Electrophysiological correlates of sustained conscious perception.

Previous research on the neural correlates of consciousness (NCC) in visual perception revealed an early event-related potential (ERP), the visual awareness negativity (VAN), to be associated with stimulus awareness. However, due to the use of brief stimulus presentations in previous studies, it remains unclear whether awareness-related negativities represent a transient onset-related response or correspond to the duration of a conscious percept. Studies are required that allow prolonged stimulus presentation under aware and unaware conditions. The present ERP study aimed to tackle this challenge by using a novel stimulation design. Male and female human participants (n = 62) performed a visual task while task-irrelevant line stimuli were presented in the background for either 500 or 1000 ms. The line stimuli sometimes contained a face, which needed so-called visual one-shot learning to be seen. Half of the participants were informed about the presence of the face, resulting in faces being perceived by the informed but not by the uninformed participants. Comparing ERPs between the informed and uninformed group revealed an enhanced negativity over occipitotemporal electrodes that persisted for the entire duration of stimulus presentation. Our results suggest that sustained visual awareness negativities (SVAN) are associated with the duration of stimulus presentation.

How and when social evaluative feedback is processed in the brain: A systematic review on ERP studies.

Social evaluative feedback informs the receiver of the other's views, which may contain judgments of personality-related traits and/or the level of likability. Such kinds of social evaluative feedback are of particular importance to humans. Event-related potentials (ERPs) can directly measure where in the processing stream feedback valence, expectancy, or contextual relevance modulate information processing. This review provides an overview and systematization of studies and early, mid-latency, and late ERP effects. Early effects were inconsistently reported for all factors. Feedback valence effects are more consistently reported for specific mid-latency ERPs (Reward Positivity, RewP, and Early Posterior Negativity, EPN) and late positivities (P3 and Late Positive Potential, LPP). Unexpected feedback consistently increased the Feedback Related Negativity (FRN) and, less consistently, decreased P3 amplitudes. Contextual relevance of the sender (e.g., human vs computer sender) or self-relatedness increased mid-latency to late ERPs. Interactions between valence and other factors were less often found, arising during mid-latency stages, where most consistent interactions showed larger EPN and P3 amplitude differences for valent feedback in a more relevant context. The ERP findings highlight that social evaluative feedback is consistently differentiated during mid-latency processing stages. The review discusses the relevance of findings, possible shortcomings of different experimental designs, and open questions. Furthermore, we suggest concrete venues for future research.

Effects of emotion on auditory ERPs are independent of manipulated target relevance.

Emotional attention describes the prioritized processing of emotional information to help humans quickly detect biologically salient stimuli and initiate appropriate reactions. Humans can also voluntarily attend to specific stimulus features that are target-relevant. Electrophysiological studies have shown specific temporal interactions between voluntary and emotional attention, while no such studies exist for natural sounds (e.g., explosions, running water, applause). In two experiments (N = 40, each), we examined event-related potentials (ERPs) toward target relevant or irrelevant negative, neutral, or positive sounds. Target relevance was induced by the instruction to respond blockwise to either negative, neutral, or positive sounds. Emotional sounds elicited increased fronto-central N1 and P2 amplitudes and a larger late positive potential (LPP), with more sustained effects for negative sounds. Target relevance increased amplitudes during an early LPP interval (400-900 ms) but did not interact with the valence of the sounds. These results show early and late ERP modulations for natural sounds, which do not interact with the target relevance of the sound valence, in contrast to findings from the visual domain. Thus, findings indicate little temporal overlap between emotional processes and target relevance effects in the auditory domain. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Stimulus awareness is associated with secondary somatosensory cortex activation in an inattentional numbness paradigm.

While inattentional blindness and deafness studies have revealed neural correlates of consciousness (NCC) without the confound of task relevance in the visual and auditory modality, comparable studies for the somatosensory modality are lacking. Here, we investigated NCC using functional magnetic resonance imaging (fMRI) in an inattentional numbness paradigm. Participants (N = 44) received weak electrical stimulation on the left hand while solving a demanding visual task. Half of the participants were informed that task-irrelevant weak tactile stimuli above the detection threshold would be applied during the experiment, while the other half expected stimuli below the detection threshold. Unexpected awareness assessments after the experiment revealed that altogether 10 participants did not consciously perceive the somatosensory stimuli during the visual task. Awareness was not significantly modulated by prior information. The fMRI data show that awareness of stimuli led to increased activation in the contralateral secondary somatosensory cortex. We found no significant effects of stimulus awareness in the primary somatosensory cortex or frontoparietal areas. Thus, our results support the hypothesis that somatosensory stimulus awareness is mainly based on activation in higher areas of the somatosensory cortex and does not require strong activation in extended anterior or posterior networks, which is usually seen when perceived stimuli are task-relevant.

Beyond facial expressions: A systematic review on effects of emotional relevance of faces on the N170.

The N170 is the most prominent electrophysiological signature of face processing. While facial expressions reliably modulate the N170, there is considerable variance in N170 modulations by other sources of emotional relevance. Therefore, we systematically review and discuss this research area using different methods to manipulate the emotional relevance of inherently neutral faces. These methods were categorized into (1) existing pre-experimental affective person knowledge (e.g., negative attitudes towards outgroup faces), (2) experimentally instructed affective person knowledge (e.g., negative person information), (3) contingency-based affective learning (e.g., fear-conditioning), or (4) the immediate affective context (e.g., emotional information directly preceding the face presentation). For all categories except the immediate affective context category, the majority of studies reported significantly increased N170 amplitudes depending on the emotional relevance of faces. Furthermore, the potentiated N170 was observed across different attention conditions, supporting the role of the emotional relevance of faces on the early prioritized processing of configural facial information, regardless of low-level differences. However, we identified several open research questions and suggest venues for further research.

Altered Putamen Activation for Social Comparison-Related Feedback in Social Anxiety Disorder: A Pilot Study.

INTRODUCTION: Social anxiety disorder (SAD) is characterized by abnormal processing of performance-related social stimuli. Previous studies have shown altered emotional experiences and activations of different sub-regions of the striatum during processing of social stimuli in patients with SAD. However, whether and to what extent social comparisons affect behavioural and neural responses to feedback stimuli in patients with SAD is unknown. MATERIALS AND METHODS: To address this issue, emotional ratings and functional magnetic resonance imaging (fMRI) responses were assessed while patients suffering from SAD and healthy controls (HC) were required to perform a choice task and received performance feedback (correct, incorrect, non-informative) that varied in relation to the performance of fictitious other participants (a few, half, or most of others had the same outcome). RESULTS: Across all performance feedback conditions, fMRI analyses revealed reduced activations in bilateral putamen when feedback was assumed to be received by only a few compared to half of the other participants in patients with SAD. Nevertheless, analysis of rating data showed a similar modulation of valence and arousal ratings in patients with SAD and HC depending on social comparison-related feedback. CONCLUSIONS: This suggests altered neural processing of performance feedback depending on social comparisons in patients with SAD.

The effects of visual perceptual load on detection performance and event-related potentials to auditory stimuli.

Load Theory states that perceptual load prevents, or at least reduces, the processing of task-unrelated stimuli. This study systematically examined the detection and neural processing of auditory stimuli unrelated to a visual foreground task. The visual task was designed to create continuous perceptual load, alternated between low and high load, and contained performance feedback to motivate participants to focus on the visual task instead of the auditory stimuli presented in the background. The auditory stimuli varied in intensity, and participants signaled their subjective perception of these stimuli without receiving feedback. Depending on stimulus intensity, we observed load effects on detection performance and P3 amplitudes of the event-related potential (ERP). N1 amplitudes were unaffected by perceptual load, as tested by Bayesian statistics. Findings suggest that visual perceptual load affects the processing of auditory stimuli in a late time window, which is associated with a lower probability of reported awareness of these stimuli.

Visual perceptual load and processing of somatosensory stimuli in primary and secondary somatosensory cortices.

Load theory assumes that neural activation to distractors in early sensory cortices is modulated by the perceptual load of a main task, regardless of whether task and distractor share the same sensory modality or not. While several studies have investigated the question of load effects on distractor processing in early sensory areas, there is no functional magnetic resonance imaging (fMRI) study regarding load effects on somatosensory stimuli. Here, we used fMRI to investigate effects of visual perceptual load on neural responses to somatosensory stimuli applied to the wrist in a study with 44 participants. Perceptual load was manipulated by an established sustained visual detection task, which avoided simultaneous target and distractor presentations. Load was operationalized by detection difficulty of subtle or clear color changes of one of 12 rotating dots. While all somatosensory stimuli led to activation in somatosensory areas SI and SII, we found no statistically significant difference in brain activation to these stimuli under high compared to low sustained visual load. Moreover, exploratory Bayesian analyses supported the absence of differences. Thus, our findings suggest a resistance of somatosensory processing to at least some forms of visual perceptual load, possibly due to behavioural relevance of discrete somatosensory stimuli and separable attentional resources for the somatosensory and visual modality.

The effects of visual working memory load on detection and neural processing of task-unrelated auditory stimuli.

While perceptual load has been proposed to reduce the processing of task-unrelated stimuli, theoretical arguments and empirical findings for other forms of task load are inconclusive. Here, we systematically investigated the detection and neural processing of auditory stimuli varying in stimulus intensity during a stimuli-unrelated visual working memory task alternating between low and high load. We found, depending on stimulus strength, decreased stimulus detection and reduced P3, but unaffected N1 amplitudes of the event-related potential to auditory stimuli under high as compared to low load. In contrast, load independent awareness effects were observed during both early (N1) and late (P3) time windows. Findings suggest a late neural effect of visual working memory load on auditory stimuli leading to lower probability of reported awareness of these stimuli.

Potentiated early neural responses to fearful faces are not driven by specific face parts.

Prioritized processing of fearful compared to neutral faces is reflected in increased amplitudes of components of the event-related potential (ERP). It is unknown whether specific face parts drive these modulations. Here, we investigated the contributions of face parts on ERPs to task-irrelevant fearful and neutral faces using an ERP-dependent facial decoding technique and a large sample of participants (N = 83). Classical ERP analyses showed typical and robust increases of N170 and EPN amplitudes by fearful relative to neutral faces. Facial decoding further showed that the absolute amplitude of these components, as well as the P1, was driven by the low-frequency contrast of specific face parts. However, the difference between fearful and neutral faces was not driven by any specific face part, as supported by Bayesian statistics. Furthermore, there were no correlations between trait anxiety and main effects or interactions. These results suggest that increased N170 and EPN amplitudes to task-irrelevant fearful compared to neutral faces are not driven by specific facial regions but represent a holistic face processing effect.

Testing stimulus exposure time as the critical factor of increased EPN and LPP amplitudes for fearful faces during perceptual distraction tasks.

Fearful facial expressions are prioritized across different information processing stages, as evident in early, intermediate, and late components of event-related brain potentials (ERPs). Recent studies showed that, in contrast to early N170 modulations, mid-latency (Early Posterior Negativity, EPN) and late (Late Positive Potential, LPP) emotional modulations depend on the attended perceptual feature. Nevertheless, several studies reported significant differences between emotional and neutral faces for the EPN or LPP components during distraction tasks. One cause for these conflicting findings might be that when faces are presented sufficiently long, participants attend to task-irrelevant features of the faces. In this registered report, we tested whether the presentation duration of faces is the critical factor for differences between reported emotional modulations during perceptual distraction tasks. To this end, 48 participants were required to discriminate the orientation of lines overlaid onto fearful or neutral faces, while face presentation varied (100 msec, 300 msec, 1,000 msec, 2,000 msec). While participants did not need to pay attention to the faces, we observed main effects of emotion for the EPN and LPP, but no interaction between emotion and presentation duration. Of note, unregistered exploratory tests per presentation duration showed no significant EPN and LPP emotion differences during short durations (100 and 300 msec) but significant differences with longer durations. While the presentation duration seems not to be a critical factor for EPN and LPP emotion effects, future studies are needed to investigate the role of threshold effects and the applied analytic designs to explain conflicting findings in the literature.

Fearful faces straight ahead or in the periphery: Early neuronal responses independently of trait anxiety.

For humans, it is vitally important to rapidly detect potentially threatening stimuli such as fearful faces in the periphery. Trait anxiety has been related to biased responses to threatening faces, which might be more pronounced for peripheral stimuli. We examined the impact of spatial location and trait anxiety on event-related potentials to fearful faces (ERPs) in a large sample (N = 80). Using a face-unrelated oddball detection task and online eye-tracking, we ensured central fixation but sustained attention to the location of fearful and neutral faces at central or peripheral locations (12° left or right). Manipulation checks showed high task performance and successful shifts of visuospatial attention indexed by prestimulus alpha lateralization of the EEG spectra. Concerning ERPs, we observed increased P1 amplitudes for fearful compared to neutral faces, independent of the spatial location. In contrast, the N170 to fearful faces was only potentiated for centrally presented faces. Finally, fearful and neutral faces did not differ during the EPN window at any spatial location. Trait anxiety was unrelated to fearful-neutral ERP differences for each examined location. Our findings show that differential P1 responses are less constrained by the stimulus location, possibly due to coarse low-level stimulus information processing. However, the relative N170 increase for fearful faces depends on the central stimulus location. Furthermore, findings support the view that differential processing at the stage of the EPN depends on attentional conditions. Finally, our results question the hypothesis that trait anxiety differences correlate with mandatorily altered processing of threatening stimuli. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Effects of emotional valence and intensity on cognitive and affective empathy after insula lesions.

The insula plays a central role in empathy. However, the complex structure of cognitive (CE) and affective empathy (AE) deficits following insular damage is not fully understood. In the present study, patients with insular lesions (n = 20) and demographically matched healthy controls (n = 24) viewed ecologically valid videos that varied in terms of valence and emotional intensity. The videos showed a person (target) narrating a personal life event. In CE conditions, subjects continuously rated the affective state of the target, while in AE conditions, they continuously rated their own affect. Mean squared error (MSE) assessed deviations between subject and target ratings. Patients differed from controls only in negative, low-intensity AE, rating their own affective state less negative than the target. This deficit was not related to trait empathy, neuropsychological or clinical parameters, or laterality of lesion. Empathic functions may be widely spared after insular damage in a naturalistic, dynamic setting, potentially due to the intact interpretation of social context by residual networks outside the lesion. The particular role of the insula in AE for negative states may evolve specifically in situations that bear higher uncertainty pointing to a threshold role of the insula in online ratings of AE.

Early neural potentiation to centrally and peripherally presented fear-conditioned faces.

For humans, it is vitally important to rapidly detect and process threatening signals regardless of whether stimuli occur at fixation or in the periphery. However, it is unknown whether eccentricity affects early neuronal electrophysiological responses to fear-conditioned stimuli. We examined early event-related potentials (ERPs) of the electroencephalogram (EEG) to fear-conditioned faces to address this question. Participants (N = 80) were presented with faces, either paired with an aversive (CS+) or neutral sound (CS-), at central or peripheral positions. We ensured constant central fixation using online eye-tracking but directed attention to either centrally or peripherally presented faces. Manipulation checks showed successful fear-conditioning (i.e., on average lower ratings in valence and higher ratings in arousal and perceived threat) and successful shifts of visuospatial attention indexed by high task performance and pre-stimulus alpha lateralization of the EEG spectra. We observed a generally increased P1 to fear-conditioned faces regardless of presentation location. An N170 difference between fear-conditioned and neutral stimuli was found but was restricted to the central location and depended on the effectivity of fear-conditioning. A similar effect was observed for the early posterior negativity (EPN). Trait anxiety was not related to differential ERP responses to CS+ versus CS- faces for any ERP component. These findings suggest that the P1 indexes early responses to centrally and peripherally presented fear-conditioned faces. Subsequent stages are modulated by the spatial location of the stimuli. This suggests different stages of neural processing of fear-conditioned faces depending on their spatial location. Finally, our results question the hypothesis that trait anxiety in healthy participants is related to altered visual processing of fear-conditioned faces.

Neural correlates of consciousness in an attentional blink paradigm with uncertain target relevance.

Several event-related potentials (ERPs) have been proposed as neural correlates of consciousness (NCC), most prominently the early visual awareness negativity (VAN) and the late P3b component. Highly influential support for the P3b comes from studies utilizing the attentional blink (AB), where conscious perception of a first visual target (T1) impairs reporting a second target (T2) presented shortly afterwards. Recent no-report studies using other paradigms suggest that the P3b component may reflect post-perceptual processes associated with decision-making rather than awareness. However, no-report studies are limited in their awareness assessment, and their conclusions have not been tested in an AB paradigm. The present study (N = 38) addressed these issues using a novel AB paradigm, which reduced decision-making processes by omitting a discrimination task on T2 stimuli and rendering their relevance uncertain. Nevertheless, awareness was assessed trial by trial. Comparing ERPs in response to seen versus unseen T2 stimuli revealed a VAN but no enhanced P3b regardless of whether they were marked as distinct from distractor stimuli or not. Our results corroborate the VAN and challenge the P3b as NCC despite rigorous trial-by-trial assessment of conscious perception. Thus, they support the idea that awareness emerges during early sensory processing.

Early and late electrophysiological correlates of gradual perceptual awareness in- and outside the Attentional Blink window.

There is an ongoing debate on the neural correlates of consciousness (NCC) in the attentional blink (AB). Theoretical accounts propose that NCC during the attentional blink occur late in the processing hierarchy and that this quality is specific to the AB. We investigated this question by recording event-related potentials during an AB experiment with faces as T2. We analyzed ERPs to T2 stimuli inside (short lag) and outside (long lag) the AB window after carefully calibrating T2 stimuli to ensure equal visibility ratings across lags. We found that the N170, the visual awareness negativity (VAN), and the P3b showed an increased amplitude for seen compared to unseen face stimuli regardless of stimulus lag and that all these components scale linearly with subjective visibility. These findings suggest similar early and late mechanisms of graded perceptual awareness within and outside the AB across perceptual (N170, VAN) and post-perceptual (P3b) processing stages.

Effects of awareness and task relevance on neurocomputational models of mismatch negativity generation.

Detection of regularities and their violations in sensory input is key to perception. Violations are indexed by an early EEG component called the mismatch negativity (MMN) - even if participants are distracted or unaware of the stimuli. On a mechanistic level, two dominant models have been suggested to contribute to the MMN: adaptation and prediction. Whether and how context conditions, such as awareness and task relevance, modulate the mechanisms of MMN generation is unknown. We conducted an EEG study disentangling influences of task relevance and awareness on the visual MMN. Then, we estimated different computational models for the generation of single-trial amplitudes in the MMN time window. Amplitudes were best explained by a prediction error model when stimuli were task-relevant but by an adaptation model when task-irrelevant and unaware. Thus, mismatch generation does not rely on one predominant mechanism but mechanisms vary with task relevance of stimuli.

Differential effects of prediction error and adaptation along the auditory cortical hierarchy during deviance processing.

Neural mismatch responses have been proposed to rely on different mechanisms, including prediction error-related activity and adaptation to frequent stimuli. However, the hierarchical cortical structure of these mechanisms is unknown. To investigate this question, we recorded hemodynamic responses while participants (N = 54) listened to an auditory oddball sequence as well as a suited control condition. In addition to effects in sensory processing areas (Heschl's gyrus, superior temporal gyrus (STG)), we found several distinct clusters that indexed deviance processing in frontal and parietal regions (anterior cingulate cortex/supplementary motor area (ACC/SMA), inferior parietal lobule (IPL), anterior insula (AI), inferior frontal junction (IFJ)). Comparing responses to the control stimulus with the deviant and standard enabled us to delineate the contributions of prediction error- or adaptation-related brain activation, respectively. We observed significant effects of adaptation in Heschl's gyrus, STG and ACC/SMA, while prediction error-related activity was observed in STG, IPL, AI and IFJ. Additional dynamic causal modeling confirmed the superiority of a hierarchical processing structure compared to a flat structure. Thus, we found that while prediction-error related processes increased with the hierarchical level of the brain area, adaptation declined. This suggests that the relative contribution of different mechanisms in deviance processing varies across the cortical hierarchy.

Effects of task load, spatial attention, and trait anxiety on neuronal responses to fearful and neutral faces.

There is an ongoing debate on how different components of the event-related potential (ERP) to threat-related facial expressions are modulated by attentional conditions and interindividual differences in trait anxiety. In the current study (N = 80), we examined ERPs to centrally presented, task-irrelevant fearful and neutral faces, while participants had to solve a face-unrelated visual task, which differed in difficulty and spatial position. Critically, we used a fixation-controlled experimental design and ensured the spatial attention manipulation by spectral analysis of the EEG. Besides the factors emotion, spatial attention, and perceptual load, we also investigated correlations between trait anxiety and ERPs. While P1 emotion effects were insignificant, the N170 was increased to fearful faces regardless of load and spatial attention conditions. During the EPN time window, a significantly increased negativity for fearful faces was observed only during low load and spatial attention to the face. We found no significant relationship between ERPs and trait anxiety, questioning the hypothesis of a general hypersensitivity toward fearful expressions in anxious individuals. These results show a high resistance of the N170 amplitude increase for fearful faces to spatial attention and task load manipulations. By contrast, the EPN modulation by fearful faces index a resource-dependent stage of the ERP, requiring both spatial attention at the location of faces and low load of the face-irrelevant task.

No trait anxiety influences on early and late differential neuronal responses to aversively conditioned faces across three different tasks.

The human brain's ability to quickly detect dangerous stimuli is crucial in selecting appropriate responses to possible threats. Trait anxiety has been suggested to moderate these processes on certain processing stages. To dissociate such different information-processing stages, research using classical conditioning has begun to examine event-related potentials (ERPs) in response to fear-conditioned (CS +) faces. However, the impact of trait anxiety on ERPs to fear-conditioned faces depending on specific task conditions is unknown. In this preregistered study, we measured ERPs to faces paired with aversive loud screams (CS +) or neutral sounds (CS -) in a large sample (N = 80) under three different task conditions. Participants had to discriminate face-irrelevant perceptual information, the gender of the faces, or the CS category. Results showed larger amplitudes in response to aversively conditioned faces for all examined ERPs, whereas interactions with the attended feature occurred for the P1 and the early posterior negativity (EPN). For the P1, larger CS + effects were observed during the perceptual distraction task, while the EPN was increased for CS + faces when deciding about the CS association. Remarkably, we found no significant correlations between ERPs and trait anxiety. Thus, fear-conditioning potentiates all ERP amplitudes, some processing stages being further modulated by the task. However, the finding that these ERP differences were not affected by individual differences in trait anxiety does not support theoretical accounts assuming increased threat processing or reduced threat discrimination depending on trait anxiety.