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Healthy social life requires relationships in different levels of personal closeness. Based on ethological, sociological, and psychological evidence, social networks have been divided into five layers, gradually increasing in size and decreasing in personal closeness. Is this division also reflected in brain processing of social networks? During functional MRI, 21 participants compared their personal closeness to different individuals. We examined the brain volume showing differential activation for varying layers of closeness and found that a disproportionately large portion of this volume (80%) exhibited preference for individuals closest to participants, while separate brain regions showed preference for all other layers. Moreover, this bipartition reflected cortical preference for different sizes of physical spaces, as well as distinct subsystems of the default mode network. Our results support a division of the neurocognitive processing of social networks into two patterns depending on personal closeness, reflecting the unique role intimately close individuals play in our social lives., (© 2024. The Author(s).)

Human communities have self-organizing properties in which specific Dunbar Numbers may be invoked to explain group attachments. By analysing Wikipedia editing histories across a wide range of subject pages, we show that there is an emergent coherence in the size of transient groups formed to edit the content of subject texts, with two peaks averaging at around N = 8 for the size corresponding to maximal contention, and at around N = 4 as a regular team. These values are consistent with the observed sizes of conversational groups, as well as the hierarchical structuring of Dunbar graphs. We use a model of bipartite trust to derive a scaling law that fits the data and may apply to all group size distributions when these are based on attraction to a seeded group process. In addition to providing further evidence that even spontaneous communities of strangers are self-organizing, the results have important implications for the governance of the Wikipedia commons and for the security of all online social platforms and associations., Competing Interests: We declare we have no competing interests., (© 2024 The Author(s).)

Music is widely recognised as a human universal, yet there is no agreed explanation for its function, or why and when it evolved. I summarise experimental evidence that the primary function of musicking lies in social bonding, both at the dyadic and community levels, via the effect that performing any form of music has on the brain's endorphin system (the principal neurohormonal basis for social bonding in primates). The many other functions associated with music-making (mate choice, pleasure, coalition signalling, etc) are all better understood as derivative of this, either as secondary selection pressures or as windows of evolutionary opportunity (exaptations). If music's function is primarily as an adjunct of the social bonding mechanism (a feature it shares with laughter, feasting, storytelling and the rituals of religion), then reverse engineering the problem suggests that the capacity for music-making most likely evolved with the appearance of archaic humans. This agrees well with anatomical evidence for the capacity to sing., Competing Interests: The author declares that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2023 Dunbar.)

The personal network of relationships is structured in circles of friendships, that go from the most intense relationships to the least intense ones. While this is a well established result, little is known about the stability of those circles and their evolution in time. To shed light on this issue, we study the temporal evolution of friendships among teenagers during two consecutive academic years by means of a survey administered on five occasions. We show that the first two circles, best friends and friends, can be clearly observed in the survey but also that being in one or the other leads to more or less stable relationships. We find that being in the same class is one of the key drivers of friendship evolution. We also observe an almost constant degree of reciprocity in the relationships, around 60%, a percentage influenced both by being in the same class and by gender homophily. Not only do our results confirm the mounting evidence supporting the circle structure of human social networks, but they also show that these structures persist in time despite the turnover of individual relationships-a fact that may prove particularly useful for understanding the social environment in middle schools., (© 2023. Springer Nature Limited.)

Doctrinal religions that involve recognised gods, more formal theologies, moral codes, dedicated religious spaces and professional priesthoods emerged in two phases during the Neolithic. Almost all of these appeared in a narrow latitudinal band (the northern Subtropical Zone). I suggest that these developments were the result of a need to facilitate community bonding in response to scalar stresses that developed as community sizes increased dramatically beyond those typical of hunter-gatherer societies. Conditions for population growth (as indexed by rainfall patterns and the difference between pathogen load and the length of the growing season) were uniquely optimised in this zone, creating an environment of ecological release in which populations could grow unusually rapidly. The relationship between latitude, religion and language in contemporary societies suggests that the peculiar characteristics of the northern (but not the southern) Subtropical Zone were especially favourable for the evolution of large scale religions as a way of enforcing community cohesion., Competing Interests: The author declares no conflicts of interest., (© The Author(s) 2023.)

In contrast to long-term relationships, far less is known about the temporal evolution of transient relationships, although these constitute a substantial fraction of people's communication networks. Previous literature suggests that ratings of relationship emotional intensity decay gradually until the relationship ends. Using mobile phone data from three countries (US, UK, and Italy), we demonstrate that the volume of communication between ego and its transient alters does not display such a systematic decay, instead showing a lack of any dominant trends. This means that the communication volume of egos to groups of similar transient alters is stable. We show that alters with longer lifetimes in ego's network receive more calls, with the lifetime of the relationship being predictable from call volume within the first few weeks of first contact. This is observed across all three countries, which include samples of egos at different life stages. The relation between early call volume and lifetime is consistent with the suggestion that individuals initially engage with a new alter so as to evaluate their potential as a tie in terms of homophily., (© 2023. The Author(s).)

Seminal studies suggest that being mimicked increases experienced social closeness and prosocial behavior to a mimicking confederate (i.e., interaction partner). Here we reexamine these results by considering the role of empathy-related traits, an indirect proxy for endorphin uptake, and their combined effects as an explanation for these results. 180 female participants were mimicked or anti-mimicked in an interaction with a confederate. The effects of being mimicked versus anti-mimicked in relation to empathy-related traits and endorphin release (assessed indirectly via pain tolerance) on experienced closeness and prosocial behavior were assessed using Bayesian analyses. Our results suggest that high individual empathy-related traits increase social closeness to the anti-mimicking and mimicking confederate and to one's romantic partner, as compared to mimicry alone. Results furthermore strongly suggest that high individual empathy-related traits increase prosocial behavior (donations and willingness to help) as compared to mimicry alone. These findings extend previous work by highlighting that empathy-related traits are more influential in creating positive effects on social closeness and prosocial behavior than a one-shot mimicking encounter., (© 2023. The Author(s).)

In anthropoid primates, social grooming is the principal mechanism (mediated by the central nervous system endorphin system) that underpins social bonding. However, the time available for social grooming is limited, and this imposes an upper limit on the size of group that can be bonded in this way. I suggest that, when hominins needed to increase the size of their groups beyond the limit that could be bonded by grooming, they co-opted laughter (a modified version of the play vocalization found widely among the catarrhine primates) as a form of chorusing to fill the gap. I show, first, that human laughter both upregulates the brain's endorphin system and increases the sense of bonding between those who laugh together. I then use a reverse engineering approach to model group sizes and grooming time requirements for fossil hominin species to search for pinch points where a phase shift in bonding mechanisms might have occurred. The results suggest that the most likely time for the origin of human-like laughter is the appearance of the genus Homo ca 2.5 Ma. This article is part of the theme issue 'Cracking the laugh code: laughter through the lens of biology, psychology and neuroscience'.

Primates live in stable social groups in which they form differentiated relationships with group members and use a range of communication including facial expressions, vocalizations and gestures. However, how these different types of communication are used to regulate social interactions, and what cognitive skills underpin this communication, is still unclear. The aim of this special issue is to examine the types of cognitive skills underpinning the flexible and complex communication that is used to maintain the bonded social relationships found in primates and humans. This article is part of the theme issue 'Cognition, communication and social bonds in primates'.

Characterizing non-human primate social complexity and its cognitive bases has proved challenging. Using principal component analyses, we show that primate social, ecological and reproductive behaviours condense into two components: socioecological complexity (including most social and ecological variables) and reproductive cooperation (comprising mainly a suite of behaviours associated with pairbonded monogamy). We contextualize these results using a meta-analysis of 44 published analyses of primate brain evolution. These studies yield two main consistent results: cognition, sociality and cooperative behaviours are associated with absolute brain volume, neocortex size and neocortex ratio, whereas diet composition and life history are consistently associated with relative brain size. We use a path analysis to evaluate the causal relationships among these variables, demonstrating that social group size is predicted by the neocortex, whereas ecological traits are predicted by the volume of brain structures other than the neocortex. That a range of social and technical behaviours covary, and are correlated with social group size and brain size, suggests that primate cognition has evolved along a continuum resulting in an increasingly flexible, domain-general capacity to solve a range of socioecological challenges culminating in a capacity for, and reliance on, innovation and social information use in the great apes and humans. This article is part of the theme issue 'Cognition, communication and social bonds in primates'.

Primate social bonds are described as being especially complex in their nature, and primates have unusually large brains for their body size compared to other mammals. Communication in primates has attracted considerable attention because of the important role it plays in social bonding. It has been proposed that differentiated social relationships are cognitively complex because primates need to continuously update their knowledge about different types of social bonds. Therefore, primates infer whether an opportunity for social interaction is rewarding (valuable to individual goals) based on their knowledge of the social relationships of the interactants. However, exposure to distraction and stress has detrimental effects on the dopaminergic system, suggesting that understanding social relationships as rewarding is affected in these conditions. This paper proposes that complex communication evolved to augment the capacity to form social relationships during stress through flexibly modifying intentionality in communication (audience checking, response waiting and elaboration). Intentional communication may upregulate dopamine dynamics to allow recognition that an interaction is rewarding during stress. By examining these associations between complexity of communication and stress, we provide new insights into the cognitive skills involved in forming social bonds in primates and the evolution of communication systems in both primates and humans. This article is part of the theme issue 'Cognition, communication and social bonds in primates'.

Group living is stressful for all mammals, and these stresses limit the size of their social groups. Humans live in very large groups by mammal standards, so how have they solved this problem? I use homicide rates as an index of within-community stress for humans living in small-scale ethnographic societies, and show that the frequency of homicide increases linearly with living-group size in hunter-gatherers. This is not, however, the case for cultivators living in permanent settlements, where there appears to be a 'glass ceiling' below which homicide rates oscillate. This glass ceiling correlates with the adoption of social institutions that allow tensions to be managed. The results suggest (a) that the transition to a settled lifestyle in the Neolithic may have been more challenging than is usually assumed and (b) that the increases in settlement size that followed the first villages necessitated the introduction of a series of social institutions designed to manage within-community discord., Competing Interests: The author declares no conflicts of interest., (© The Author(s) 2022.)

Humans are social animals and the interpersonal bonds formed between them are crucial for their development and well being in a society. These relationships are usually structured into several layers (Dunbar's layers of friendship) depending on their significance in an individual's life with closest friends and family being the most important ones taking major part of their time and communication effort. However, we have little idea how the initiation and termination of these relationships occurs across the lifespan. Mobile phones, in particular, have been used extensively to shed light on the different types of social interactions between individuals and to explore this, we analyse a national cellphone database to determine how and when changes in close relationships occur in the two genders. In general, membership of this inner circle of intimate relationships is extremely stable, at least over a three-year period. However, around 1-4% of alters change every year, with the rate of change being higher among 17-21 year olds than older adults. Young adult females terminate more of their opposite-gender relationships, while older males are more persistent in trying to maintain relationships in decline. These results emphasise the variability in relationship dynamics across age and gender, and remind us that individual differences play an important role in the structure of social networks. Overall, our study provides a holistic understanding of the dynamic nature of close relationships during different stages of human life., (© 2022. The Author(s).)

Professional religious specialists centralised religious authority in early human societies and represented some of the earliest instances of formalised social leadership. These individuals played a central role in the emergence of organised religion and transitions to more stratified human societies. Evolutionary theories highlight a range of environmental, economic and social factors that are potentially causally related to the emergence of professional religious specialists in human history. There remains little consensus over the relative importance of these factors and whether professional religious specialists were the outcome or driver of increased socio-cultural complexity. We built a global dataset of hunter-gatherer societies and developed a novel method of exploratory phylogenetic path analysis. This enabled us to systematically identify the factors associated with the emergence of professional religious specialists and infer the directionality of causal dependencies. We find that environmental predictability, environmental richness, pathogen load, social leadership and food storage systems are all correlated with professional religious specialists. However, only food storage is directly related to the emergence of professional religious specialists. Our findings are most consistent with the claim that the early stages of organised religion were the outcome rather than driver of increased socio-economic complexity., Competing Interests: The authors are not aware of any conflicts of interest., (© The Author(s) 2022.)

Fictional storytelling has played an important role in human cultural life since earliest times, and we are willing to invest significant quantities of time, mental effort and money in it. Nonetheless, the psychological mechanisms that make this possible, and how they relate to the mechanisms that underpin real-world social relationships, remain understudied. We explore three factors: identification (the capacity to identify with a character), moral approval and causal attribution with respect to a character's behaviour in live performances of two plays from the European literary canon. There were significant correlations between the extent to which subjects identified with a character and their moral approval of that character's behaviour that was independent of the way the play was directed. However, the subjects' psychological explanations for a character's behaviour (attribution) were independent of whether or not they identified with, or morally approved of, the character. These data extend previous findings by showing that moral approval plays an important role in facilitating identification even in live drama. Despite being transported by an unfolding drama, audiences do not necessarily become biased in their psychological understanding of why characters behaved as they did. The psychology of drama offers significant insights into the psychological processes that underpin our everyday social world., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Teasdale, Maguire, Budelmann and Dunbar.)

Humans deploy a number of specific behaviours for forming social bonds, one of which is laughter. However, two questions have not yet been investigated with respect to laughter: (1) Does laughter increase the sense of bonding to those with whom we laugh? and (2) Does laughter facilitate prosocial generosity? Using changes in pain threshold as a proxy for endorphin upregulation in the brain and a standard economic game (the Dictator Game) as an assay of prosociality, we show that laughter does trigger the endorphin system and, through that, seems to enhance social bonding, but it does not reliably influence donations to others. This suggests that social bonding and prosociality may operate via different mechanisms, or on different time scales, and relate to different functional objectives., Competing Interests: The authors have declared that no competing interests exist.

Human behaviour follows a 24-h rhythm and is known to be governed by the individual chronotypes. Due to the widespread use of technology in our daily lives, it is possible to record the activities of individuals through their different digital traces. In the present study we utilise a large mobile phone communication dataset containing time stamps of calls and text messages to study the circadian rhythms of anonymous users in a European country. After removing the effect of the synchronization of East-West sun progression with the calling activity, we used two closely related approaches to heuristically compute the chronotypes of the individuals in the dataset, to identify them as morning persons or "larks" and evening persons or "owls". Using the computed chronotypes we showed how the chronotype is largely dependent on age with younger cohorts being more likely to be owls than older cohorts. Moreover, our analysis showed how on average females have distinctly different chronotypes from males. Younger females are more larkish than males while older females are more owlish. Finally, we also studied the period of low calling activity for each of the users which is considered as a marker of their sleep period during the night. We found that while "extreme larks" tend to sleep more than "extreme owls" on the weekends, we do not observe much variation between them on weekdays. In addition, we have observed that women tend to sleep even less than males on weekdays while there is not much difference between them on the weekends., (© 2021. The Author(s).)

The C-tactile (CLTM) peripheral nervous system is involved in social bonding in primates and humans through its capacity to trigger the brain's endorphin system. Since the mammalian cochlea has an unusually high density of similar neurons (type-II spiral ganglion neurons, SGNs), we hypothesise that their function may have been exploited for social bonding by co-opting head movements in response to music and other rhythmic movements of the head in social contexts. Music provides one of many cultural behavioural mechanisms for 'virtual grooming' in that it is used to trigger the endorphin system with many people simultaneously so as to bond both dyadic relationships and large groups. Changes in pain threshold across an activity are a convenient proxy assay for endorphin uptake in the brain, and we use this, in two experiments, to show that pain thresholds are higher when nodding the head than when sitting still., (© 2021. The Author(s).)

Religious rituals are associated with health benefits, potentially produced via social bonding. It is unknown whether secular rituals similarly increase social bonding. We conducted a field study with individuals who celebrate secular rituals at Sunday Assemblies and compared them with participants attending Christian rituals. We assessed levels of social bonding and affect before and after the rituals. Results showed the increase in social bonding taking place in secular rituals is comparable to religious rituals. We also found that both sets of rituals increased positive affect and decreased negative affect, and that the change in positive affect predicted the change in social bonding observed. Together these results suggest that secular rituals might play a similar role to religious ones in fostering feelings of social connection and boosting positive affect., Competing Interests: The authors have declared that no competing interests exist.

Humans survive and thrive through social exchange. Yet, social dependency also comes at a cost. Perceived social isolation, or loneliness, affects physical and mental health, cognitive performance, overall life expectancy, and increases vulnerability to Alzheimer's disease-related dementias. Despite severe consequences on behavior and health, the neural basis of loneliness remains elusive. Using the UK Biobank population imaging-genetics cohort (n = ~40,000, aged 40-69 years when recruited, mean age = 54.9), we test for signatures of loneliness in grey matter morphology, intrinsic functional coupling, and fiber tract microstructure. The loneliness-linked neurobiological profiles converge on a collection of brain regions known as the 'default network'. This higher associative network shows more consistent loneliness associations in grey matter volume than other cortical brain networks. Lonely individuals display stronger functional communication in the default network, and greater microstructural integrity of its fornix pathway. The findings fit with the possibility that the up-regulation of these neural circuits supports mentalizing, reminiscence and imagination to fill the social void.

Religious rituals are universal human practices that play a seminal role in community bonding. In two experiments, we tested the role of mu-opioids as the active factor fostering social bonding. We used a mu-opioid blocker (naltrexone) in two double-blind studies of rituals from different religious traditions. We found the same effect across both studies, with naltrexone leading to significantly lower social bonding compared with placebo. These studies suggest that mu-opioids play a significant role in experiences of social bonding within ritual contexts.

Extensive empirical evidence suggests that there is a maximal number of people with whom an individual can maintain stable social relationships (the Dunbar number). We argue that this arises as a consequence of a natural phase transition in the dynamic self-organization among N individuals within a social system. We present the calculated size dependence of the scaling properties of complex social network models to argue that this collective behavior is an enhanced form of collective intelligence. Direct calculation establishes that the complexity of social networks as measured by their scaling behavior is nonmonotonic, peaking around 150, thereby providing a theoretical basis for the value of the Dunbar number. Thus, we establish a theory-based bridge spanning the gap between sociology and psychology., Competing Interests: The authors declare no competing interest., (Copyright © 2020 the Author(s). Published by PNAS.)

In human and nonhuman primates, sex differences typically explain much interindividual variability. Male and female behaviors may have played unique roles in the likely coevolution of increasing brain volume and more complex social dynamics. To explore possible divergence in social brain morphology between men and women living in different social environments, we applied probabilistic generative modeling to ~10,000 UK Biobank participants. We observed strong volume effects especially in the limbic system but also in regions of the sensory, intermediate, and higher association networks. Sex-specific brain volume effects in the limbic system were linked to the frequency and intensity of social contact, such as indexed by loneliness, household size, and social support. Across the processing hierarchy of neural networks, different conditions for social interplay may resonate in and be influenced by brain anatomy in sex-dependent ways., (Copyright © 2020 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. Distributed under a Creative Commons Attribution NonCommercial License 4.0 (CC BY-NC).)

Most studies of social cognition have focused on dyadic relationships, and rather few have looked at how we engage with individuals in the wider social world into which we are embedded. Here we use principle component analysis (PCA) and path analysis to explore how different aspects of human sociality interact. We demonstrate two distinct clusters in both sexes relating to (i) romantic relationships and (ii) wider social engagement, such as that with the local community. These two domains of relationship were associated with different dispositional traits: individual variation in impulsivity in the former, and in empathy and avoidant attachment in the latter. Although these clusters were broadly similar across both sexes, clearer differentiation is evident in males. In females only, support network size was positively related to the anxious dimension of attachment and, unlike in males, was not related to feelings of inclusion in the local community. This suggests that support networks may play different roles in the two sexes, indicating a productive line of future research. These findings have important practical applications: loneliness interventions that target the specific type of relationship that is felt to be lacking and the associated dispositional traits are likely to be more effective than more generic approaches., Competing Interests: The authors have declared that no competing interests exist.