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.

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.

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.

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

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

Dissociating the Neural Correlates of Consciousness and Task Relevance in Face Perception Using Simultaneous EEG-fMRI.

Current theories of visual consciousness disagree about whether it emerges during early stages of processing in sensory brain regions or later when a widespread frontoparietal network becomes involved. Moreover, disentangling conscious perception from task-related postperceptual processes (e.g., report) and integrating results across different neuroscientific methods remain ongoing challenges. The present study addressed these problems using simultaneous EEG-fMRI and a specific inattentional blindness paradigm with three physically identical phases in female and male human participants. In phase 1, participants performed a distractor task during which line drawings of faces and control stimuli were presented centrally. While some participants spontaneously noticed the faces in phase 1, others remained inattentionally blind. In phase 2, all participants were made aware of the task-irrelevant faces but continued the distractor task. In phase 3, the faces became task-relevant. Bayesian analysis of brain responses demonstrated that conscious face perception was most strongly associated with activation in fusiform gyrus (fMRI) as well as the N170 and visual awareness negativity (EEG). Smaller awareness effects were revealed in the occipital and prefrontal cortex (fMRI). Task-relevant face processing, on the other hand, led to strong, extensive activation of occipitotemporal, frontoparietal, and attentional networks (fMRI). In EEG, it enhanced early negativities and elicited a pronounced P3b component. Overall, we provide evidence that conscious visual perception is linked with early processing in stimulus-specific sensory brain areas but may additionally involve prefrontal cortex. In contrast, the strong activation of widespread brain networks and the P3b are more likely associated with task-related processes.SIGNIFICANCE STATEMENT How does our brain generate visual consciousness-the subjective experience of what it is like to see, for example, a face? To date, it is hotly debated whether it emerges early in sensory brain regions or later when a widespread frontoparietal network is activated. Here, we use simultaneous fMRI and EEG for high spatial and temporal resolution and demonstrate that conscious face perception is predominantly linked to early and occipitotemporal processes, but also prefrontal activity. Task-related processes (e.g., decision-making), on the other hand, elicit brain-wide activations including late and strong frontoparietal activity. These findings challenge numerous previous studies and highlight the importance of investigating the neural correlates of consciousness in the absence of task relevance.

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.

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.

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.

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.

Differential Effects of Awareness and Task Relevance on Early and Late ERPs in a No-Report Visual Oddball Paradigm.

To date it is poorly understood how and when deviance processing interacts with awareness and task relevance. Furthermore, an important issue in the study of consciousness is the prevalent confound of conscious perception with the requirement of reporting it. This study addresses these topics using a no-report inattentional blindness paradigm with a visual oddball sequence of geometrical shapes presented to male and female human participants. Electrophysiological responses were obtained in three physically identical Phases A-C that differed only with respect to the instructions: (A) participants were uninformed about the shapes and attended an unrelated foreground task (inattentional blind), (B) were informed about the shapes but still attended the foreground task, and (C) attended the shapes. Conscious processing of shapes was indexed by the visual awareness negativity but not a P3. Deviance processing was associated with the visual mismatch negativity independently of consciousness and task relevance. The oddball P3, however, only emerged when the stimuli were task relevant, and was absent for consciously perceived but task irrelevant stimuli. The P3 thus does not represent a reliable marker of stimulus awareness. This result pattern supports the view of hierarchical predictive processing, where lower levels display automatic deviance processing, whereas higher levels require attention and task relevance.SIGNIFICANCE STATEMENT To react to potentially important changes in our environment it is fundamental to detect deviations from regularities of sensory input. It has yet to be understood how awareness and task relevance of this input interact with deviance processing. We investigated the role of awareness in deviance detection while at the same time circumventing the confound of awareness and report by means of a no-report paradigm. Our results suggest that early processes are elicited automatically, whereas, contrary to prominent theories, late processes do not depend on awareness but on task-based attention.

Dissociating neural correlates of consciousness and task relevance during auditory processing.

In recent years, several ERP components have been identified as potential neural correlates of consciousness (NCC), including early negativities and late positivities. Based on experiments in the visual modality, it has recently been shown that awareness is often confounded with reporting it, possibly overestimating the NCC. It is unknown whether similar constraints also exist in the auditory modality. In order to address this gap, we presented spoken words in a sustained inattentional deafness paradigm. Electrophysiological responses were obtained in three physically identical experimental conditions that differed only with respect to the participants' instructions. Participants were either left uninformed or informed about the presence of spoken words while confronted with an auditory distractor task (U/I condition), informed about the words while exposed to the same task as before (I condition), or requested to respond to the now task-relevant speech stimuli (TR condition). After completion of the U/I condition, only informed participants reported awareness of the words. In ERPs, awareness of words in the U/I and I condition was accompanied by an anterior auditory awareness negativity (AAN). Only when stimuli were task-relevant, i.e., during the TR condition, late positivities emerged. Taken together, these results indicate that early negativities but not late positivities index awareness across sensory modalities. Thus, they provide evidence for a recurrent processing framework, which highlights the importance of early sensory processing in conscious perception.

Fearful face scrambles increase early visual sensory processing in the absence of face information.

Emotional facial expressions elicit distinct increased early electrophysiological responses. Many studies report even emotional modulations of very early sensory processing at about 80 and 100 ms after stimulus presentation, indexed by the P1. These early effects are often interpreted to index differential responses to biologically relevant expressions. Since specific spatial frequencies differ between fearful and neutral expressions, it has recently been suggested that these early modulations are substantially driven by such low-level visual differences. However, it remains unclear whether similar P1 effects are also observed in experiments in which no recognizable face information is presented at all. This study investigated this question and explored also whether any effects depend on colour information and attentional conditions. Participants (N = 20) performed a continuous perceptual task of low or high difficultly and were presented with task-irrelevant black/white and colour images of fearful and neutral faces, rendered unrecognizable by doing Fourier phase transformation. ERP findings revealed increased P1 amplitudes for fearful scrambles regardless of experimental conditions. Taken together, our findings show early emotional effects in the absence of any facial expression. Specific low-level frequency information seems to increase P1 amplitudes which thus might have implications for the interpretation of very early sensory emotional expression effects.

Electrophysiological responses to negative evaluative person-knowledge: Effects of individual differences.

Faces transmit rich information about a unique personal identity. Recent studies examined how negative evaluative information affects event-related potentials (ERPs), the relevance of individual differences, such as trait anxiety, neuroticism, or agreeableness, for these effects is unclear. In this preregistered study, participants (N = 80) were presented with neutral faces, either associated with highly negative or neutral biographical information. Faces were shown under three different task conditions that varied the attentional focus on face-unrelated features, perceptual face information, or emotional information. Results showed a task-independent increase of the N170 component for faces associated with negative information, while interactions occurred for the Early Posterior Negativity (EPN) and the Late Positive Potential (LPP), showing ERP differences only when paying attention to the evaluative information. Trait anxiety and neuroticism did not influence ERP differences. Low agreeableness increased EPN differences during perceptual distraction. Thus, we observed that low agreeableness leads to early increased processing of potentially hostile faces, although participants were required to attend to a face-unrelated feature.

Let's Talk About Each Other: Neural Responses to Dissenting Personality Evaluations Based on Real Dyadic Interactions.

Dyadic interactions are associated with the exchange of personality-related messages, which can be congruent or incongruent with one's self-view. In the current preregistered study (N = 52), we investigated event-related potentials (ERPs) toward real social evaluations in order to uncover the neural mechanisms underlying the processing of congruent and incongruent evaluative feedback. Participants interacted first, and then during an electroencephalogram (EEG) session, they received evaluations from their interaction partner that were either congruent or incongruent with their own ratings. Findings show potentiated processing of self-related incongruent negative evaluations at early time points (N1) followed by increased processing of both incongruent negative and positive evaluations at midlatency time windows (early posterior negativity) and a prioritized processing of self-related incongruent positive evaluations at late time points (feedback-related P3, late positive potential). These findings reveal that, after real social interactions, evaluative feedback about oneself that violates one's self-view modulates all processing stages with an early negativity and a late positivity bias.

Charged With a Crime: The Neuronal Signature of Processing Negatively Evaluated Faces Under Different Attentional Conditions.

Our brains rapidly respond to human faces and can differentiate between many identities, retrieving rich semantic emotional-knowledge information. Studies provide a mixed picture of how such information affects event-related potentials (ERPs). We systematically examined the effect of feature-based attention on ERP modulations to briefly presented faces of individuals associated with a crime. The tasks required participants (N = 40 adults) to discriminate the orientation of lines overlaid onto the face, the age of the face, or emotional information associated with the face. Negative faces amplified the N170 ERP component during all tasks, whereas the early posterior negativity (EPN) and late positive potential (LPP) components were increased only when the emotional information was attended to. These findings suggest that during early configural analyses (N170), evaluative information potentiates face processing regardless of feature-based attention. During intermediate, only partially resource-dependent, processing stages (EPN) and late stages of elaborate stimulus processing (LPP), attention to the acquired emotional information is necessary for amplified processing of negatively evaluated faces.

Inter-individual differences in trait anxiety shape the functional connectivity between the bed nucleus of the stria terminalis and the amygdala during brief threat processing.

An influential framework suggests that the central nucleus of the amygdala (CeA) is involved in phasic responses to threat, while the bed nucleus of the stria terminalis (BNST) mediates sustained anxiety. However, this model has been questioned, proposing that the role of the BNST is not limited to sustained threat contexts. Rather, amygdala and BNST also seem to work in concert in the processing of discrete and briefly presented threat-related stimuli, likely dependent on inter-individual differences in anxiety. A direct test of this assumption with sufficient experimental power is missing in human research and the degree to which individual differences in trait anxiety moderate phasic responses and functional connectivity of amygdala and BNST during threat processing remains unclear. The current event-related functional magnetic resonance imaging (fMRI) study investigated activation and connectivity of amygdala and BNST, as well as modulating effects of trait anxiety, during processing of briefly presented threat-related relative to neutral standardized pictures in 93 psychiatrically healthy individuals. Both amygdala and BNST activation was increased during presentation of threat-related relative to neutral pictures. Furthermore, functional connectivity between BNST and amygdala in response to threat was positively associated with trait anxiety. These findings suggest that amygdala and BNST form a functional unit during phasic threat processing whereby their connectivity is shaped by inter-individual differences in trait anxiety.

Early brain responses to affective faces: A simultaneous EEG-fMRI study.

The spatio-temporal neural basis of earliest differentiation between emotional and neutral facial expressions is a matter of debate. The present study used concurrent electroencephalography (EEG) and functional magnetic resonance imaging (fMRI) in order to investigate the 'when' and 'where' of earliest prioritization of emotional over neutral expressions. We measured event-related potentials (ERPs) and blood oxygen dependent (BOLD) signal changes in response to facial expressions of varying emotional intensity and different valence categories. Facial expressions were presented superimposed by two horizontal bars and participants engaged in a focal bars task (low load, high load), in order to manipulate the availability of attentional resources during face perception. EEG data revealed the earliest expression effects in the P1 range (76-128 ms) as a parametric function of stimulus arousal independent of load conditions. Conventional fMRI data analysis also demonstrated significant modulations as a function of stimulus arousal, independent of load, in amygdala, superior temporal sulcus, fusiform gyrus and lateral occipital cortex. Correspondingly, EEG-informed fMRI analysis revealed a significant positive correlation between single-trial P1 amplitudes and BOLD responses in amygdala and lateral posterior occipital cortex. Our results are in line with the hypothesis of the amygdala as fast responding relevance detector and corresponding effects in early visual face processing areas across facial expressions and load conditions.

No differential effects to facial expressions under continuous flash suppression: An event-related potentials study.

In recent years continuous flash suppression (CFS) has become a popular "blinding" technique for the investigation of nonconscious affective processing since it elicits potent and long-lasting suppression of conscious visual perception. While the majority of studies provides some positive evidence for nonconscious affective processing, there are also studies reporting their absence. Several methodological variations may give rise to this discrepancy: with respect to the experimental paradigm these variations pertain to the likelihood of residual stimulus visibility on the level of individual participants and single trials. Concerning the statistical analysis they relate to the procedures applied to assess whether detection performance is at chance level and whether the outcome measure does or does not depend on the affective stimulus category. In the present study we determined individual eye dominance and individual stimulus contrast in pretests, measured objective and subjective awareness online and applied Bayesian statistics to estimate the likelihood for the null hypothesis. Event-related potentials (ERPs) were measured while participants were subjected to fearful, happy, and neutral faces in a conscious as well as in a nonconscious CFS condition. In the conscious condition, expected emotion effects were observed in the ERP components N170/EPN and LPP. However, despite high statistical power, no effects of emotional expression were found in the nonconscious condition and the absence of nonconscious affective processing under the tested conditions was substantially more likely than its presence. We discuss whether CFS disrupts affective processing completely if thoroughly applied or whether positive and negative findings should be integrated under a two-threshold framework of nonconscious processing.

Brain responses to disorder-related visual threat in panic disorder.

Panic disorder (PD) patients show aberrant neural responses to threatening stimuli in an extended fear network, but results are only partially comparable, and studies implementing disorder-related visual scenes are lacking as stimuli. The neural responses and functional connectivity to a newly developed set of disorder-related, ecologically valid scenes as compared with matched neutral visual scenes, using event-related functional magnetic resonance imaging (fMRI) in 26 PD patients and 26 healthy controls (HC) were investigated. PD patients versus HC showed hyperactivation in an extended fear network comprising brainstem, insula, thalamus, anterior, and mid-cingulate cortex and (dorso-)medial prefrontal cortex for disorder-related versus neutral scenes. Amygdala differences between groups failed significance. Subjective levels of anxiety significantly correlated with brainstem activation in PD patients. Analysis of functional connectivity by means of beta series correlation revealed no emotion-specific alterations in connectivity in PD patients versus HC. The results suggest that subjective anxiety evoked by external stimuli is directly related to altered activation in the homeostatic alarm system in PD. With novel disorder-related stimuli, the study sheds new light on the neural underpinnings of pathological threat processing in PD. Hum Brain Mapp 37:4439-4453, 2016. © 2016 Wiley Periodicals, Inc.