Current State of EEG/ERP Microstate Research.

The analysis of EEG microstates for investigating rapid whole-brain network dynamics during rest and tasks has become a standard practice in the EEG research community, leading to a substantial increase in publications across various affective, cognitive, social and clinical neuroscience domains. Recognizing the growing significance of this analytical method, the authors aim to provide the microstate research community with a comprehensive discussion on methodological standards, unresolved questions, and the functional relevance of EEG microstates. In August 2022, a conference was hosted in Bern, Switzerland, which brought together many researchers from 19 countries. During the conference, researchers gave scientific presentations and engaged in roundtable discussions aiming at establishing steps toward standardizing EEG microstate analysis methods. Encouraged by the conference's success, a special issue was launched in Brain Topography to compile the current state-of-the-art in EEG microstate research, encompassing methodological advancements, experimental findings, and clinical applications. The call for submissions for the special issue garnered 48 contributions from researchers worldwide, spanning reviews, meta-analyses, tutorials, and experimental studies. Following a rigorous peer-review process, 33 papers were accepted whose findings we will comprehensively discuss in this Editorial.

EEG-Meta-Microstates: Towards a More Objective Use of Resting-State EEG Microstate Findings Across Studies.

Over the last decade, EEG resting-state microstate analysis has evolved from a niche existence to a widely used and well-accepted methodology. The rapidly increasing body of empirical findings started to yield overarching patterns of associations of biological and psychological states and traits with specific microstate classes. However, currently, this cross-referencing among apparently similar microstate classes of different studies is typically done by "eyeballing" of printed template maps by the individual authors, lacking a systematic procedure. To improve the reliability and validity of future findings, we present a tool to systematically collect the actual data of template maps from as many published studies as possible and present them in their entirety as a matrix of spatial similarity. The tool also allows importing novel template maps and systematically extracting the findings associated with specific microstate maps from ongoing or published studies. The tool also allows importing novel template maps and systematically extracting the findings associated with specific microstate maps in the literature. The analysis of 40 included sets of template maps indicated that: (i) there is a high degree of similarity of template maps across studies, (ii) similar template maps were associated with converging empirical findings, and (iii) representative meta-microstates can be extracted from the individual studies. We hope that this tool will be useful in coming to a more comprehensive, objective, and overarching representation of microstate findings.

MICROSTATELAB: The EEGLAB Toolbox for Resting-State Microstate Analysis.

Microstate analysis is a multivariate method that enables investigations of the temporal dynamics of large-scale neural networks in EEG recordings of human brain activity. To meet the enormously increasing interest in this approach, we provide a thoroughly updated version of the first open source EEGLAB toolbox for the standardized identification, visualization, and quantification of microstates in resting-state EEG data. The toolbox allows scientists to (i) identify individual, mean, and grand mean microstate maps using topographical clustering approaches, (ii) check data quality and detect outlier maps, (iii) visualize, sort, and label individual, mean, and grand mean microstate maps according to published maps, (iv) compare topographical similarities of group and grand mean microstate maps and quantify shared variances, (v) obtain the temporal dynamics of the microstate classes in individual EEGs, (vi) export quantifications of these temporal dynamics of the microstates for statistical tests, and finally, (vii) test for topographical differences between groups and conditions using topographic analysis of variance (TANOVA). Here, we introduce the toolbox in a step-by-step tutorial, using a sample dataset of 34 resting-state EEG recordings that are publicly available to follow along with this tutorial. The goals of this manuscript are (a) to provide a standardized, freely available toolbox for resting-state microstate analysis to the scientific community, (b) to allow researchers to use best practices for microstate analysis by following a step-by-step tutorial, and (c) to improve the methodological standards of microstate research by providing previously unavailable functions and recommendations on critical decisions required in microstate analyses.

EEG Microstates in Social and Affective Neuroscience.

Social interactions require both the rapid processing of multifaceted socio-affective signals (e.g., eye gaze, facial expressions, gestures) and their integration with evaluations, social knowledge, and expectations. Researchers interested in understanding complex social cognition and behavior face a "black box" problem: What are the underlying mental processes rapidly occurring between perception and action and why are there such vast individual differences? In this review, we promote electroencephalography (EEG) microstates as a powerful tool for both examining socio-affective states (e.g., processing whether someone is in need in a given situation) and identifying the sources of heterogeneity in socio-affective traits (e.g., general willingness to help others). EEG microstates are identified by analyzing scalp field maps (i.e., the distribution of the electrical field on the scalp) over time. This data-driven, reference-independent approach allows for identifying, timing, sequencing, and quantifying the activation of large-scale brain networks relevant to our socio-affective mind. In light of these benefits, EEG microstates should become an indispensable part of the methodological toolkit of laboratories working in the field of social and affective neuroscience.

A Self-Controlled Mind Is Reflected by Stable Mental Processing.

Self-control-the ability to inhibit inappropriate impulses-predicts economic, physical, and psychological well-being. However, recent findings demonstrate low correlations among self-control measures, raising the question of what self-control actually is. Here, we examined the idea that people high in self-control show more stable mental processing, characterized by processing steps that are fewer in number but longer lasting because of fewer interruptions by distracting impulses. To test this hypothesis, we relied on resting electroencephalography microstate analysis, a method that provides access to the stream of mental processing by assessing the sequential activation of neural networks. Across two samples (Study 1: N = 58 male adults from Germany; Study 2: N = 101 adults from Canada, 58 females), the temporal stability of resting networks (i.e., longer durations and fewer occurrences) was positively associated with self-reported self-control and a neural index of inhibitory control, and it was negatively associated with risk-taking behavior. These findings suggest that stable mental processing represents a core feature of a self-controlled mind.

Acute alcohol intoxication modulates the temporal dynamics of resting electroencephalography networks.

This study aimed to provide a currently missing link between general intoxication-induced changes in overall brain activity and the multiple cognitive control deficits typically observed during acute alcohol intoxication. For that purpose, we analyzed the effects of acute alcohol intoxication (1.1‰) on the four archetypal electroencephalography (EEG) resting networks (i.e., microstates A-D) and their temporal dynamics (e.g., coverage and transitions from one microstate to another), as well as on self-reported resting-state cognition in n = 22 healthy young males using a counterbalanced within-subject design. Our microstate analyses indicated that alcohol increased the coverage of the visual processing-related microstate B at the expense of the autonomic processing-related microstate C. Add-on exploratory analyses revealed that alcohol increased transitions from microstate C to microstate B and decreased bidirectional transitions between microstate C and the attention-related microstate D. In line with the observed alcohol-induced decrease of the autonomic processing-related microstate C, participants reported decreases of their somatic awareness during intoxication, which were positively associated with more transitions from microstate C to microstate B. In sum, the observed effects provide mechanistic insights into how alcohol might hamper cognitive processing by generally prioritizing the bottom-up processing of visual stimuli over top-down internal information processing. The fact that this was found during the resting state further proves that alcohol-induced changes in brain activity are continuously present and do not only emerge during demanding situations or tasks.

Clocking the social mind by identifying mental processes in the IAT with electrical neuroimaging.

Why do people take longer to associate the word "love" with outgroup words (incongruent condition) than with ingroup words (congruent condition)? Despite the widespread use of the implicit association test (IAT), it has remained unclear whether this IAT effect is due to additional mental processes in the incongruent condition, or due to longer duration of the same processes. Here, we addressed this previously insoluble issue by assessing the spatiotemporal evolution of brain electrical activity in 83 participants. From stimulus presentation until response production, we identified seven processes. Crucially, all seven processes occurred in the same temporal sequence in both conditions, but participants needed more time to perform one early occurring process (perceptual processing) and one late occurring process (implementing cognitive control to select the motor response) in the incongruent compared with the congruent condition. We also found that the latter process contributed to individual differences in implicit bias. These results advance understanding of the neural mechanics of response time differences in the IAT: They speak against theories that explain the IAT effect as due to additional processes in the incongruent condition and speak in favor of theories that assume a longer duration of specific processes in the incongruent condition. More broadly, our data analysis approach illustrates the potential of electrical neuroimaging to illuminate the temporal organization of mental processes involved in social cognition.

Individual differences in inhibitory control--relationship between baseline activation in lateral PFC and an electrophysiological index of response inhibition.

The capacity to inhibit inappropriate responses is crucial for goal-directed behavior. Inhibiting such responses seems to come more easily to some of us than others, however. From where do these individual differences originate? Here, we measured 263 participants' neural baseline activation using resting electroencephalogram. Then, we used this stable neural marker to predict a reliable electrophysiological index of response inhibition capacity in the cued Continuous Performance Test, the NoGo-Anteriorization (NGA). Using a source-localization technique, we found that resting delta, theta, and alpha1 activity in the left middle frontal gyrus and resting alpha1 activity in the right inferior frontal gyrus were negatively correlated with the NGA. As a larger NGA is thought to represent better response inhibition capacity, our findings demonstrate that lower levels of resting slow-wave oscillations in the lateral prefrontal cortex, bilaterally, are associated with a better response inhibition capacity.

Normalized affective responsiveness following deep brain stimulation of the medial forebrain bundle in depression.

Deep brain stimulation (DBS) of the supero-lateral medial forebrain bundle (slMFB) is associated with rapid and sustained antidepressant effects in treatment-resistant depression (TRD). Beyond that, improvements in social functioning have been reported. However, it is unclear whether social skills, the basis of successful social functioning, are systematically altered following slMFB DBS. Therefore, the current study investigated specific social skills (affective empathy, compassion, and theory of mind) in patients with TRD undergoing slMFB DBS in comparison to healthy subjects. 12 patients with TRD and 12 age- and gender-matched healthy subjects (5 females) performed the EmpaToM, a video-based naturalistic paradigm differentiating between affective empathy, compassion, and theory of mind. Patients were assessed before and three months after DBS onset and compared to an age- and gender-matched sample of healthy controls. All data were analyzed using non-parametric Mann-Whitney U tests. DBS treatment significantly affected patients' affective responsiveness towards emotional versus neutral situations (i.e. affective empathy): While their affective responsiveness was reduced compared to healthy subjects at baseline, they showed normalized affective responsiveness three months after slMFB DBS onset. No effects occurred in other domains with persisting deficits in compassion and intact socio-cognitive skills. Active slMFB DBS resulted in a normalized affective responsiveness in patients with TRD. This specific effect might represent one factor supporting the resumption of social activities after recovery from chronic depression. Considering the small size of this unique sample as well as the explorative nature of this study, future studies are needed to investigate the robustness of these effects.

Impartiality in humans is predicted by brain structure of dorsomedial prefrontal cortex.

The moral force of impartiality (i.e. the equal treatment of all human beings) is imperative for providing justice and fairness. Yet, in reality many people become partial during intergroup interactions; they demonstrate a preferential treatment of ingroup members and a discriminatory treatment of outgroup members. Some people, however, do not show this intergroup bias. The underlying sources of these inter-individual differences are poorly understood. Here we demonstrate that the larger the gray matter volume and thickness of the dorsomedial prefrontal cortex (DMPFC), the more individuals in the role of an uninvolved third-party impartially punish outgroup and ingroup perpetrators. Moreover, we show evidence for a possible mechanism that explains the impact of DMPFC's gray matter volume on impartiality, namely perspective-taking. Large gray matter volume of DMPFC seems to facilitate equal perspective-taking of all sides, which in turn leads to impartial behavior. This is the first evidence demonstrating that brain structure of the DMPFC constitutes an important source underlying an individual's propensity for impartiality.

Oxytocin has sex-specific effects on trust and underlying neurophysiological processes.

The neuropeptide oxytocin (OT) regulates mammalian social approach behavior across sexes. Yet most OT studies in humans exclusively investigated men. Here, we studied sex differences in OT's effects on human trust behavior in 144 heterosexual participants (73 women, 71 men). Participants received 24 international units of intranasal OT or placebo treatment and played a trust game in the role of the investor while undergoing electroencephalography. Trustees were represented by photos of the other sex gradually varying in their pre-rated intensities of facial features signaling attractiveness and threat. On a behavioral level, we observed that OT increased trust in men and reduced it in women when trustees showed weak signals of attractiveness and threat. Correspondingly, on the neurophysiological level, we noted that OT intensified the P100 in male participants, but dampened it in female ones. Our findings demonstrate OT's sex- and context-specific effects on social approach behavior and an underlying early visual attention-related brain process. This evidence demonstrates the need to consider psychobiological mechanisms of sexual dimorphism in human OT research.

Social high performers under stress behave more prosocially and detect happy emotions better in a male sample.

Psychosocial stress is increasing in society, impacting our lives in all social domains. However, the conditions under which stress facilitates ("tend-and-befriend") or hinders ("fight-or-flight") social approach remain elusive. We tested whether previous heterogeneous findings might be resolved by accounting for individual differences in social performance under stress. For that purpose, we introduce the novel Trier Social Stress Test (TSST) social performance index that was aggregated across ratings from two independent observers. Moreover, we apply an innovative setup enabling electroencephalographic (EEG) data to be measured inside an electrically-shielded cabin during stress, namely the TSST-EEG. Relying on a sample of 59 healthy male participants, we collected behavioral (i.e., sharing resources with others) and cognitive (i.e., detecting facial emotional expressions) approach patterns while participants experienced either acute psychosocial stress (n = 31) or no stress (control condition; n = 28) and while EEG was being recorded. During stress exposure, high-performing participants behaved more prosocially, and differentiated better between happy and neutral emotions on both behavioral and neurophysiological levels (revealed by intensity differences in a N170-like response). Overall, our findings demonstrate the added value of both the novel TSST social performance index and the novel TSST-EEG setup. By showing that high social performance during the TSST is associated with behavioral, cognitive, and neurophysiological approach patterns, our study offers valuable insights into adaptive or maladaptive psychobiological mechanisms in coping with psychosocial stress. Future stress research should address the role of social performance differences during stress in social interaction to better understand the behavioral consequences of psychosocial stress in humans.

Impaired socio-affective, but intact socio-cognitive skills in patients with treatment-resistant, recurrent depression.

BACKGROUND: Social withdrawal is a key symptom of depression. The resulting loss of social reinforcement in turn contributes to chronic, recurrent courses of the disease. However, it is not clear whether depressed patients have less motivation to socially interact, or whether their skills in doing so are impaired. The current study investigates potential skill deficits in patients with treatment-resistant depression (TRD). METHODS: 15 TRD patients and 19 age- and sex-matched healthy controls performed the EmpaToM, a paradigm which includes naturalistic video stimuli of either neutral or emotional valence and which differentiates between socio-affective (affective empathy, compassion) and socio-cognitive (theory of mind) skills. RESULTS: Controlling for the baseline affective state in neutral situations, TRD patients displayed significantly reduced affective empathy towards emotional situations compared to healthy controls. Furthermore, TRD patients were less compassionate in both neutral and emotional situations. In contrast, socio-cognitive skill performances did not differ between patients and healthy controls. LIMITATIONS: Further studies might explore socio-affective and socio-cognitive skills in TRD patients using socio-affective/-cognitive tasks involving face-to-face social interactions. CONCLUSION: Our study revealed a specific socio-affective deficit in TRD patients, while showing intact socio-cognitive skills. Patients were less able to affectively resonate with others (affective empathy) and exhibited generally reduced feelings of compassion. These deficits might interfere with providing and receiving social support. Our study contributes to a better understanding of the underlying causes of social withdrawal and stresses the need to specifically address pervasive socio-affective deficits in psychotherapy of TRD patients.

No evidence that gaze anxiety predicts gaze avoidance behavior during face-to-face social interaction.

Eye contact is an indispensable social signal, yet for some individuals it is also a source of discomfort they fear and avoid. However, it is still unknown whether gaze anxiety actually produces avoidant gaze behavior in naturalistic, face-to-face interactions. Here, we relied on a novel dual eye-tracking setup that allows us to assess interactive gaze behavior. To investigate the effect of gaze anxiety on gaze behavior, we a priori created groups of participants reporting high or low levels of gaze anxiety. These participants (n = 51) then performed a semi-standardized interaction with a previously unknown individual reporting a medium level of gaze anxiety. The gaze behavior of both groups did not differ in either classical one-way, eye-tracking parameters (e.g. unilateral eye gaze), or interactive, two-way ones (e.g. mutual gaze). Furthermore, the subjective ratings of both participants' interaction did not differ between groups. Gaze anxious individuals seem to exhibit normal gaze behavior which does not hamper the perceived quality of interactions in a naturalistic face-to-face setup. Our findings point to the existence of cognitive distortions in gaze anxious individuals whose exterior behavior might be less affected than feared by their interior anxiety.

EmBody/EmFace as a new open tool to assess emotion recognition from body and face expressions.

Nonverbal expressions contribute substantially to social interaction by providing information on another person's intentions and feelings. While emotion recognition from dynamic facial expressions has been widely studied, dynamic body expressions and the interplay of emotion recognition from facial and body expressions have attracted less attention, as suitable diagnostic tools are scarce. Here, we provide validation data on a new open source paradigm enabling the assessment of emotion recognition from both 3D-animated emotional body expressions (Task 1: EmBody) and emotionally corresponding dynamic faces (Task 2: EmFace). Both tasks use visually standardized items depicting three emotional states (angry, happy, neutral), and can be used alone or together. We here demonstrate successful psychometric matching of the EmBody/EmFace items in a sample of 217 healthy subjects with excellent retest reliability and validity (correlations with the Reading-the-Mind-in-the-Eyes-Test and Autism-Spectrum Quotient, no correlations with intelligence, and given factorial validity). Taken together, the EmBody/EmFace is a novel, effective (< 5 min per task), highly standardized and reliably precise tool to sensitively assess and compare emotion recognition from body and face stimuli. The EmBody/EmFace has a wide range of potential applications in affective, cognitive and social neuroscience, and in clinical research studying face- and body-specific emotion recognition in patient populations suffering from social interaction deficits such as autism, schizophrenia, or social anxiety.

Facial threat affects trust more strongly than facial attractiveness in women than it does in men.

Trust is essential in initiating social relationships. Due to the differential evolution of sex hormones as well as the fitness burdens of producing offspring, evaluations of a potential mating partner's trustworthiness likely differ across sexes. Here, we explore unknown sex-specific effects of facial attractiveness and threat on trusting other-sex individuals. Ninety-three participants (singles; 46 women) attracted by the other sex performed an incentivized trust game. They had to decide whether to trust individuals of the other sex represented by a priori-created face stimuli gradually varying in the intensities of both attractiveness and threat. Male and female participants trusted attractive and unthreatening-looking individuals more often. However, whereas male participants' trust behavior was affected equally by attractiveness and threat, female participants' trust behavior was more strongly affected by threat than by attractiveness. This indicates that a partner's high facial attractiveness might compensate for high facial threat in male but not female participants. Our findings suggest that men and women prioritize attractiveness and threat differentially, with women paying relatively more attention to threat cues inversely signaling parental investment than to attractiveness cues signaling reproductive fitness. This difference might be attributable to an evolutionary, biologically sex-specific decision regarding parental investment and reproduction behavior.

Oxytocin changes behavior and spatio-temporal brain dynamics underlying inter-group conflict in humans.

Inter-group conflicts drive human discrimination, mass migration, and violence, but their psychobiological mechanisms remain largely unknown. Here, we investigated whether the neuropeptide oxytocin modulates behavior and spatio-temporal brain dynamics in naturalistic inter-group conflict. Eighty-six male members of natural rival social groups received either oxytocin or placebo intranasally. In a decision-making paradigm involving real monetary stakes, participants could sacrifice their own resources to modulate the monetary gains and losses of in- and out-group members. Oxytocin eliminated the reduction in out-group gains - particularly in individuals with low emotional empathy, whereas those given placebo exhibited this negative social behavior. Our spatio-temporal analysis of event-related potentials elicited by outcome valuation revealed that oxytocin replaced a neurophysiological process associated with the negative valuation of out-group gains via a process associated with positive valuation between 200-500ms after outcome presentation. Oxytocin thus seems to modulate inter-group behavior in humans via a specific alteration of valuation-related brain dynamics.

Temporal dynamics of resting EEG networks are associated with prosociality.

As prosociality is key to facing many of our societies' global challenges (such as fighting a global pandemic), we need to better understand why some individuals are more prosocial than others. The present study takes a neural trait approach, examining whether the temporal dynamics of resting EEG networks are associated with inter-individual differences in prosociality. In two experimental sessions, we collected 55 healthy males' resting EEG, their self-reported prosocial concern and values, and their incentivized prosocial behavior across different reward domains (money, time) and social contexts (collective, individual). By means of EEG microstate analysis we identified the temporal coverage of four canonical resting networks (microstates A, B, C, and D) and their mutual communication in order to examine their association with an aggregated index of prosociality. Participants with a higher coverage of microstate A and more transitions from microstate C to A were more prosocial. Our study demonstrates that temporal dynamics of intrinsic brain networks can be linked to complex social behavior. On the basis of previous findings on links of microstate A with sensory processing, our findings suggest that participants with a tendency to engage in bottom-up processing during rest behave more prosocially than others.

The Trust Game for Couples (TGC): A new standardized paradigm to assess trust in romantic relationships.

Trust between couples is a prerequisite for stable and satisfactory romantic relationships. However, there has been no valid research tool to assess partner-specific trust behavior including costly investments in the trustworthiness of the romantic partner. We here present a comprehensive validation of the newly developed Trust Game for Couples (TGC) by means of various self-report and implicit relationship-related measures. The TGC operationalizes trust by measuring an individual's willingness to invest his or her own financial resources in pro-relationship attitudes of their romantic partner (collected by dichotomous responses to relationship-relevant items, e.g., answering yes to "I am absolutely sure that I love my partner"). Thirty-five healthy couples between 20 and 34 years completed the TGC in an interactive (both partners present), but anonymous setting (no information on the partner's responses revealed). Trust, as measured by the TGC, correlates positively with self-reported trust, satisfaction, and felt closeness in the relationship, but not with general interpersonal trust, confirming both its convergent and discriminant validity. In addition to explicit criteria for construct validity, implicit measures of partner valence and confidence explained variance in the TGC, demonstrating that it constitutes an economical measure of implicit and explicit ingredients of trust between couples. In sum, the TGC provides a novel, specific behavioral tool for a sensitive assessment of trust in dyadic relationships with potential for numerous research fields.

Oxytocin modulates the temporal dynamics of resting EEG networks.

Oxytocin is a key modulator of social interaction, but we possess little knowledge of its underlying effects on neuropsychological processes. We used a spatio-temporal EEG microstates analysis to reveal oxytocin's effects on the temporal dynamics of intrinsically generated activity in neural networks. Given oxytocin's known anxiolytic effects, we hypothesized that it increases the temporal stability of the four archetypal EEG resting networks. Eighty-six male participants had received oxytocin or placebo intranasally before we recorded their resting EEG. As hypothesized, oxytocin globally increased the average duration of the four archetypal resting networks and specifically decreased the occurrence and coverage of an autonomic processing-related network to benefit greater coverage of an attention-related network. Moreover, these neurophysiological changes were more pronounced in participants with high anxiety levels and strong subjectively experienced effects of the oxytocin administration. In sum, our study shows that oxytocin reduces rapid switching among neural resting networks by increasing their temporal stability. Specifically, it seems to reduce the brain's need for preparing the internally-oriented processing of autonomic information, thus enabling the externally-oriented processing of social information. Changes in the temporal dynamics of resting networks might underlie oxytocin's anxiolytic effects - potentially informing innovative psychobiological treatment strategies.