Two common and distinct forms of variation in human functional brain networks.

The cortex has a characteristic layout with specialized functional areas forming distributed large-scale networks. However, substantial work shows striking variation in this organization across people, which relates to differences in behavior. While most previous work treats individual differences as linked to boundary shifts between the borders of regions, here we show that cortical 'variants' also occur at a distance from their typical position, forming ectopic intrusions. Both 'border' and 'ectopic' variants are common across individuals, but differ in their location, network associations, properties of subgroups of individuals, activations during tasks, and prediction of behavioral phenotypes. Border variants also track significantly more with shared genetics than ectopic variants, suggesting a closer link between ectopic variants and environmental influences. This work argues that these two dissociable forms of variation-border shifts and ectopic intrusions-must be separately accounted for in the analysis of individual differences in cortical systems across people.

In the face of ambiguity: intrinsic brain organization in development predicts one's bias toward positivity or negativity.

Exacerbated negativity bias, including in responses to ambiguity, represents a common phenotype of internalizing disorders. Individuals differ in their propensity toward positive or negative appraisals of ambiguity. This variability constitutes one's valence bias, a stable construct linked to mental health. Evidence suggests an initial negativity in response to ambiguity that updates via regulatory processes to support a more positive bias. Previous work implicates the amygdala and prefrontal cortex, and regions of the cingulo-opercular system, in this regulatory process. Nonetheless, the neurodevelopmental origins of valence bias remain unclear. The current study tests whether intrinsic brain organization predicts valence bias among 119 children and adolescents (6 to 17 years). Using whole-brain resting-state functional connectivity, a machine-learning model predicted valence bias (r = 0.20, P = 0.03), as did a model restricted to amygdala and cingulo-opercular system features (r = 0.19, P = 0.04). Disrupting connectivity revealed additional intra-system (e.g. fronto-parietal) and inter-system (e.g. amygdala to cingulo-opercular) connectivity important for prediction. The results highlight top-down control systems and bottom-up perceptual processes that influence valence bias in development. Thus, intrinsic brain organization informs the neurodevelopmental origins of valence bias, and directs future work aimed at explicating related internalizing symptomology.

Proposing a model whereby negative valence bias increases the risk for more severe dysphoric posttraumatic stress disorder and depression symptomology.

Experiencing trauma increases risk for posttraumatic stress disorder (PTSD) and depression, and individuals who experience psychopathology after a traumatic event often experience symptoms from both disorders. Because a tendency to view events in a more negative light and a propensity toward threat appraisals are risk factors for both PTSD and depression, negative valence bias-a tendency to appraise emotional ambiguity as having a more negative (less positive) meaning-may be a transdiagnostic risk factor. In other words, we expect individuals with a negative valence bias experience greater PTSD and depression symptoms. We measured valence bias and self-reported PTSD and depression symptoms in a sample of college students in 2021 (n = 287; 72.5% reported experiencing trauma). Although valence bias was not associated with PTSD symptoms as a whole, we found in our exploratory model that more negative bias was associated with greater dysphoria-related PTSD symptoms and greater depression symptoms (indirect effect p = .03). Thus, we propose a model whereby a more negative valence bias contributes to increased susceptibility for maladaptive stress responses, which may be associated with greater likelihood of symptoms of dysphoria-related PTSD and depression. These findings suggest that valence bias represents a transdiagnostic affective risk factor, warranting future research examining the impacts of bias-altering interventions (e.g., mindfulness-based treatments) as a means for managing symptoms in individuals with heightened dysphoria-related PTSD and/or depression symptoms. Additionally, in post hoc analyses it emerged that Latinx participants displayed a more negative valence bias, indicating the need for more research in diverse samples. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Concussion-Related Disruptions to Hub Connectivity in the Default Mode Network Are Related to Symptoms and Cognition.

Concussions present with a myriad of symptomatic and cognitive concerns; however, the relationship between these functional disruptions and the underlying changes in the brain are not yet well understood. Hubs, or brain regions that are connected to many different functional networks, may be specifically disrupted after concussion. Given the implications in concussion research, we quantified hub disruption within the default mode network (DMN) and between the DMN and other brain networks. We collected resting-state functional magnetic resonance imaging data from collegiate student-athletes (n = 44) at three time points: baseline (before beginning their athletic season), acute post-injury (approximately 48h after a diagnosed concussion), and recovery (after starting return-to-play progression, but before returning to contact). We used self-reported symptoms and computerized cognitive assessments collected across similar time points to link these functional connectivity changes to clinical outcomes. Concussion resulted in increased connectivity between regions within the DMN compared with baseline and recovery, and this post-injury connectivity was more positively related to symptoms and more negatively related to visual memory performance compared with baseline and recovery. Further, concussion led to decreased connectivity between DMN hubs and visual network non-hubs relative to baseline and recovery, and this post-injury connectivity was more negatively related to somatic symptoms and more positively related to visual memory performance compared with baseline and recovery. Relationships between functional connectivity, symptoms, and cognition were not significantly different at baseline versus recovery. These results highlight a unique relationship between self-reported symptoms, visual memory performance, and acute functional connectivity changes involving DMN hubs after concussion in athletes. This may provide evidence for a disrupted balance of within- and between-network communication highlighting possible network inefficiencies after concussion. These results aid in our understanding of the pathophysiological disruptions after concussion and inform our understanding of the associations between disruptions in brain connectivity and specific clinical presentations acutely post-injury.

The role of trait reappraisal in response to emotional ambiguity: A systematic review and meta-analysis.

Individuals exhibit a systematic valence bias-a specific form of interpretation bias-in response to emotional ambiguity. Accumulating evidence suggests most people initially respond to emotional ambiguity negatively and differ only in subsequent responses. We hypothesized that trait-level cognitive reappraisal-an emotion regulation strategy involving the reinterpretation of affective meaning of stimuli-might explain individual differences in valence bias. To answer this question, we conducted a random-effects meta-analysis of 14 effect sizes from 13 prior studies (n = 2,086), identified via Google Scholar searches. We excluded studies (a) in languages other than English, (b) from non-peer-reviewed sources, or (c) nonempirical sources. We included studies with (a) the Emotion Regulation Questionnaire, (b) a putative measure of valence bias prior to any study-specific manipulations, and (c) adult human participants (i.e., 17+). Supporting our prediction, we found individuals with higher trait reappraisal exhibited a less negative bias (r = -.18, z = -4.04, p < .001), whereas there was a smaller, opposite effect for trait expressive suppression (r = .10, z = 2.14, p = .03). The effects did, however, vary across tasks with stronger effects observed among studies using the scrambled sentences task compared to the valence bias task. Although trait reappraisal accounted for only a small amount of variance, reappraisal may be one mechanism contributing to variability in response to ambiguity. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Specialized late cingulo-opercular network activation elucidates the mechanisms underlying decisions about ambiguity.

Cortical task control networks, including the cingulo-opercular (CO) network play a key role in decision-making across a variety of functional domains. In particular, the CO network functions in a performance reporting capacity that supports successful task performance, especially in response to errors and ambiguity. In two studies testing the contribution of the CO network to ambiguity processing, we presented a valence bias task in which masked clearly and ambiguously valenced emotional expressions were slowly revealed over several seconds. This slow reveal task design provides a window into the decision-making mechanisms as they unfold over the course of a trial. In the main study, the slow reveal task was administered to 32 young adults in the fMRI environment and BOLD time courses were extracted from regions of interest in three control networks. In a follow-up study, the task was administered to a larger, online sample (n = 81) using a more extended slow reveal design with additional unmasking frames. Positive judgments of surprised faces were uniquely accompanied by slower response times and strong, late activation in the CO network. These results support the initial negativity hypothesis, which posits that the default response to ambiguity is negative and positive judgments are associated with a more effortful controlled process, and additionally suggest that this controlled process is mediated by the CO network. Moreover, ambiguous trials were characterized by a second CO response at the end of the trial, firmly placing CO function late in the decision-making process.

Face coverings differentially alter valence judgments of emotional expressions.

Face masks that prevent disease transmission obscure facial expressions, impairing nonverbal communication. We assessed the impact of lower (masks) and upper (sunglasses) face coverings on emotional valence judgments of clearly valenced (fearful, happy) and ambiguously valenced (surprised) expressions, the latter of which have both positive and negative meaning. Masks, but not sunglasses, impaired judgments of clearly valenced expressions compared to faces without coverings. Drift diffusion models revealed that lower, but not upper, face coverings slowed evidence accumulation and affected differences in non-judgment processes (i.e., stimulus encoding, response execution time) for all expressions. Our results confirm mask-interference effects in nonverbal communication. The findings have implications for nonverbal and intergroup communication, and we propose guidance for implementing strategies to overcome mask-related interference.

Tendency to Share Positive Emotions Buffers Loneliness-Related Negativity in the Context of Shared Adversity.

Loneliness is associated with adverse outcomes, and the COVID-19 pandemic threatened to increase loneliness. How loneliness-related outcomes unfold, though, varies across individuals. Individuals' sense of social connectedness and engagement with others to regulate emotional experiences (interpersonal emotion regulation; IER) may modulate loneliness-related outcomes. Individuals failing to maintain social connectedness and/or regulate emotions may be at heightened risk. We assessed how loneliness, social connectedness, and IER related to valence bias, a tendency to categorize ambiguity as more positive or negative. Loneliness was associated with a more negative valence bias among individuals reporting above average social connectedness but who shared positive emotion less often (z=-3.19, p=.001). These findings suggest that sharing positive emotional experiences may buffer loneliness-related outcomes during shared adverse experiences.

Negative interpretation bias connects to real-world daily affect: A multistudy approach.

Negative interpretation bias, the tendency to appraise ambiguous stimuli as threatening, shapes our emotional lives. Various laboratory tasks, which differ in stimuli features and task procedures, can quantify negative interpretation bias. However, it is unknown whether these tasks globally predict individual differences in real-world negative (NA) and positive (PA) affect. Across two studies, we tested whether different lab-based negative interpretation bias tasks predict daily NA and PA, measured via mobile phone across months. To quantify negative interpretation bias, Study 1 (N = 69) used a verbal, self-referential task whereas Study 2 (N = 110) used a perceptual, emotional image task with faces and scenes. Across tasks, negative interpretation bias was linked to heightened daily NA. However, only negative interpretation bias in response to ambiguous faces was related to decreased daily PA. These results illustrate the ecological validity of negative interpretation bias tasks and highlight converging and unique relationships between distinct tasks and naturalistic emotion. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Think again: the role of reappraisal in reducing negative valence bias.

Stimuli such as surprised faces are ambiguous in that they are associated with both positive and negative outcomes. Interestingly, people differ reliably in whether they evaluate these and other ambiguous stimuli as positive or negative, and we have argued that a positive evaluation relies in part on a biasing of the appraisal processes via reappraisal. To further test this idea, we conducted two studies to evaluate whether increasing the cognitive accessibility of reappraisal through a brief emotion regulation task would lead to an increase in positive evaluations of ambiguity. Supporting this prediction, we demonstrated that cuing reappraisal, but not in three other forms of emotion regulation (Study 1a-d; n = 120), increased positive evaluations of ambiguous faces. In a sign of robustness, we also found that the effect of reappraisal generalised from ambiguous faces to ambiguous scenes (Study 2; n = 34). Collectively, these findings suggest that reappraisal may play a key role in determining responses to ambiguous stimuli. We discuss these findings in the context of affective flexibility, and suggest that valence bias (i.e. the tendency to evaluate ambiguity more positively or negatively) represents a novel approach to measuring implicit emotion regulation.

Surprise as an Emotion: A Response to Ortony.

We write in response to an article published in this journal by Andrew Ortony titled "Are All 'Basic Emotions' Emotions? A Problem for the (Basic) Emotions Construct." The author claimed that "for all its elevated status as a basic emotion, surprise fails to satisfy the minimal requirements that [he] proposed for something to be an emotion, and if it is not an emotion, it cannot possibly be a basic emotion." Although we acknowledge the concerns brought forth by Ortony, we respectfully disagree with his conclusion about surprise. To make a case against the assertion that surprise is valence-free, we summarize an extensive body of work showing that surprise is indeed valenced-in a specific manner (i.e., ambiguously valenced)-and that it meets all of Ortony's criteria for an emotion. In other words, rather than being described as neither positive nor negative, this emotion is either positive or negative. We consider the data with respect to surprise as a basic emotion, and we dispute the definitions of basic emotions as "widely divergent." Future work is needed to continue defining an emotion, and a basic emotion, but we believe this is a worthy effort toward shaping a still evolving field.

Affective flexibility as a developmental building block of cognitive reappraisal: An fMRI study.

Cognitive reappraisal is a form of emotion regulation that involves reinterpreting the meaning of a stimulus, often to downregulate one's negative affect. Reappraisal typically recruits distributed regions of prefrontal and parietal cortex to generate new appraisals and downregulate the emotional response in the amygdala. In the current study, we compared reappraisal ability in an fMRI task with affective flexibility in a sample of children and adolescents (ages 6-17, N = 76). Affective flexibility was defined as variability in valence interpretations of ambiguous (surprised) facial expressions from a second behavioral task. Results demonstrated that age and affective flexibility predicted reappraisal ability, with an interaction indicating that flexibility in children (but not adolescents) supports reappraisal success. Using a region of interest-based analysis of participants' BOLD time courses, we also found dissociable reappraisal-related brain mechanisms that support reappraisal success and affective flexibility. Specifically, late increases in middle prefrontal cortex activity supported reappraisal success and late decreases in amygdala activity supported flexibility. Together, these results suggest that our novel measure of affective flexibility - the ability to see multiple interpretations of an ambiguous emotional cue - may represent part of the developmental building blocks of cognitive reappraisal ability.

Task-irrelevant emotional faces impact BOLD responses more for prosaccades than antisaccades in a mixed saccade fMRI task.

Cognitive control allows individuals to flexibly and efficiently perform tasks by attending to relevant stimuli while inhibiting distraction from irrelevant stimuli. The antisaccade task assesses cognitive control by requiring participants to inhibit a prepotent glance towards a peripheral stimulus and generate an eye movement to the mirror image location. This task can be administered with various contextual manipulations to investigate how factors such as trial timing or emotional content interact with cognitive control. In the current study, 26 healthy adults completed a mixed antisaccade and prosaccade fMRI task that included task irrelevant emotional faces and gap/overlap timing. The results showed typical antisaccade and gap behavioral effects with greater BOLD activation in frontal and parietal brain regions for antisaccade and overlap trials. Conversely, there were no differences in behavior based on the emotion of the task irrelevant face, but trials with neutral faces had greater activation in widespread visual regions than trials with angry faces, particularly for prosaccade and overlap trials. Together, these effects suggest that a high level of cognitive control and inhibition was required throughout the task, minimizing the impact of the face presentation on saccade behavior, but leading to increased attention to the neutral faces on overlap prosaccade trials when both the task cue (look towards) and emotion stimulus (neutral, non-threatening) facilitated disinhibition of visual processing.

Interpersonal emotion regulation mitigates the link between trait neuroticism and a more negative valence bias.

Emerging research suggests that trait neuroticism is associated with enhanced attention to and perception of negative emotional stimuli, increasing the risk for multiple forms of psychopathology including depression and anxiety. However, modifiable factors such as certain forms of emotion regulation have the potential to weaken this association. In a large sample (N = 1252), we investigated the link between neuroticism and valence bias in response to stimuli that have the potential for both positive and negative interpretations and examined the moderating role of interpersonal emotion regulation. Primary tests of hypotheses demonstrated that increased neuroticism was associated with a more negative valence bias in response to ambiguity, but only for individuals who are less likely to rely on interpersonal resources to regulate negative affect. Supplemental analyses suggest that this moderation effect of interpersonal emotion regulation might depend on the nature of the stimuli, and that regulation of positive emotions-not just negative affect-can also contribute to a less negative valence bias. Taken together, results suggest that individuals who are high in neuroticism, but consistently rely on interpersonal relationships to regulate their emotions, are better able to override the bias toward negativity that can occur when appraising ambiguity.

Reappraisal-related downregulation of amygdala BOLD activation occurs only during the late trial window.

During cognitive reappraisal, an individual reinterprets the meaning of an emotional stimulus to regulate the intensity of their emotional response. Prefrontal cortex activity has been found to support reappraisal and is putatively thought to downregulate the amygdala response to these stimuli. The timing of these regulation-related responses during the course of a trial, however, remains poorly understood. In the current fMRI study, participants were instructed to view or reappraise negative images and then rate how negative they felt following each image. The hemodynamic response function was estimated in 11 regions of interest for the entire time course of the trial including image viewing and rating. Notably, within the amygdala there was no evidence of downregulation in the early (picture viewing) window of the trial, only in the late (rating) window, which also correlated with a behavioral measure of reappraisal success. With respect to the prefrontal regions, some (e.g., inferior frontal gyrus) showed reappraisal-related activation in the early window, whereas others (e.g., middle frontal gyrus) showed increased activation primarily in the late window. These results highlight the temporal dynamics of different brain regions during emotion regulation and suggest that the amygdala response to negative images need not be immediately dampened to achieve successful cognitive reappraisal.

Mindfulness-based stress reduction triggers a long-term shift toward more positive appraisals of emotional ambiguity.

Reducing negative impacts of stress, for example through mindfulness training, benefits physical and psychological well-being and is becoming ever more crucial owing to large-scale societal uncertainties (e.g., COVID-19). Whereas extensive research has focused on mindfulness-related reductions in self-reported negativity, essentially no research has targeted task-based behavioral outcomes throughout long-term mindfulness trainings. Responses to emotionally ambiguous signals (e.g., surprised expressions), which might be appraised as either positive or negative, provide a nuanced assessment of one's emotional bias across diverse contexts, offering unique leverage for assessing the effects of mindfulness. Here, we compared the effects of short- and long-term training via Mindfulness-Based Stress Reduction on ratings of faces with a relatively clear (angry, happy) and ambiguous (surprised) valence. Ratings became more positive for ambiguity from the start (Week 1) to end of training (Week 8; p < .001), but there were no short-term effects (from a single class session). This shift toward positivity continued through an additional 8-week follow-up (Week 16; p < .001). Notably, posttraining valence bias (Week 8) was uniquely predicted by the nonreactivity facet of mindfulness (p = .01). Together, mindfulness promotes a relatively long-lasting shift toward positivity bias, which is uniquely supported by reduced emotional reactivity. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Spring Break or Heart Break? Extending Valence Bias to Emotional Words.

Ambiguous stimuli are useful for assessing emotional bias. For example, surprised faces could convey a positive or negative meaning, and the degree to which an individual interprets these expressions as positive or negative represents their "valence bias." Currently, the most well-validated ambiguous stimuli for assessing valence bias include nonverbal signals (faces and scenes), overlooking an inherent ambiguity in verbal signals. This study identified 32 words with dual-valence ambiguity (i.e., relatively high intersubject variability in valence ratings and relatively slow response times) and length-matched clearly valenced words (16 positive, 16 negative). Preregistered analyses demonstrated that the words-based valence bias correlated with the bias for faces, r s (213) = .27, p < .001, and scenes, r s (204) = .46, p < .001. That is, the same people who interpret ambiguous faces/scenes as positive also interpret ambiguous words as positive. These findings provide a novel tool for measuring valence bias and greater generalizability, resulting in a more robust measure of this bias.

Reappraisal-but not Suppression-Tendencies Determine Negativity Bias After Laboratory and Real-World Stress Exposure.

Higher reactivity to stress exposure is associated with an increased tendency to appraise ambiguous stimuli as negative. However, it remains unknown whether tendencies to use emotion regulation strategies-such as cognitive reappraisal, which involves altering the meaning or relevance of affective stimuli-can shape individual differences regarding how stress affects perceptions of ambiguity. Here, we examined whether increased reappraisal use is one factor that can determine whether stress exposure induces increased negativity bias. In Study 1, healthy participants (n = 43) rated the valence of emotionally ambiguous (surprised) faces before and after an acute stress or control manipulation and reported reappraisal habits. Increased negativity ratings were milder for stressed individuals that reported more habitual reappraisal use. In Study 2 (n = 97), we extended this investigation to real-world perceived stress before and during the COVID-19 pandemic. We found that reappraisal tendency moderates the relationship between perceived stress and increased negativity bias. Collectively, these findings suggest that the propensity to reappraise determines negativity bias when evaluating ambiguity under stress. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s42761-021-00059-5.

Positivity effect in aging: evidence for the primacy of positive responses to emotional ambiguity.

Older compared to younger adults show greater amygdala activity to positive emotions, and are more likely to interpret emotionally ambiguous stimuli (e.g., surprised faces) as positive. While some evidence suggests this positivity effect results from a top-down, effortful mechanism, others suggest it may emerge as the default or initial response. The amygdala is a key node in rapid, bottom-up processing and patterns of amygdala activity over time (e.g., habituation) can shed light on the mechanisms underlying the positivity effect. Younger and older adults passively viewed neutral and surprised faces in an MRI. Only in older adults, amygdala habituation was associated with the tendency to interpret surprised faces as positive or negative (valence bias), where a more positive bias was associated with greater habituation. Interestingly, although a positive bias in younger adults was associated with slower responses, consistent with an initial negativity hypothesis in younger adults, older adults showed faster categorizations of positivity. Together, we propose that there may be a switch to a primacy of positivity in aging.