Learning rules of engagement for social exchange within and between groups.

Globalizing economies and long-distance trade rely on individuals from different cultural groups to negotiate agreement on what to give and take. In such settings, individuals often lack insight into what interaction partners deem fair and appropriate, potentially seeding misunderstandings, frustration, and conflict. Here, we examine how individuals decipher distinct rules of engagement and adapt their behavior to reach agreements with partners from other cultural groups. Modeling individuals as Bayesian learners with inequality aversion reveals that individuals, in repeated ultimatum bargaining with responders sampled from different groups, can be more generous than needed. While this allows them to reach agreements, it also gives rise to biased beliefs about what is required to reach agreement with members from distinct groups. Preregistered behavioral (N = 420) and neuroimaging experiments (N = 49) support model predictions: Seeking equitable agreements can lead to overly generous behavior toward partners from different groups alongside incorrect beliefs about prevailing norms of what is appropriate in groups and cultures other than one's own.

Partisans process policy-based and identity-based messages using dissociable neural systems.

Political partisanship is often conceived as a lens through which people view politics. Behavioral research has distinguished two types of "partisan lenses"-policy-based and identity-based-that may influence peoples' perception of political events. Little is known, however, about the mechanisms through which partisan discourse appealing to policy beliefs or targeting partisan identities operate within individuals. We addressed this question by collecting neuroimaging data while participants watched videos of speakers expressing partisan views. A "partisan lens effect" was identified as the difference in neural synchrony between each participant's brain response and that of their partisan ingroup vs. outgroup. When processing policy-based messaging, a partisan lens effect was observed in socio-political reasoning and affective responding brain regions. When processing negative identity-based attacks, a partisan lens effect was observed in mentalizing and affective responding brain regions. These data suggest that the processing of political discourse that appeals to different forms of partisanship is supported by related but distinguishable neural-and therefore psychological-mechanisms, which may have implications for how we characterize partisanship and ameliorate its deleterious impacts.

The Self-Concept Is Represented in the Medial Prefrontal Cortex in Terms of Self-Importance.

Knowledge about one's personality, the self-concept, shapes human experience. Social cognitive neuroscience has made strides addressing the question of where and how the self is represented in the brain. The answer, however, remains elusive. We conducted two functional magnetic resonance imaging experiments (the second preregistered) with human male and female participants employing a self-reference task with a broad range of attributes and carrying out a searchlight representational similarity analysis (RSA). The importance of attributes to self-identity was represented in the medial prefrontal cortex (mPFC), whereas mPFC activation was unrelated both to self-descriptiveness of attributes (experiments 1 and 2) and importance of attributes to a friend's self-identity (experiment 2). Our research provides a comprehensive answer to the abovementioned question: The self-concept is conceptualized in terms of self-importance and represented in the mPFC.SIGNIFICANCE STATEMENT The self-concept comprises beliefs about who one is as an individual (e.g., personality traits, physical characteristics, desires, likes/dislikes, and social roles). Despite researchers' efforts in the last two decades to understand where and how the self-concept is stored in the brain, the question remains elusive. Using a neuroimaging technique, we found that a brain region called medial prefrontal cortex (mPFC) shows differential but systematic activation patterns depending on the importance of presented word stimuli to a participant's self-concept. Our findings suggest that one's sense of the self is supported by neural populations in the mPFC, each of which is differently sensitive to distinct levels of the personal importance of incoming information.

Virtual (Zoom) Interactions Alter Conversational Behavior and Interbrain Coherence.

A growing number of social interactions are taking place virtually on videoconferencing platforms. Here, we explore potential effects of virtual interactions on observed behavior, subjective experience, and neural "single-brain" and "interbrain" activity via functional near-infrared spectroscopy neuroimaging. We scanned a total of 36 human dyads (72 participants, 36 males, 36 females) who engaged in three naturalistic tasks (i.e., problem-solving, creative-innovation, socio-emotional task) in either an in-person or virtual (Zoom) condition. We also coded cooperative behavior from audio recordings. We observed reduced conversational turn-taking behavior during the virtual condition. Given that conversational turn-taking was associated with other metrics of positive social interaction (e.g., subjective cooperation and task performance), this measure may be an indicator of prosocial interaction. In addition, we observed altered patterns of averaged and dynamic interbrain coherence in virtual interactions. Interbrain coherence patterns that were characteristic of the virtual condition were associated with reduced conversational turn-taking. These insights can inform the design and engineering of the next generation of videoconferencing technology.SIGNIFICANCE STATEMENT Videoconferencing has become an integral part of our lives. Whether this technology impacts behavior and neurobiology is not well understood. We explored potential effects of virtual interaction on social behavior, brain activity, and interbrain coupling. We found that virtual interactions were characterized by patterns of interbrain coupling that were negatively implicated in cooperation. Our findings are consistent with the perspective that videoconferencing technology adversely affects individuals and dyads during social interaction. As virtual interactions become even more necessary, improving the design of videoconferencing technology will be crucial for supporting effective communication.

Sisterhood predicts similar neural processing of a film.

Relationships between humans are essential for how we see the world. Using fMRI, we explored the neural basis of homophily, a sociological concept that describes the tendency to bond with similar others. Our comparison of brain activity between sisters, friends and acquaintances while they watched a movie, indicate that sisters' brain activity is more similar than that of friends and friends' activity is more similar than that of acquaintances. The increased similarity in brain activity measured as inter-subject correlation (ISC) was found both in higher-order brain areas including the default-mode network (DMN) and sensory areas. Increased ISC could not be explained by genetic relation between sisters neither by similarities in eye-movements, emotional experiences, and physiological activity. Our findings shed light on the neural basis of homophily by revealing that similarity in brain activity in the DMN and sensory areas is the stronger the closer is the relationship between the people.

Brain System for Social Categorization by Narrative Roles.

The cognitive system applies categorical thinking to facilitate perception of the rich environment around us. In person cognition, research has focused on the roles of gender, race, age, or appearance in social categorical thinking. Here we investigated how narrative roles, as portrayed by different cinematic characters, are categorized in the neurocognitive system. Under functional MRI, 17 human participants (7 females) were asked to make different judgments regarding personality traits of 16 renowned cinematic characters representing four roles: hero, sidekick, mentor, and villain. Classification analysis showed a brain network, comprising the dorsal medial prefrontal cortex, the precuneus and the temporoparietal junction bilaterally, and the left occipital face area (OFA), to discriminate among the four roles. No such classification was found between other individual attributes including age or the associated film. Moreover, regions overlapping the default mode network (DMN) were found to better discriminate between roles, rather than the individual characters, while the OFA was found to better discriminate between individuals. These results demonstrate the inherent role of roles in person cognition, and suggest an intimate relation between roles categorization and self-referential activity.SIGNIFICANCE STATEMENT Social categorization, the assignment of different people in our social network to subgroups, is a powerful strategy in social cognition. How is this managed by the brain? We provide evidence that different characters from different stories, representing similar roles in their corresponding narrative, elicit similar brain activation patterns, as revealed by functional MRI. Unlike previous studies of social categorization, these brain activations were similar to those elicited by social cognition rather than face processing, and included regions at the prefrontal cortex, the precuneus, and the temporoparietal junction. The identified brain network significantly overlapped the default mode network. We suggest that social categorization by roles is fundamental to the cognitive system, relying on brain regions related to social cognition.

Quantification of inter-brain coupling: A review of current methods used in haemodynamic and electrophysiological hyperscanning studies.

Hyperscanning is a form of neuroimaging experiment where the brains of two or more participants are imaged simultaneously whilst they interact. Within the domain of social neuroscience, hyperscanning is increasingly used to measure inter-brain coupling (IBC) and explore how brain responses change in tandem during social interaction. In addition to cognitive research, some have suggested that quantification of the interplay between interacting participants can be used as a biomarker for a variety of cognitive mechanisms aswell as to investigate mental health and developmental conditions including schizophrenia, social anxiety and autism. However, many different methods have been used to quantify brain coupling and this can lead to questions about comparability across studies and reduce research reproducibility. Here, we review methods for quantifying IBC, and suggest some ways moving forward. Following the PRISMA guidelines, we reviewed 215 hyperscanning studies, across four different brain imaging modalities: functional near-infrared spectroscopy (fNIRS), functional magnetic resonance (fMRI), electroencephalography (EEG) and magnetoencephalography (MEG). Overall, the review identified a total of 27 different methods used to compute IBC. The most common hyperscanning modality is fNIRS, used by 119 studies, 89 of which adopted wavelet coherence. Based on the results of this literature survey, we first report summary statistics of the hyperscanning field, followed by a brief overview of each signal that is obtained from each neuroimaging modality used in hyperscanning. We then discuss the rationale, assumptions and suitability of each method to different modalities which can be used to investigate IBC. Finally, we discuss issues surrounding the interpretation of each method.

Reading the mind in the eyes of Black and White people: Interracial contact and perceived race affects brain activity when inferring mental states.

Inferring others' mental states, or mentalizing, is a critical social cognitive ability that underlies humans' remarkable capacity for complex social interactions. Recent work suggests that interracial contact shapes the recruitment of brain regions involved in mentalizing during impression formation. However, it remains unclear how a target's perceived racial group and a perceiver's previous contact with that racial group shapes mental state inferences. In this study, we examined brain activity in regions of interest associated with mentalizing and race perception among self-identified White perceivers who varied in lifetime contact while they inferred secondary emotions from perceived White eyes and perceived Black eyes (i.e., the Reading the Mind in the Eyes test). The interaction between lifetime contact and perceived target race predicted activity in the superior temporal sulcus (STS), a region consistently implicated in mental state inferences from perceptual cues, tracking eye gaze, and biological motion. Low and average contact White perceivers showed more left STS activity when inferring mental states from perceived White eyes than perceived Black eyes, whereas high contact White perceivers showed similar left STS activity regardless of perceived target race. These results indicate that interracial contact decreases racial biases in the recruitment of regions involved in mentalizing when inferring mental states from perceptual cues.

Training volitional control of the theory of mind network with real-time fMRI neurofeedback.

Is there a way improve our ability to understand the minds of others? Towards addressing this question, here, we conducted a single-arm, proof-of-concept study to evaluate whether real-time fMRI neurofeedback (rtfMRI-NF) from the temporo-parietal junction (TPJ) leads to volitional control of the neural network subserving theory of mind (ToM; the process by which we attribute and reason about the mental states of others). As additional aims, we evaluated the strategies used to self-regulate the network and whether volitional control of the ToM network was moderated by participant characteristics and associated with improved performance on behavioral measures. Sixteen participants underwent fMRI while completing a task designed to individually-localize the TPJ, and then three separate rtfMRI-NF scans during which they completed multiple runs of a training task while receiving intermittent, activation-based feedback from the TPJ, and one run of a transfer task in which no neurofeedback was provided. Region-of-interest analyses demonstrated volitional control in most regions during the training tasks and during the transfer task, although the effects were smaller in magnitude and not observed in one of the neurofeedback targets for the transfer task. Text analysis demonstrated that volitional control was most strongly associated with thinking about prior social experiences when up-regulating the neural signal. Analysis of behavioral performance and brain-behavior associations largely did not reveal behavior changes except for a positive association between volitional control in RTPJ and changes in performance on one ToM task. Exploratory analysis suggested neurofeedback-related learning occurred, although some degree of volitional control appeared to be conferred with the initial self-regulation strategy provided to participants (i.e., without the neurofeedback signal). Critical study limitations include the lack of a control group and pre-rtfMRI transfer scan, which prevents a more direct assessment of neurofeedback-induced volitional control, and a small sample size, which may have led to an overestimate and/or unreliable estimate of study effects. Nonetheless, together, this study demonstrates the feasibility of training volitional control of a social cognitive brain network, which may have important clinical applications. Given the study's limitations, findings from this study should be replicated with more robust experimental designs.

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.

Social Relationship Strength Modulates the Similarity of Brain-to-Brain Representations of Group Members.

Within our societies, humans form co-operative groups with diverse levels of relationship quality among individual group members. In establishing relationships with others, we use attitudes and beliefs about group members and the group as a whole to establish relationships with particular members of our social networks. However, we have yet to understand how brain responses to group members facilitate relationship quality between pairs of individuals. We address this here using a round-robin interpersonal perception paradigm in which each participant was both a perceiver and target for every other member of their group in a set of 20 unique groups of between 5 and 6 members in each (total N = 111). Using functional magnetic resonance imaging, we show that measures of social relationship strength modulate the brain-to-brain multivoxel similarity patterns between pairs of participants' responses when perceiving other members of their group in regions of the brain implicated in social cognition. These results provide evidence for a brain mechanism of social cognitive processes serving interpersonal relationship strength among group members.

The Anatomy of Friendship: Neuroanatomic Homophily of the Social Brain among Classroom Friends.

Homophily refers to the tendency to like similar others. Here, we ask if homophily extends to brain structure. Specifically: do children who like one another have more similar brain structures? We hypothesized that neuroanatomic similarity tied to friendship is most likely to pertain to brain regions that support social cognition. To test this hypothesis, we analyzed friendship network data from 1186 children in 49 classrooms. Within each classroom, we identified "friendship distance"-mutual friends, friends-of-friends, and more distantly connected or unconnected children. In total, 125 children (mean age = 7.57 years, 65 females) also had good quality neuroanatomic magnetic resonance imaging scans from which we extracted properties of the "social brain." We found that similarity of the social brain varied by friendship distance: mutual friends showed greater similarity in social brain networks compared with friends-of-friends (ß = 0.65, t = 2.03, P = 0.045) and even more remotely connected peers (ß = 0.77, t = 2.83, P = 0.006); friends-of-friends did not differ from more distantly connected peers (ß = -0.13, t = -0.53, P = 0.6). We report that mutual friends have similar "social brain" networks, adding a neuroanatomic dimension to the adage that "birds of a feather flock together."

The neurobiology of hatred: Tools of Dialogue© intervention for youth reared amidst intractable conflict impacts brain, behaviour, and peacebuilding attitudes.

Myths, drama, and sacred texts have warned against the fragile nature of human love; the closer the affiliative bond, the quicker it can turn into hatred, suggesting similarities in the neurobiological underpinnings of love and hatred. Here, I offer a theoretical account on the neurobiology of hatred based on our model on the biology of human attachments and its three foundations; the oxytocin system, the "affiliative brain", comprising the neural network sustaining attachment, and biobehavioural synchrony, the process by which humans create a coupled biology through coordinated action. These systems mature in mammals in the context of the mother-infant bond and then transfer to support life within social groups. During this transition, they partition to support affiliation and solidarity to one's group and fear and hatred towards out-group based on minor variations in social behaviour. I present the Tools of Dialogue© intervention for outgroup members based on social synchrony. Applied to Israeli and Palestinian youth and implementing RCT, we measured social behaviour, attitudes, hormones, and social brain response before and after the 8-session intervention. Youth receiving the intervention increased reciprocity and reduced hostile behaviour towards outgroup, attenuated the neural marker of prejudice and increased neural empathic response, reduced cortisol and elevated oxytocin, and adapted attitudes of compromise. These neural changes predicted peacebuilding support 7 years later, when young adults can engage in civil responsibilities. Our intervention, the first to show long-term effects of inter-group intervention on brain and behaviour, demonstrates how social synchrony can tilt the neurobiology of hatred towards the pole of affiliation.

Charting the social neuroscience of human attachment (SoNeAt).

This introduction aims to set out the potential as well as some of the pitfalls of the newly emerging area of the Social Neuroscience of Human Attachment (SoNeAt). To organize and interconnect the burgeoning empirical studies in this line of research, including those in this special issue, we outline a programmatic framework including an extension of our conceptual proposals NAMA and NAMDA to guide future research. We hope that this special issue will act as a stimulus for redoubling our efforts advancing the newly emerging SoNeAt area bridging attachment theory and social neuroscience.

Adult attachment is related to maternal neural response to infant cues: an ERP study.

Maternal attachment security is an important predictor of caregiving . However, little is known regarding the neurobiological mechanisms by which attachment influences processing of infant cues, a critical component of caregiving. We examined whether attachment security, measured by the Adult Attachment Interview, might relate to neural responses to infant cues using event-related potentials. Secure (n=35) and insecure (n=24) mothers viewed photographs of infant faces and heard recordings of infant vocalizations while electroencephalography was recorded. We examined initial processing of infant faces (N170) and cries (N100), and attentional allocation to infant faces and cries (P300). Secure mothers were significantly faster than insecure mothers to orient to infant cries (N100), structurally encode their own infant's face (N170), and attend to infant faces (P300). These differences may elucidate mechanisms underlying how attachment may shape neural processing of infant cues and highlight the use ofsocial neuroscientific approaches in examining clinically relevant aspects of attachment.

Constructing Others' Beliefs from One's Own Using Medial Frontal Cortex.

Many daily choices are based on one's own knowledge. However, when predicting other people's behavior, we need to consider the differences between our knowledge and other people's presumed knowledge. Social agents need a mechanism to use privileged information for their own behavior but exclude it from predictions of others. Using fMRI, we investigated the neural implementation of such social and personal predictions in healthy human volunteers of both sexes by manipulating privileged and shared information. The medial frontal cortex appeared to have an important role in flexibly making decisions using privileged information for oneself or predicting others' behavior. Specifically, we show that ventromedial PFC tracked the state of the world independent of the type of decision (personal, social), whereas dorsomedial regions adjusted their frame of reference to the use of privileged or shared information. Sampling privileged evidence not available to another person also relied on specific interactions between temporoparietal junction area and frontal pole.SIGNIFICANCE STATEMENT What we know about the minds of others and how we use that information is crucial to understanding social interaction. Mentalizing, or reading the minds of others, is argued to be particularly well developed in the human and crucially affected in some disorders. However, the intractable nature of human interactions makes it very difficult to study these processes. Here, we present a way to objectively quantify the information people have about others and to investigate how their brain deals with this information. This shows that people use similar areas in the brain related to nonsocial decision-making when making decisions in social situations and modify this information processing by the knowledge about others use these to modify their information processing according to the knowledge of others.

Perceiving social injustice during arrests of Black and White civilians by White police officers: An fMRI investigation.

From social media to courts of law, recordings of interracial police officer-civilian interactions are now widespread and publicly available. People may be motivated to preferentially understand the dynamics of these interactions when they perceive injustice towards those whose communities experience disproportionate policing relative to others (e.g., non-White racial/ethnic groups). To explore these questions, two studies were conducted (study 1 neuroimaging n = 69 and study 2 behavioral n = 58). The fMRI study examined White participants' neural activity when viewing real-world videos with varying degrees of aggression or conflict of White officers arresting a Black or White civilian. Activity in brain regions supporting social cognition was greater when viewing Black (vs. White) civilians involved in more aggressive police encounters. Additionally, although an independent sample of perceivers rated videos featuring Black and White civilians as similar in overall levels of aggression when civilian race was obscured, participants in the fMRI study (where race was not obscured) rated officers as more aggressive and their use of force as less legitimate when the civilian was Black. In study 2, participants who had not viewed the videos also reported that they believe police are generally more unjustly aggressive towards Black compared with White civilians. These findings inform our understanding of how perceptions of conflict with the potential for injustice shape social cognitive engagement when viewing arrests of Black and White individuals by White police officers.

Prefrontal cortical activation associated with prospective memory while walking around a real-world street environment.

Rostral PFC (area 10) activation is common during prospective memory (PM) tasks. But it is not clear what mental processes these activations index. Three candidate explanations from cognitive neuroscience theory are: (i) monitoring of the environment; (ii) spontaneous intention retrieval; (iii) a combination of the two. These explanations make different predictions about the temporal and spatial patterns of activation that would be seen in rostral PFC in naturalistic settings. Accordingly, we plotted functional events in PFC using portable fNIRS while people were carrying out a PM task outside the lab and responding to cues when they were encountered, to decide between these explanations. Nineteen people were asked to walk around a street in London, U.K. and perform various tasks while also remembering to respond to prospective memory (PM) cues when they detected them. The prospective memory cues could be either social (involving greeting a person) or non-social (interacting with a parking meter) in nature. There were also a number of contrast conditions which allowed us to determine activation specifically related to the prospective memory components of the tasks. We found that maintaining both social and non-social intentions was associated with widespread activation within medial and right hemisphere rostral prefrontal cortex (BA 10), in agreement with numerous previous lab-based fMRI studies of prospective memory. In addition, increased activation was found within lateral prefrontal cortex (BA 45 and 46) when people were maintaining a social intention compared to a non-social one. The data were then subjected to a GLM-based method for automatic identification of functional events (AIDE), and the position of the participants at the time of the activation events were located on a map of the physical space. The results showed that the spatial and temporal distribution of these events was not random, but aggregated around areas in which the participants appeared to retrieve their future intentions (i.e., where they saw intentional cues), as well as where they executed them. Functional events were detected most frequently in BA 10 during the PM conditions compared to other regions and tasks. Mobile fNIRS can be used to measure higher cognitive functions of the prefrontal cortex in "real world" situations outside the laboratory in freely ambulant individuals. The addition of a "brain-first" approach to the data permits the experimenter to determine not only when haemodynamic changes occur, but also where the participant was when it happened. This can be extremely valuable when trying to link brain and cognition.

Technologically-assisted communication attenuates inter-brain synchrony.

The transition to technologically-assisted communication has permeated all facets of human social life; yet, its impact on the social brain is still unknown and the effects may be particularly intense during periods of developmental transitions. Applying a two-brain perspective, the current preregistered study utilized hyperscanning EEG to measure brain-to-brain synchrony in 62 mother-child pairs at the transition to adolescence (child age; M = 12.26, range 10-14) during live face-to-face interaction versus technologically-assisted remote communication. The live interaction elicited 9 significant cross-brain links between densely inter-connected frontal and temporal areas in the beta range [14-30 Hz]. Mother's right frontal region connected with the child's right and left frontal, temporal, and central regions, suggesting its regulatory role in organizing the two-brain dynamics. In contrast, the remote interaction elicited only 1 significant cross-brain-cross-hemisphere link, attenuating the robust right-to-right-brain connectivity during live social moments that communicates socio-affective signals. Furthermore, while the level of social behavior was comparable between the two interactions, brain-behavior associations emerged only during the live exchange. Mother-child right temporal-temporal synchrony linked with moments of shared gaze and the degree of child engagement and empathic behavior correlated with right frontal-frontal synchrony. Our findings indicate that human co-presence is underpinned by specific neurobiological processes that should be studied in depth. Much further research is needed to tease apart whether the "Zoom fatigue" experienced during technological communication may stem, in part, from overload on more limited inter-brain connections and to address the potential cost of social technology for brain maturation, particularly among youth.

The neural computation of human prosocial choices in complex motivational states.

Motives motivate human behavior. Most behaviors are driven by more than one motive, yet it is unclear how different motives interact and how such motive combinations affect the neural computation of the behaviors they drive. To answer this question, we induced two prosocial motives simultaneously (multi-motive condition) and separately (single motive conditions). After the different motive inductions, participants performed the same choice task in which they allocated points in favor of the other person (prosocial choice) or in favor of themselves (egoistic choice). We used fMRI to assess prosocial choice-related brain responses and drift diffusion modeling to specify how motive combinations affect individual components of the choice process. Our results showed that the combination of the two motives in the multi-motive condition increased participants' choice biases prior to the behavior itself. On the neural level, these changes in initial prosocial bias were associated with neural responses in the bilateral dorsal striatum. In contrast, the efficiency of the prosocial decision process was comparable between the multi-motive and the single-motive conditions. These findings provide insights into the computation of prosocial choices in complex motivational states, the motivational setting that drives most human behaviors.