Structural and Cognitive Mechanisms of Group Cohesion in Primates.

Group-living creates stresses that, all else equal, naturally lead to group fragmentation, and hence loss of the benefits that group-living provides. How species that live in large stable groups counteract these forces is not well understood. I use comparative data on grooming networks and cognitive abilities in primates to show that living in large, stable groups has involved a series of structural solutions designed to create chains of 'friendship' (friends-of-friends effects), increased investment in bonding behaviours (made possible by dietary adjustments) to ensure that coalitions work effectively, and neuronally expensive cognitive skills of the kind known to underpin social relationships in humans. The first ensures that individuals synchronise their activity cycles; the second allows the stresses created by group-living to be defused; and the third allows a large number of weak ties to be managed. Between them, these create a form of multilevel sociality based on strong versus weak ties similar to that found in human social networks. In primates, these strategies appear successively at quite specific group sizes, suggesting that they are solutions to 'glass ceilings' that would otherwise limit the range of group sizes that animals can live in (and hence the habitats they can occupy). This sequence maps closely onto the grades now known to underpin the Social Brain Hypothesis and the fractal pattern that is known to optimise information flow round networks.

Lifehistory Trade-Offs Influence Women's Reproductive Strategies.

Objective: In a UK national census sample, women from the upper and lower socioeconomic (SES) classes achieve parity in completed family size, despite marked differences in both birth rates and offspring survival rates. We test the hypothesis that women adopt reproductive strategies that manipulate age at first reproduction to achieve this. Methods: We use a Monte-Carlo modeling approach parameterized with current UK lifehistory data to simulate the reproductive lifehistories of 64,000 individuals from different SES classes, with parameter values at each successive time step drawn from a statistical distribution defined by the census data. Results: We show that, if they are to achieve parity with women in the higher socioeconomic classes, women in lower socioeconomic classes must begin reproducing 5.65 years earlier on average than women in the higher SES classes in order to offset the higher class-specific mortality and infertility rates that they experience. The model predicts very closely the observed differences in age at first reproduction in the census data. Conclusions: Opting to delay reproduction in order to purse an education-based professional career may be a high risk strategy that many lower SES women are unwilling and unable to pursue. As a result, reproducing as early as possible may be the best strategy available to them.

Why did doctrinal religions first appear in the Northern Subtropical Zone?

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.

Laughter and its role in the evolution of human social bonding.

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'.

Nonverbal Auditory Cues Allow Relationship Quality to be Inferred During Conversations.

The claim that nonverbal cues provide more information than the linguistic content of a conversational exchange (the Mehrabian Conjecture) has been widely cited and equally widely disputed, mainly on methodological grounds. Most studies that have tested the Conjecture have used individual words or short phrases spoken by actors imitating emotions. While cue recognition is certainly important, speech evolved to manage interactions and relationships rather than simple information exchange. In a cross-cultural design, we tested participants' ability to identify the quality of the interaction (rapport) in naturalistic third party conversations in their own and a less familiar language, using full auditory content versus audio clips whose verbal content has been digitally altered to differing extents. We found that, using nonverbal content alone, people are 75-90% as accurate as they are with full audio cues in identifying positive vs negative relationships, and 45-53% as accurate in identifying eight different relationship types. The results broadly support Mehrabian's claim that a significant amount of information about others' social relationships is conveyed in the nonverbal component of speech. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s10919-021-00386-y.

Managing the stresses of group-living in the transition to village life.

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.

How Audiences Engage With Drama: Identification, Attribution and Moral Approval.

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.

Laughter influences social bonding but not prosocial generosity to friends and strangers.

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.

Cochlear SGN neurons elevate pain thresholds in response to music.

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.

Blocking mu-opioid receptors inhibits social bonding in rituals.

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.

Structure and function in human and primate social networks: implications for diffusion, network stability and health.

The human social world is orders of magnitude smaller than our highly urbanized world might lead us to suppose. In addition, human social networks have a very distinct fractal structure similar to that observed in other primates. In part, this reflects a cognitive constraint, and in part a time constraint, on the capacity for interaction. Structured networks of this kind have a significant effect on the rates of transmission of both disease and information. Because the cognitive mechanism underpinning network structure is based on trust, internal and external threats that undermine trust or constrain interaction inevitably result in the fragmentation and restructuring of networks. In contexts where network sizes are smaller, this is likely to have significant impacts on psychological and physical health risks.

Physical Contact and Loneliness: Being Touched Reduces Perceptions of Loneliness.

OBJECTIVE: This article addresses the connection between loneliness and physical contact. Evolutionary and psychological research has shown that touch is an important part of bond-building and emotion communication; loneliness is intimately related to these elements as well. In this paper, we ask whether physical contact reduces feelings of loneliness -which might derive from evolutionary ancient bonding mechanisms-despite a cultural context that is relatively non-tactile. METHOD: An experimental study (40 participants, 13 males) tested for observable effects of touch on loneliness scores in a low-contact culture to analyse whether they respond positively to that stimulus despite cultural training against it. RESULTS: Participants exposed to physical contact reported significantly lower neglect scores from their close relationships in a short loneliness scale, thus suggesting that there is an underlying mechanism that persists despite enculturation. The effects were particularly strong among single people, which could mean that lower loneliness among married people might be partly explained by the regular availability of physical contact. Participants in the experimental condition also showed a faster reduction in heart rate, interpreted as a sign of physiological wellbeing. CONCLUSIONS: These findings help to specify mechanisms within the evolutionary theoretical framework of loneliness that link internal feelings to environmental cues. This article aims at contributing to a more complex discussion on the interactions between emotions, cultural practices and psychological well-being.

Relating size and functionality in human social networks through complexity.

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.

Time Constraints Do Not Limit Group Size in Arboreal Guenons but Do Explain Community Size and Distribution Patterns.

To understand how species will respond to environmental changes, it is important to know how those changes will affect the ecological stress that animals experience. Time constraints can be used as indicators of ecological stress. Here we test whether time constraints can help us understand group sizes, distribution patterns, and community sizes of forest guenons (Cercopithecus/Allochrocebus). Forest guenons typically live in small to medium sized one-male-multifemale groups and often live in communities with multiple forest guenon species. We developed a time-budget model using published data on time budgets, diets, body sizes, climate, and group sizes to predict maximum ecologically tolerable group and community sizes of forest guenons across 202 sub-Saharan African locations. The model correctly predicted presence/absence at 83% of these locations. Feeding-foraging time (an indicator of competition) limited group sizes, while resting and moving time constraints shaped guenon biogeography. Predicted group sizes were greater than observed group sizes but comparable to community sizes, suggesting community sizes are set by competition among guenon individuals irrespective of species. We conclude that time constraints and intraspecific competition are unlikely to be the main determinants of relatively small group sizes in forest guenons. Body mass was negatively correlated with moving time, which may give larger bodied species an advantage over smaller bodied species under future conditions when greater fragmentation of forests is likely to lead to increased moving time. Resting time heavily depended on leaf consumption and is likely to increase under future climatic conditions when leaf quality is expected to decrease.

Trade-off between fertility and predation risk drives a geometric sequence in the pattern of group sizes in baboons.

Group-living offers both benefits (protection against predators, access to resources) and costs (increased ecological competition, the impact of group size on fertility). Here, we use cluster analysis to detect natural patternings in a comprehensive sample of baboon groups, and identify a geometric sequence with peaks at approximately 20, 40, 80 and 160. We suggest (i) that these form a set of demographic oscillators that set habitat-specific limits to group size and (ii) that the oscillator arises from a trade-off between female fertility and predation risk.

Social structure as a strategy to mitigate the costs of group living: a comparison of gelada and guereza monkeys.

In mammals, and especially primates, group size and social complexity are typically correlated. However, we have no general explanation why this is so. I suggest that the answer may lie in one of the costs of group living: mammalian reproductive endocrinology is extremely sensitive to stress, and forms one of the hidden costs of living in groups. Fertility declines with group size widely across the social mammals, including primates, and will ultimately place a constraint on group size. However, some species seem to have been able to mitigate this cost by forming bonded relationships that reduce the impact of experienced aggression, even if rates of aggression remain high. The downside is that they reduce network connectivity and hence risk fragmenting the group by providing fracture lines for group fission. To explore this, I compare network indices and fertility patterns across the same range of group sizes for two species of Old World monkeys, Colobus guereza and Theropithecus gelada: the former relatively unsocial, the latter intensely social with frequent use of grooming-based alliances. Compared to those of the guereza, gelada social networks lose density more slowly, maintain connectedness more effectively and are less likely to fragment as they increase in size. Although fertility declines with group size in both species, in gelada the impact of this effect is deferred to larger group sizes. The differences in fertility and network structure both predict the very different maximum group sizes typical of these two species, as well as the typical sizes at which their groups undergo fission. This finding may explain aspects of wider mammalian sociality.

The structural and functional brain networks that support human social networks.

Social skills rely on a specific set of cognitive processes, raising the possibility that individual differences in social networks are related to differences in specific brain structural and functional networks. Here, we tested this hypothesis with multimodality neuroimaging. With diffusion MRI (DMRI), we showed that differences in structural integrity of particular white matter (WM) tracts, including cingulum bundle, extreme capsule and arcuate fasciculus were associated with an individual's social network size (SNS). A voxel-based morphology analysis demonstrated correlations between gray matter (GM) volume and SNS in limbic and temporal lobe regions. These structural changes co-occured with functional network differences. As a function of SNS, dorsomedial and dorsolateral prefrontal cortex showed altered resting-state functional connectivity with the default mode network (DMN). Finally, we integrated these three complementary methods, interrogating the relationship between social GM clusters and specific WM and resting-state networks (RSNs). Probabilistic tractography seeded in these GM nodes utilized the SNS-related WM pathways. Further, the spatial and functional overlap between the social GM clusters and the DMN was significantly closer than other control RSNs. These integrative analyses provide convergent evidence of the role of specific circuits in SNS, likely supporting the adaptive behavior necessary for success in extensive social environments.

Primate social group sizes exhibit a regular scaling pattern with natural attractors.

Primate groups vary considerably in size across species. Nonetheless, the distribution of mean species group size has a regular scaling pattern with preferred sizes approximating 2.5, 5, 15, 30 and 50 individuals (although strepsirrhines lack the latter two), with a scaling ratio of approximately 2.5 similar to that observed in human social networks. These clusters appear to form distinct social grades that are associated with rapid evolutionary change, presumably in response to intense environmental selection pressures. These findings may have wider implications for other highly social mammal taxa.

The Anatomy of Friendship.

Friendship is the single most important factor influencing our health, well-being, and happiness. Creating and maintaining friendships is, however, extremely costly, in terms of both the time that has to be invested and the cognitive mechanisms that underpin them. Nonetheless, personal social networks exhibit many constancies, notably in their size and their hierarchical structuring. Understanding the processes that give rise to these patterns and their evolutionary origins requires a multidisciplinary approach that combines social and neuropsychology as well as evolutionary biology.