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.

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.

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.

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.

Lateralized deficits in arousal processing after insula lesions: Behavioral and autonomic evidence.

A large body of evidence ascribes a pivotal role in emotion processing to the insular cortex. However, the complex structure and lateralization of emotional deficits following insular damage are not understood. Here, we investigated emotional ratings of valence and arousal and skin conductance responses (SCR) to a graded series of emotionally arousing scenes in patients with left (n = 10) or right (n = 9) insular damage and in healthy controls (n = 18). We found a significant reduction in overall SCRs, arousal ratings and valence extremity scores in right-lesioned patients, as compared to left-lesioned patients and healthy controls. The degree of right insular damage was significantly correlated with the degree of arousal, SCR and extremity attenuation. Additional analyses of correlations between subjective arousal ratings resp. SCR and normative arousal ratings revealed that both lesion groups had evaluative and physiological difficulties to discover changes in stimulus arousal. Although no group differences emerged on overall ratings of valence, analysis of correlations between subjective and normative valence ratings displayed markedly reduced accuracy in right-lesioned patients, as compared to left-lesioned patients and healthy controls. Our findings support the hypothesis that the left and right insulae subserve different functions in emotion processing, potentially due to asymmetrical representations of autonomic information in the left and right human forebrain. The right insula may serve as integral node for sympathetic arousal and cognitive-affective processing.

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.

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.

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.

Amygdala enlargement and emotional responses in (autoimmune) temporal lobe epilepsy.

Temporal lobe epilepsy with amygdala enlargement (TLE-AE) is increasingly recognized as a distinct adult electroclinical syndrome. However, functional consequences of morphological alterations of the amygdala in TLE-AE are poorly understood. Here, two emotional stimulation designs were employed to investigate subjective emotional rating and skin conductance responses in a sample of treatment-naïve patients with suspected or confirmed autoimmune TLE-AE (n = 12) in comparison to a healthy control group (n = 16). A subgroup of patients completed follow-up measurements after treatment. As compared to healthy controls, patients with suspected or confirmed autoimmune TLE-AE showed markedly attenuated skin conductance responses and arousal ratings, especially pronounced for anxiety-inducing stimuli. The degree of right amygdala enlargement was significantly correlated with the degree of autonomic arousal attenuation. Furthermore, a decline of amygdala enlargement following prompt aggressive immunotherapy in one patient suffering from severe confirmed autoimmune TLE-AE with a very recent clinical onset was accompanied by a significant improvement of autonomic responses. Findings suggest dual impairments of autonomic and cognitive discrimination of stimulus arousal as hallmarks of emotional processing in TLE-AE. Emotional responses might, at least partially, recover after successful treatment, as implied by first single case data.

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.