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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).

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., (© 2022 The Authors. Psychophysiology published by Wiley Periodicals LLC on behalf of Society for Psychophysiological Research.)

Previous studies on the associations between trait anxiety and amygdalar responses to threat stimuli have resulted in mixed findings, possibly due to sample characteristics, specific tasks, and analytical methods. The present functional magnetic resonance imaging (fMRI) study aimed to investigate linear or non-linear associations between trait anxiety and amygdalar responses in a sample of participants with low, medium, and high trait anxiety scores. During scanning, participants were presented with threat-related or neutral pictures and had either to solve an emotional task or an emotional-unrelated distraction task. Results showed that only during the explicit task trait anxiety was associated with right amygdalar responses to threat-related pictures as compared to neutral pictures. The best model was a cubic model with increased amygdala responses for very low and medium trait anxiety values but decreased amygdala activation for very high trait anxiety values. The findings imply a non-linear relation between trait anxiety and amygdala activation depending on task conditions., (© 2021. The Author(s).)

Fearful facial expressions are prioritized across different stages of information processing as reflected by early, mid-latency, and late components of event-related brain potentials (ERP). Trait anxiety has been proposed to modulate these responses, but it is yet unclear how such modulations depend on feature-based attention. In this preregistered study (N = 80), we investigated the effects of trait anxiety on ERP differences between fearful and neutral faces across three different tasks. Participants had to discriminate either the orientation of lines overlaid onto the faces, the gender of the face, or the emotional expression, thus increasing attention to emotionally relevant facial features across the tasks. Fearful versus neutral faces elicited increased P1 and N170 amplitudes across tasks and potentiated amplitudes when attention was directed to faces (early posterior negativity [EPN]) or the expression (EPN and late positive potential). Higher trait anxiety was related to smaller EPN differences between fearful and neutral faces during the perceptual discrimination task. This early relationship suggests reduced instead of amplified processing of fearful faces for high trait anxious participants under perceptual distraction., (© 2021 The Authors. Psychophysiology published by Wiley Periodicals LLC on behalf of Society for Psychophysiological Research.)

In a previous study, we investigated the resting-state fMRI effective connectivity (EC) between the bed nucleus of the stria terminalis (BNST) and the laterobasal (LB), centromedial (CM), and superficial (SF) amygdala. We found strong negative EC from all amygdala nuclei to the BNST, while the BNST showed positive EC to the amygdala. However, the validity of these findings remains unclear, since a reproduction in different samples has not been done. Moreover, the association of EC with measures of anxiety offers deeper insight, due to the known role of the BNST and amygdala in fear and anxiety. Here, we aimed to reproduce our previous results in three additional samples. We used spectral Dynamic Causal Modeling to estimate the EC between the BNST, the LB, CM, and SF, and its association with two measures of self-reported anxiety. Our results revealed consistency over samples with regard to the negative EC from the amygdala nuclei to the BNST, while the positive EC from BNST to the amygdala was also found, but weaker and more heterogenic. Moreover, we found the BNST-BNST EC showing a positive and the CM-BNST EC, showing a negative association with anxiety. Our study suggests a reproducible pattern of negative EC from the amygdala to the BNST along with weaker positive EC from the BNST to the amygdala. Moreover, less BNST self-inhibition and more inhibitory influence from the CM to the BNST seems to be a pattern of EC that is related to higher anxiety., (© 2020 The Authors. Human Brain Mapping published by Wiley Periodicals LLC.)

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., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Sustained anticipatory anxiety is central to Generalized Anxiety Disorder (GAD). During anticipatory anxiety, phasic threat responding appears to be mediated by the amygdala, while sustained threat responding seems related to the bed nucleus of the stria terminalis (BNST). Although sustained anticipatory anxiety in GAD patients was proposed to be associated with BNST activity alterations, firm evidence is lacking. We aimed to explore temporal characteristics of BNST and amygdala activity during threat anticipation in GAD patients. Nineteen GAD patients and nineteen healthy controls (HC) underwent functional magnetic resonance imaging (fMRI) during a temporally unpredictable threat anticipation paradigm. We defined phasic and a systematic variation of sustained response models for blood oxygen level-dependent responses during threat anticipation, to disentangle temporally dissociable involvement of the BNST and the amygdala. GAD patients relative to HC responded with increased phasic amygdala activity to onset of threat anticipation and with elevated sustained BNST activity that was delayed relative to the onset of threat anticipation. Both the amygdala and the BNST displayed altered responses during threat anticipation in GAD patients, albeit with different time courses. The results for the BNST activation hint towards its role in sustained threat responding, and contribute to a deeper understanding of pathological sustained anticipatory anxiety in GAD., (© The Author (2017). Published by Oxford University Press.)

Social context influences social decisions and outcome processing, partially depending on inter-individual differences. The present study investigated social context-dependent modulation of behavior and feedback processing in the ultimatum game (UG) in relation to inter-individual differences in social anxiety. Thirty-two healthy adults completed the UG both under social observation and without observation. Offers were allegedly either randomly generated by the computer or drawn from a pool of offers from previous human players. Overall, fewer unfair than fair offers were accepted. Observation decreased acceptance rates for unfair offers. The feedback-locked feedback-related negativity (FRN) but not the P3 was modulated by observation and fairness, with stronger differential coding of unfair/fair under observation. This effect was strongly correlated with individual levels of social anxiety, with higher levels associated with stronger differential fairness coding in the FRN under observation. Behavioral findings support negative reciprocity in the UG, suggesting that (implicit) social norms overwrite explicit task instructions even in the absence of (alleged) social interaction. Observation enhances this effect. Fairness coding in the FRN was modulated by observation as a function of social anxiety, supporting the notion that altered sensitivity to equality in a social context may contribute to social avoidance in socially anxious individuals.

It is as yet unknown if behavioral and neural correlates of performance monitoring in socially anxious individuals are affected by whether feedback is provided by a person or a computer. This fMRI study investigated modulation of feedback processing by feedback source (person vs. computer) in participants with high (HSA) (N=16) and low social anxiety (LSA) (N=16). Subjects performed a choice task in which they were informed that they would receive positive or negative feedback from a person or the computer. Subjective ratings indicated increased arousal and anxiety in HSA versus LSA, most pronounced for social and negative feedback. FMRI analyses yielded hyperactivation in ventral medial prefrontal cortex (vmPFC)/anterior cingulate cortex (ACC) and insula for social relative to computer feedback, and in mPFC/ventral ACC for positive relative to negative feedback in HSA as compared to LSA. These activation patterns are consistent with increased interoception and self-referential processing in social anxiety, especially during processing of positive feedback. Increased ACC activation in HSA to positive feedback may link to unexpectedness of (social) praise as posited in social anxiety disorder (SAD) psychopathology. Activation in rostral ACC showed a reversed pattern, with decreased activation to positive feedback in HSA, possibly indicating altered action values depending on feedback source and valence. The present findings corroborate a crucial role of mPFC for performance monitoring in social anxiety., (Copyright © 2016 Elsevier Inc. All rights reserved.)

The recognition of faces across incidences is a complex function of the human brain and a crucial ability for communication and daily interactions. This first study on ERP correlates of emotional face learning in social anxiety disorder (SAD) investigates whether the known attentional bias for threatening faces leads to a corresponding memory bias. Therefore, 21 patients with SAD and 21 healthy controls (HCs) learned faces with emotional facial expressions (neutral, happy, and angry) and were later asked to recognize these out of novel identities all presented with a neutral facial expression. EEG was recorded throughout. Behaviorally, the faces' emotional expression modulated later recognition in terms of accuracy, response times, signal detection parameters and ratings of valence, but with better performance for happy than angry faces in HC as well as in SAD. In the learning phase, attention- and memory-associated event-related potentials (ERPs) P100, N170, P200, N250/EPN, and LPP indicated enhanced processing of angry faces, which was restricted to patients with SAD in N250/EPN and LPP. In the test phase, familiarity effects emerged in N250, FN400 and LPP. While N250 was affected by learned-angry faces, FN400 and LPP reflected image learning of neutral faces, which was restricted to SAD in LPP. We replicated the attentional bias to threatening faces, which was not restricted to early ERP components, but was prolonged to later stages of conscious processing, especially in SAD. In contrast to what had been expected, sustained hypervigilance to the emotional content seems to have impaired the processing of the facial identity, resulting in a happy face advantage at the behavioral level. This could be explained by prominent models assuming separate processing of facial emotion and identity. Hypervigilance in SAD might be a disadvantage in those studies focusing on other aspects of face processing than emotion., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Socially anxious individuals tend to shift their attention away from external socially threatening cues and instead become highly self-focused. Such heightened self-focused attention has been suggested to be involved in the development and maintenance of social anxiety disorder. This study used functional magnetic resonance imaging to investigate the neural correlates of self-focused attention in 16 high socially anxious (HSA) and 16 low socially anxious (LSA) individuals. Participants were instructed to focus their attention either inwardly or outwardly during a simulated social situation. Results indicate hyperactivation of medial prefrontal cortex (mPFC), temporo-parietal junction (TPJ) and temporal pole during inward vs outward attention in HSA compared with LSA participants. Furthermore, activation of mPFC, right anterior insula, TPJ and posterior cingulate cortex was positively correlated with the trait of self-focused attention in HSA subjects. Results highlight the prominent role of the mPFC and other cortical structures in abnormal self-focused attention in social anxiety. Finally, findings for the insula suggest increased processing of bodily states that is related to the amount of habitual self-focused attention in social anxiety., (© The Author (2014). Published by Oxford University Press. For Permissions, please email: journals.permissions@oup.com.)

Fear of negative evaluation, such as negative social performance feedback, is the core symptom of social anxiety. The present study investigated the neural correlates of anticipation and perception of social performance feedback in social anxiety. High (HSA) and low (LSA) socially anxious individuals were asked to give a speech on a personally relevant topic and received standardized but appropriate expert performance feedback in a succeeding experimental session in which neural activity was measured during anticipation and presentation of negative and positive performance feedback concerning the speech performance, or a neutral feedback-unrelated control condition. HSA compared to LSA subjects reported greater anxiety during anticipation of negative feedback. Functional magnetic resonance imaging results showed deactivation of medial prefrontal brain areas during anticipation of negative feedback relative to the control and the positive condition, and medial prefrontal and insular hyperactivation during presentation of negative as well as positive feedback in HSA compared to LSA subjects. The results indicate distinct processes underlying feedback processing during anticipation and presentation of feedback in HSA as compared to LSA individuals. In line with the role of the medial prefrontal cortex in self-referential information processing and the insula in interoception, social anxiety seems to be associated with lower self-monitoring during feedback anticipation, and an increased self-focus and interoception during feedback presentation, regardless of feedback valence., (© 2014 Wiley Periodicals, Inc.)

Some people search for intense sensations such as being scared by frightening movies while others do not. The brain mechanisms underlying such inter-individual differences are not clear. Testing theoretical models, we investigated neural correlates of anxiety and the personality trait sensation seeking in 40 subjects who watched threatening and neutral scenes from scary movies during functional magnetic resonance imaging. Threat versus neutral scenes induced increased activation in anterior cingulate cortex, insula, thalamus, and visual areas. Movie-induced anxiety correlated positively with activation in dorsomedial prefrontal cortex, indicating a role for this area in the subjective experience of being scared. Sensation seeking-scores correlated positively with brain activation to threat versus neutral scenes in visual areas and in thalamus and anterior insula, i.e. regions involved in the induction and representation of arousal states. For the insula and thalamus, these outcomes were partly due to an inverse relation between sensation seeking scores and brain activation during neutral film clips. These results support models predicting cerebral hypoactivation in high sensation seekers during neutral stimulation, which may be compensated by more intense sensations such as watching scary movies., (2009 Wiley-Liss, Inc.)

Based on theoretical models, we investigated the dynamics of brain activation during anticipatory anxiety using functional magnetic resonance imaging and a combined parametric/correlational design. Subjects (16 females) anticipated the application of electrical shocks of varying intensity resulting in four different threat levels. The parametric analysis revealed an inverted U-function of activation in the ventral anterior cingulate cortex (ACC) depending on the level of threat. Furthermore, the correlation analysis showed that the association between anxiety and brain activation in the pregenual ACC was, as a tendency, positive during moderate threat but clearly negative during strong threat. Moreover, during strong threat, a positive correlation between anxiety and activation was observed in the dorsal ACC, somatosensory cortex, motor cortex, and hippocampus. These findings suggest threat dependent dynamics of brain activation in the ACC; with increased attentional avoidance during moderate threat and a switch to hypervigilant action readiness in the most anxious subjects during strong threat.

Anticipatory anxiety during expectation of phobogenic stimuli is an integral part of abnormal behaviour in phobics. The neural basis of anticipatory anxiety in specific phobia is unknown. Using functional magnetic resonance imaging (fMRI), we explored brain activation in subjects with spider phobia and in non-phobic subjects, while participants anticipated the presentation of either neutral or phobogenic visual stimuli. Subjective ratings indicated that anticipation of phobia-related stimuli was associated with increased anxiety in phobics but not in healthy subjects. FMRI results showed increased activation of the dorsal anterior cingulate cortex (ACC), insula, thalamus, and visual areas in phobics compared to controls during anticipation of phobia-relevant versus anticipation of neutral stimulation. Furthermore, for this contrast, we found also increased activation of the bed nucleus of the stria terminalis (BNST). This particular finding supports models, which propose, based on animal experiments, a critical involvement of the BNST in anticipatory anxiety. Finally, correlation analysis revealed that subjective anxiety of phobics correlated significantly with activation in rostral and dorsal ACC and the anterior medial prefrontal cortex. Thus, activation in different ACC regions and the medial prefrontal cortex seems to be specifically associated with the severity of experienced anticipatory anxiety in subjects with spider phobia.

BackgroundPanic disorder (PD) patients are constantly concerned about future panic attacks and exhibit general hypersensitivity to unpredictable threat. We aimed to reveal phasic and sustained brain responses and functional connectivity of the amygdala and the bed nucleus of the stria terminalis (BNST) during threat anticipation in PD.MethodsUsing functional magnetic resonance imaging (fMRI), we investigated 17 PD patients and 19 healthy controls (HC) during anticipation of temporally unpredictable aversive and neutral sounds. We used a phasic and sustained analysis model to disentangle temporally dissociable brain activations.ResultsPD patients compared with HC showed phasic amygdala and sustained BNST responses during anticipation of aversive v. neutral stimuli. Furthermore, increased phasic activation was observed in anterior cingulate cortex (ACC), insula and prefrontal cortex (PFC). Insula and PFC also showed sustained activation. Functional connectivity analyses revealed partly distinct phasic and sustained networks.ConclusionsWe demonstrate a role for the BNST during unpredictable threat anticipation in PD and provide first evidence for dissociation between phasic amygdala and sustained BNST activation and their functional connectivity. In line with a hypersensitivity to uncertainty in PD, our results suggest time-dependent involvement of brain regions related to fear and anxiety. [ABSTRACT FROM AUTHOR]