJECTIVES: The present randomized placebo-controlled double-blind crossover study aimed to investigate the effect of a single dose of modafinil (200 mg) on everyday moral decision making and its neural correlates, which have been linked to the ventro- and dorsomedial PFC. METHODS: Healthy male study participants were presented with short stories describing everyday moral or neutral dilemmas. Each moral dilemma required a decision between a personal desire and a moral standard, while the neutral dilemmas required decisions between two personal desires. The participants underwent this task twice, once under the influence of modafinil and once under placebo. Brain activity associated with the processing of the dilemmas was assessed by means of functional magnetic resonance imaging. RESULTS: For the processing of moral vs. neutral dilemmas, activations were found in a network of brain regions linked to social cognitive processes including, among others, the bilateral medial PFC, the insula, and the precuneus. Modafinil was found to increase the number of moral decisions and had no effect on brain activity associated with dilemma processing. Exploratory analyses revealed reduced response-locked activity in the dorsomedial PFC for moral compared to neutral dilemmas under modafinil, but not under placebo. CONCLUSIONS: The results are discussed in terms of altered predictions of others' emotional states under modafinil, possibly due to higher processing efficiency.

The role of experience for abstract concepts: Expertise modulates the electrophysiological correlates of mathematical word processing.

Embodied theories assign experience a crucial role in shaping conceptual representations. Supporting evidence comes mostly from studies on concrete concepts, where e.g., motor expertise facilitated action concept processing. This study examined experience-dependent effects on abstract concept processing. We asked participants with high and low mathematical expertise to perform a lexical decision task on mathematical and nonmathematical abstract words, while acquiring event-related potentials. Analyses revealed an interaction of expertise and word type on the amplitude of a fronto-central N400 and a centro-parietal late positive component (LPC). For mathematical words, we found a trend for a lower N400 and a significantly higher LPC amplitude in experts compared to nonexperts. No differences between groups were found for nonmathematical words. The results suggest that expertise affects the processing stages of semantic integration and memory retrieval specifically for expertise-related concepts. This study supports the generalization of experience-dependent conceptual processing mechanisms to the abstract domain.

The Effect of Training-Induced Visual Imageability on Electrophysiological Correlates of Novel Word Processing.

The concreteness effect (CE) describes a processing advantage for concrete over abstract words. Electrophysiologically, the CE manifests in higher N400 and N700 amplitudes for concrete words. The contribution of the stimulus-inherent imageability to the electrophysiological correlates of the CE is not yet fully unraveled. This EEG study focused on the role of imageability irrespective of concreteness by examining the effects of training-induced visual imageability on the processing of novel words. In two training sessions, 21 healthy participants learned to associate novel words with pictures of novel objects as well as electron-microscopical structures and were additionally familiarized with novel words without any picture association. During a post-training EEG session, participants categorized trained novel words with or without picture association, together with real concrete and abstract words. Novel words associated with novel object pictures during the training elicited a higher N700 than familiarized novel words without picture-association. Crucially, this training-induced N700 effect resembled the CE found for real words. However, a CE on the N400 was found for real words, but no effect of imageability in novel words. The results suggest that the N400 CE for real words depends on the integration of multiple semantic features, while mere visual imageability might contribute to the CE in the N700 time window.

ERP correlates of processing the auditory consequences of own versus observed actions.

Research has so far focused on neural mechanisms that allow us to predict the sensory consequences of our own actions, thus also contributing to ascribing them to ourselves as agents. Less attention has been devoted to processing the sensory consequences of observed actions ascribed to another human agent. Focusing on audition, there is consistent evidence of a reduction of the auditory N1 ERP for self- versus externally generated sounds, while ERP correlates of processing sensory consequences of observed actions are mainly unexplored. In a between-groups ERP study, we compared sounds generated by self-performed (self group) or observed (observation group) button presses with externally generated sounds, which were presented either intermixed with action-generated sounds or in a separate condition. Results revealed an overall reduction of the N1 amplitude for processing action- versus externally generated sounds in both the intermixed and the separate condition, with no difference between the groups. Further analyses, however, suggested that an N1 attenuation effect relative to the intermixed condition at frontal electrode sites might exist only for the self but not for the observation group. For both groups, we found a reduction of the P2 amplitude for processing action- versus all externally generated sounds. We discuss whether the N1 and the P2 reduction can be interpreted in terms of predictive mechanisms for both action execution and observation, and to what extent these components might reflect also the feeling of (self) agency and the judgment of agency (i.e., ascribing agency either to the self or to others).