He Who Seeks Finds (Bodily Signals): German Validation of the Interoceptive Attention Scale (IATS) and its Relationship with Subclinical Psychopathology.

Alterations in interoception have been linked to psychopathology. Recent findings suggest that both the attention to and the accuracy of, interoceptive perceptions may be oppositely related to subclinical symptomatology. Thus, providing well-validated tools that tap into these interoceptive processes is crucial for understanding the relation between interoceptive processing and subclinical psychopathology. In the current study (N = 642), we aimed to (1) validate the German version of the Interoceptive Attention Scale (IATS; Gabriele et al., 2022), and (2) test the differential association of self-reported interoceptive attention and accuracy with subclinical symptomatology, including alexithymia, depressive, and anxious symptomatology. We observed that a one-factor solution is a well-fitting model for the IATS. Further, the IATS showed good internal consistency, convergent, and divergent validity, but poor test-retest reliability. Self-reported interoceptive attention and accuracy were unrelated to each other. However, IATS scores were positively related to all measures of psychopathology (except depressive symptomatology), whereas self-reported interoceptive accuracy scores showed negative or nonsignificant relations with these. Our data suggest that the IATS is a good instrument to measure self-report interoceptive attention in the German population. Further, we highlight the need to distinguish between constructs of interoception to better understand the relation between interoception and psychopathology.

Transcutaneous auricular vagus nerve stimulation modulates the processing of interoceptive prediction error signals and their role in allostatic regulation.

It has recently been suggested that predictive processing principles may apply to interoception, defined as the processing of hormonal, autonomic, visceral, and immunological signals. In the current study, we aimed at providing empirical evidence for the role of cardiac interoceptive prediction errors signals on allostatic adjustments, using transcutaneous auricular vagus nerve stimulation (taVNS) as a tool to modulate the processing of interoceptive afferents. In a within-subject design, participants performed a cardiac-related interoceptive task (heartbeat counting task) under taVNS and sham stimulation, spaced 1-week apart. We observed that taVNS, in contrast to sham stimulation, facilitated the maintenance of interoceptive accuracy levels over time (from the initial, stimulation-free, baseline block to subsequent stimulation blocks), suggesting that vagus nerve stimulation may have helped to maintain engagement to cardiac afferent signals. During the interoceptive task, taVNS compared to sham, produced higher heart-evoked potentials (HEP) amplitudes, a potential readout measure of cardiac-related prediction error processing. Further analyses revealed that the positive relation between interoceptive accuracy and allostatic adjustments-as measured by heart rate variability (HRV)-was mediated by HEP amplitudes. Providing initial support for predictive processing accounts of interoception, our results suggest that the stimulation of the vagus nerve may increase the precision with which interoceptive signals are processed, favoring their influence on allostatic adjustments.

Attention-affect interactions and triarchic psychopathy: New electrophysiological insights from the late positive potential.

One of the most prominent characteristics of psychopathy is a reduced processing of emotionally relevant information. However, it is still unclear how attentional mechanisms may modulate this deficit. The current study aimed to examine the impact of attentional focus on emotion processing in relation to the triarchic constructs of boldness, meanness, and disinhibition. Participants performed two tasks in which pleasant, neutral, and unpleasant framed pictures were presented. In the first task, participants were required to indicate the color of the frame (alternative-focus task), whereas in the second task they were instructed to indicate the emotional category of the image (affect-focus task). The Late Positive Potential (LPP) was used as an index of sustained engagement of attention to affective material. Confirming a successful task manipulation, we observed reduced LPP amplitudes, particularly for affective relevant material, in the alternative-focus task compared to the affect-focus task. Most interestingly, our results evidenced that trait meanness scores were associated with blunted elaborative processing of affective material (both appetitive and aversive) when this information was task-relevant (affect-focus task), but not when it was task-irrelevant (alternative-focus task). These findings indicate that high mean individuals are characterized by blunted elaborative processing of affective stimuli when their motivational relevance is determined in a top-down manner (i.e., when it is task-relevant). Our results highlight the need for further studying of the bottom-up and top-down dynamics of emotional attention in psychopathy.

Evidence for a modulating effect of transcutaneous auricular vagus nerve stimulation (taVNS) on salivary alpha-amylase as indirect noradrenergic marker: A pooled mega-analysis.

BACKGROUND: Non-invasive transcutaneous auricular vagus nerve stimulation (taVNS) has received tremendous attention as a potential neuromodulator of cognitive and affective functions, which likely exerts its effects via activation of the locus coeruleus-noradrenaline (LC-NA) system. Reliable effects of taVNS on markers of LC-NA system activity, however, have not been demonstrated yet. METHODS: The aim of the present study was to overcome previous limitations by pooling raw data from a large sample of ten taVNS studies (371 healthy participants) that collected salivary alpha-amylase (sAA) as a potential marker of central NA release. RESULTS: While a meta-analytic approach using summary statistics did not yield any significant effects, linear mixed model analyses showed that afferent stimulation of the vagus nerve via taVNS increased sAA levels compared to sham stimulation (b = 0.16, SE = 0.05, p = 0.001). When considering potential confounders of sAA, we further replicated previous findings on the diurnal trajectory of sAA activity. CONCLUSION(S): Vagal activation via taVNS increases sAA release compared to sham stimulation, which likely substantiates the assumption that taVNS triggers NA release. Moreover, our results highlight the benefits of data pooling and data sharing in order to allow stronger conclusions in research.

New insights on the correspondence between subjective affective experience and physiological responses from representational similarity analysis.

Classical views suggest that experienced affect is related to a specific bodily response, whereas recent perspectives challenge this view postulating that similar affective experiences rather evoke different physiological responses. To further advance this debate in the field, we used representational similarity analysis to investigate the correspondence between subjective affect (arousal and valence ratings) and physiological reactions (skin conductance response [SCR], startle blink response, heart rate, and corrugator activity) across various emotion induction contexts (picture viewing task, sound listening task, and imagery task). Significant similarities were exclusively observed between SCR and arousal in the picture viewing task. However, none of the other physiological measures showed a significant relation with valence and arousal ratings in any of the tasks. These findings are discussed within the framework of the Populations hypothesis, suggesting that physiological responses do not depend on the experienced affect but are directly associated with the context in which they are evoked.

Memory advantage for untrustworthy faces: Replication across lab- and web-based studies.

The Covid-19 pandemic imposed new constraints on empirical research and forced researchers to transfer from traditional laboratory research to the online environment. This study tested the validity of a web-based episodic memory paradigm by comparing participants' memory performance for trustworthy and untrustworthy facial stimuli in a supervised laboratory setting and an unsupervised web setting. Consistent with previous results, we observed enhanced episodic memory for untrustworthy compared to trustworthy faces. Most importantly, this memory bias was comparable in the online and the laboratory experiment, suggesting that web-based procedures are a promising tool for memory research.

The Role of Interoceptive Sensibility and Emotional Conceptualization for the Experience of Emotions.

The theory of constructed emotions suggests that different psychological components, including core affect (mental and neural representations of bodily changes), and conceptualization (meaning-making based on prior experiences and semantic knowledge), are involved in the formation of emotions. However, little is known about their role in experiencing emotions. In the current study, we investigated how individual differences in interoceptive sensibility and emotional conceptualization (as potential correlates of these components) interact to moderate three important aspects of emotional experiences: emotional intensity (strength of emotion felt), arousal (degree of activation), and granularity (ability to differentiate emotions with precision). To this end, participants completed a series of questionnaires assessing interoceptive sensibility and emotional conceptualization and underwent two emotion experience tasks, which included standardized material (emotion differentiation task; ED task) and self-experienced episodes (day reconstruction method; DRM). Correlational analysis showed that individual differences in interoceptive sensibility and emotional conceptualization were related to each other. Principal Component Analysis (PCA) revealed two independent factors that were referred to as sensibility and monitoring. The Sensibility factor, interpreted as beliefs about the accuracy of an individual in detecting internal physiological and emotional states, predicted higher granularity for negative words. The Monitoring factor, interpreted as the tendency to focus on the internal states of an individual, was negatively related to emotional granularity and intensity. Additionally, Sensibility scores were more strongly associated with greater well-being and adaptability measures than Monitoring scores. Our results indicate that independent processes underlying individual differences in interoceptive sensibility and emotional conceptualization contribute to emotion experiencing.

Establishment of Emotional Memories Is Mediated by Vagal Nerve Activation: Evidence from Noninvasive taVNS.

Emotional memories are better remembered than neutral ones, but the mechanisms leading to this memory bias are not well understood in humans yet. Based on animal research, it is suggested that the memory-enhancing effect of emotion is based on central noradrenergic release, which is triggered by afferent vagal nerve activation. To test the causal link between vagus nerve activation and emotional memory in humans, we applied continuous noninvasive transcutaneous auricular vagus nerve stimulation (taVNS) during exposure to emotional arousing and neutral scenes and tested subsequent, long-term recognition memory after 1 week. We found that taVNS, compared with sham, increased recollection-based memory performance for emotional, but not neutral, material. These findings were complemented by larger recollection-related brain potentials (parietal ERP Old/New effect) during retrieval of emotional scenes encoded under taVNS, compared with sham. Furthermore, brain potentials recorded during encoding also revealed that taVNS facilitated early attentional discrimination between emotional and neutral scenes. Extending animal research, our behavioral and neural findings confirm a modulatory influence of the vagus nerve in emotional memory formation in humans.SIGNIFICANCE STATEMENT Emotionally relevant information elicits stronger and more enduring memories than nonrelevant information. Animal research has shown that this memory-enhancing effect of emotion is related to the noradrenergic activation in the brain, which is triggered by afferent fibers of the vagus nerve (VN). In the current study, we show that noninvasive transcutaneous auricular VN stimulation enhances recollection-based memory formation specifically for emotionally relevant information as indicated by behavioral and electrophysiological indices. These human findings give novel insights into the mechanisms underlying the establishment of emotional episodic memories by confirming the causal link between the VN and memory formation which may help understand the neural mechanisms underlying disorders associated with altered memory functions and develop treatment options.

Transcutaneous Vagus Nerve Stimulation (tVNS) Improves High-Confidence Recognition Memory but Not Emotional Word Processing.

Previous clinical research found that invasive vagus nerve stimulation (VNS) enhanced word recognition memory in epileptic patients, an effect assumed to be related to the activation of brainstem arousal systems. In this study, we applied non-invasive transcutaneous auricular VNS (tVNS) to replicate and extend the previous work. Using a single-blind, randomized, between-subject design, 60 healthy volunteers received active or sham stimulation during a lexical decision task, in which emotional and neutral stimuli were classified as words or non-words. In a subsequent recognition memory task (1 day after stimulation), participants' memory performance on these words and their subjective memory confidence were tested. Salivary alpha-amylase (sAA) levels, a putative indirect measure of central noradrenergic activation, were also measured before and after stimulation. During encoding, pleasant words were more accurately detected than neutral and unpleasant words. However, no tVNS effects were observed on task performance or on overall sAA level changes. tVNS also did not modulate overall recognition memory, which was particularly enhanced for pleasant emotional words. However, when hit rates were split based on confidence ratings reflecting familiarity- and recollection-based memory, higher recollection-based memory performance (irrespective of emotional category) was observed during active stimulation than during sham stimulation. To summarize, we replicated prior findings of enhanced processing and memory for emotional (pleasant) words. Whereas tVNS showed no effects on word processing, subtle effects on recollection-based memory performance emerged, which may indicate that tVNS facilitates hippocampus-mediated consolidation processes.

International Consensus Based Review and Recommendations for Minimum Reporting Standards in Research on Transcutaneous Vagus Nerve Stimulation (Version 2020).

Given its non-invasive nature, there is increasing interest in the use of transcutaneous vagus nerve stimulation (tVNS) across basic, translational and clinical research. Contemporaneously, tVNS can be achieved by stimulating either the auricular branch or the cervical bundle of the vagus nerve, referred to as transcutaneous auricular vagus nerve stimulation(VNS) and transcutaneous cervical VNS, respectively. In order to advance the field in a systematic manner, studies using these technologies need to adequately report sufficient methodological detail to enable comparison of results between studies, replication of studies, as well as enhancing study participant safety. We systematically reviewed the existing tVNS literature to evaluate current reporting practices. Based on this review, and consensus among participating authors, we propose a set of minimal reporting items to guide future tVNS studies. The suggested items address specific technical aspects of the device and stimulation parameters. We also cover general recommendations including inclusion and exclusion criteria for participants, outcome parameters and the detailed reporting of side effects. Furthermore, we review strategies used to identify the optimal stimulation parameters for a given research setting and summarize ongoing developments in animal research with potential implications for the application of tVNS in humans. Finally, we discuss the potential of tVNS in future research as well as the associated challenges across several disciplines in research and clinical practice.

Neural correlates of emotion-attention interactions: From perception, learning, and memory to social cognition, individual differences, and training interventions.

Due to their ability to capture attention, emotional stimuli tend to benefit from enhanced perceptual processing, which can be helpful when such stimuli are task-relevant but hindering when they are task-irrelevant. Altered emotion-attention interactions have been associated with symptoms of affective disturbances, and emerging research focuses on improving emotion-attention interactions to prevent or treat affective disorders. In line with the Human Affectome Project's emphasis on linguistic components, we also analyzed the language used to describe attention-related aspects of emotion, and highlighted terms related to domains such as conscious awareness, motivational effects of attention, social attention, and emotion regulation. These terms were discussed within a broader review of available evidence regarding the neural correlates of (1) Emotion-Attention Interactions in Perception, (2) Emotion-Attention Interactions in Learning and Memory, (3) Individual Differences in Emotion-Attention Interactions, and (4) Training and Interventions to Optimize Emotion-Attention Interactions. This comprehensive approach enabled an integrative overview of the current knowledge regarding the mechanisms of emotion-attention interactions at multiple levels of analysis, and identification of emerging directions for future investigations.

Item and source memory for emotional associates is mediated by different retrieval processes.

Recent event-related potential (ERP) data showed that neutral objects encoded in emotional background pictures were better remembered than objects encoded in neutral contexts, when recognition memory was tested one week later. In the present study, we investigated whether this long-term memory advantage for items is also associated with correct memory for contextual source details. Furthermore, we were interested in the possibly dissociable contribution of familiarity and recollection processes (using a Remember/Know procedure). The results revealed that item memory performance was mainly driven by the subjective experience of familiarity, irrespective of whether the objects were previously encoded in emotional or neutral contexts. Correct source memory for the associated background picture, however, was driven by recollection and enhanced when the content was emotional. In ERPs, correctly recognized old objects evoked frontal ERP Old/New effects (300-500 ms), irrespective of context category. As in our previous study (Ventura-Bort et al., 2016b), retrieval for objects from emotional contexts was associated with larger parietal Old/New differences (600-800 ms), indicating stronger involvement of recollection. Thus, the results suggest a stronger contribution of recollection-based retrieval to item and contextual background source memory for neutral information associated with an emotional event.

Behavioral and neural evidence of enhanced long-term memory for untrustworthy faces.

In daily life, we automatically form impressions of other individuals on basis of subtle facial features that convey trustworthiness. Because these face-based judgements influence current and future social interactions, we investigated how perceived trustworthiness of faces affects long-term memory using event-related potentials (ERPs). In the current study, participants incidentally viewed 60 neutral faces differing in trustworthiness, and one week later, performed a surprise recognition memory task, in which the same old faces were presented intermixed with novel ones. We found that after one week untrustworthy faces were better recognized than trustworthy faces and that untrustworthy faces prompted early (350-550 ms) enhanced frontal ERP old/new differences (larger positivity for correctly remembered old faces, compared to novel ones) during recognition. Our findings point toward an enhanced long-lasting, likely familiarity-based, memory for untrustworthy faces. Even when trust judgments about a person do not necessarily need to be accurate, a fast access to memories predicting potential harm may be important to guide social behaviour in daily life.

Enhanced spontaneous retrieval of cues from emotional events: An ERP study.

Recent evidence points to enhanced episodic memory retrieval not only for emotional items but also for neutral information encoded in emotional contexts. However, prior research only tested instructed explicit recognition, and hence here we investigated whether memory retrieval is also heightened for cues from emotional contexts when retrieval is not explicitly probed. During the first session of a two-session experiment, neutral objects were presented on different background scenes varying in emotional and neutral contents. One week later, objects were presented again (with no background) intermixed with novel objects. In both sessions, participants were instructed to attentively watch the stimuli (free viewing procedure), and during the second session, ERPs were also collected to measure the ERP Old/New effect, an electrophysiological correlate of episodic memory retrieval. Analyses were performed using cluster-based permutation tests in order to identify reliable spatio-temporal ERP differences. Based on this approach, old relative to new objects, were associated with larger ERP positivity in an early (364-744 ms) and late time window (760-1148 ms) over distinct central electrode clusters. Interestingly, significant late ERP Old/New differences were only observed for objects previously encoded with emotional, but not neutral scenes (504 to 1144 ms). Because these ERP differences were observed in a non-instructed retrieval context, our results indicate that long-term, spontaneous retrieval for neutral objects, is particularly heightened if encoded within emotionally salient contextual information. These findings may assist in understanding mechanisms underlying spontaneous retrieval of emotional associates and the utility of ERPs to study maladaptive involuntary memories in trauma- and stress-related disorders.

Chronic stress and emotion: Differential effects on attentional processing and recognition memory.

Previous research indicates that acute stress around the time of learning facilitates attention and memory for emotionally salient information. Despite accumulating evidence for these acute stress effects, less is known about the role of chronic stress. In the present study, we therefore tested emotional and neutral scene processing and later recognition memory in female participants using hair cortisol concentrations as a biological marker for chronic stress. Event-related potentials recorded during picture viewing indicated enhanced late positive potentials (LPPs) for emotional, relative to neutral contents. These brain potentials varied as a function of long-term hair cortisol levels: hair-cortisol levels were positively related to overall LPP amplitudes. Results from recognition memory testing one week after encoding revealed better memory for emotional relative to neutral scenes. Hair-cortisol levels, however, were related to poorer memory accuracy. Taken together, our results indicate that chronic stress enhanced attentional processing during encoding of new stimuli and impaired later recognition memory. Results are discussed with regard to putatively opposite effects of chronic stress on certain brain regions (e.g., amygdala and hippocampus).

Effects of Transcutaneous Vagus Nerve Stimulation (tVNS) on the P300 and Alpha-Amylase Level: A Pilot Study.

Recent research suggests that the P3b may be closely related to the activation of the locus coeruleus-norepinephrine (LC-NE) system. To further study the potential association, we applied a novel technique, the non-invasive transcutaneous vagus nerve stimulation (tVNS), which is speculated to increase noradrenaline levels. Using a within-subject cross-over design, 20 healthy participants received continuous tVNS and sham stimulation on two consecutive days (stimulation counterbalanced across participants) while performing a visual oddball task. During stimulation, oval non-targets (standard), normal-head (easy) and rotated-head (difficult) targets, as well as novel stimuli (scenes) were presented. As an indirect marker of noradrenergic activation we also collected salivary alpha-amylase (sAA) before and after stimulation. Results showed larger P3b amplitudes for target, relative to standard stimuli, irrespective of stimulation condition. Exploratory post hoc analyses, however, revealed that, in comparison to standard stimuli, easy (but not difficult) targets produced larger P3b (but not P3a) amplitudes during active tVNS, compared to sham stimulation. For sAA levels, although main analyses did not show differential effects of stimulation, direct testing revealed that tVNS (but not sham stimulation) increased sAA levels after stimulation. Additionally, larger differences between tVNS and sham stimulation in P3b magnitudes for easy targets were associated with larger increase in sAA levels after tVNS, but not after sham stimulation. Despite preliminary evidence for a modulatory influence of tVNS on the P3b, which may be partly mediated by activation of the noradrenergic system, additional research in this field is clearly warranted. Future studies need to clarify whether tVNS also facilitates other processes, such as learning and memory, and whether tVNS can be used as therapeutic tool.

Transcutaneous vagus nerve stimulation (tVNS) enhances conflict-triggered adjustment of cognitive control.

Response conflicts play a prominent role in the flexible adaptation of behavior as they represent context-signals that indicate the necessity for the recruitment of cognitive control. Previous studies have highlighted the functional roles of the affectively aversive and arousing quality of the conflict signal in triggering the adaptation process. To further test this potential link with arousal, participants performed a response conflict task in two separate sessions with either transcutaneous vagus nerve stimulation (tVNS), which is assumed to activate the locus coeruleus-noradrenaline (LC-NE) system, or with neutral sham stimulation. In both sessions the N2 and P3 event-related potentials (ERP) were assessed. In line with previous findings, conflict interference, the N2 and P3 amplitude were reduced after conflict. Most importantly, this adaptation to conflict was enhanced under tVNS compared to sham stimulation for conflict interference and the N2 amplitude. No effect of tVNS on the P3 component was found. These findings suggest that tVNS increases behavioral and electrophysiological markers of adaptation to conflict. Results are discussed in the context of the potentially underlying LC-NE and other neuromodulatory (e.g., GABA) systems. The present findings add important pieces to the understanding of the neurophysiological mechanisms of conflict-triggered adjustment of cognitive control.

Event-related potentials of emotional and neutral memories: The role of encoding position and delayed testing.

Previous research found that memory is not only better for emotional information but also for neutral information that has been encoded in the context of an emotional event. In the present ERP study, we investigated two factors that may influence memory for neutral and emotional items: temporal proximity between emotional and neutral items during encoding, and retention interval (immediate vs. delayed). Forty-nine female participants incidentally encoded 36 unpleasant and 108 neutral pictures (36 neutral pictures preceded an unpleasant picture, 36 followed an unpleasant picture, and 36 neutral pictures were preceded and followed by neutral pictures) and participated in a recognition memory task either immediately (N = 24) or 1 week (N = 25) after encoding. Results showed better memory for emotional pictures relative to neutral pictures. In accordance, enhanced centroparietal old/new differences (500-900 ms) during recognition were observed for unpleasant compared to neutral pictures, most pronounced for the 1-week interval. Picture position effects, however, were only subtle. During encoding, late positive potentials for neutral pictures were slightly lower for neutral pictures following unpleasant ones, but only at trend level. To summarize, we could replicate and extend previous ERP findings showing that emotionally arousing events are better recollected than neutral events, particularly when memory is tested after longer retention intervals. Picture position during encoding, however, had only small effects on elaborative processing and no effects on memory retrieval.

When neutral turns significant: brain dynamics of rapidly formed associations between neutral stimuli and emotional contexts.

The ability to associate neutral stimuli with motivationally relevant outcomes is an important survival strategy. In this study, we used event-related potentials (ERPs) to investigate brain dynamics of associative emotional learning when participants were confronted with multiple heterogeneous information. Participants viewed 144 different objects in the context of 144 different emotional and neutral background scenes. During each trial, neutral objects were shown in isolation and then paired with the background scene. All pairings were presented twice to compare ERPs in response to neutral objects before and after single association. After single pairing, neutral objects previously encoded in the context of emotional scenes evoked a larger P100 over occipital electrodes compared to objects that were previously paired with neutral scenes. Likewise, larger late positive potentials (LPPs) were observed over parieto-occipital electrodes (450-750 ms) for objects previously associated with emotional relative to neutral contexts. The LPP - but not P100 - enhancement was also related to subjective object/context binding. Taken together, our ERP data provide evidence for fast emotional associative learning, as reflected by heightened perceptual and sustained elaborative processing for neutral information previously encountered in emotional contexts. These findings could assist in understanding binding mechanisms in stress and anxiety, as well as in addiction and eating-related disorders.