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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., (Crown Copyright © 2018. Published by Elsevier B.V. All rights reserved.)

Previous behavioural studies have shown that humans act more altruistically towards kin. Whether and how knowledge of genetic relatedness translates into differential neurocognitive evaluation of observed social interactions has remained an open question. Here, we investigated how the human brain is engaged when viewing a moral dilemma between genetic vs. non-genetic sisters. During functional magnetic resonance imaging, a movie was shown, depicting refusal of organ donation between two sisters, with subjects guided to believe the sisters were related either genetically or by adoption. Although 90% of the subjects self-reported that genetic relationship was not relevant, their brain activity told a different story. Comparing correlations of brain activity across all subject pairs between the two viewing conditions, we found significantly stronger inter-subject correlations in insula, cingulate, medial and lateral prefrontal, superior temporal, and superior parietal cortices, when the subjects believed that the sisters were genetically related. Cognitive functions previously associated with these areas include moral and emotional conflict regulation, decision making, and mentalizing, suggesting more similar engagement of such functions when observing refusal of altruism from a genetic sister. Our results show that mere knowledge of a genetic relationship between interacting persons robustly modulates social cognition of the perceiver.

The question as to why primates have evolved unusually large brains has received much attention, with many alternative proposals all supported by evidence. We review the main hypotheses, the assumptions they make and the evidence for and against them. Taking as our starting point the fact that every hypothesis has sound empirical evidence to support it, we argue that the hypotheses are best interpreted in terms of a framework of evolutionary causes (selection factors), consequences (evolutionary windows of opportunity) and constraints (usually physiological limitations requiring resolution if large brains are to evolve). Explanations for brain evolution in birds and mammals generally, and primates in particular, have to be seen against the backdrop of the challenges involved with the evolution of coordinated, cohesive, bonded social groups that require novel social behaviours for their resolution, together with the specialized cognition and neural substrates that underpin this. A crucial, but frequently overlooked, issue is that fact that the evolution of large brains required energetic, physiological and time budget constraints to be overcome. In some cases, this was reflected in the evolution of 'smart foraging' and technical intelligence, but in many cases required the evolution of behavioural competences (such as coalition formation) that required novel cognitive skills. These may all have been supported by a domain-general form of cognition that can be used in many different contexts.This article is part of the themed issue 'Physiological determinants of social behaviour in animals'., (© 2017 The Authors.)

In non-human primates, opioid-receptor blockade increases social grooming, and the endogenous opioid system has therefore been hypothesized to support maintenance of long-term relationships in humans as well. Here we tested whether social touch modulates opioidergic activation in humans using in vivo positron emission tomography (PET). Eighteen male participants underwent two PET scans with [11C]carfentanil, a ligand specific to μ-opioid receptors (MOR). During the social touch scan, the participants lay in the scanner while their partners caressed their bodies in a non-sexual fashion. In the baseline scan, participants lay alone in the scanner. Social touch triggered pleasurable sensations and increased MOR availability in the thalamus, striatum, and frontal, cingulate, and insular cortices. Modulation of activity of the opioid system by social touching might provide a neurochemical mechanism reinforcing social bonds between humans., (Copyright © 2016 Elsevier Inc. All rights reserved.)

Sociality is primarily a coordination problem. However, the social (or communication) complexity hypothesis suggests that the kinds of information that can be acquired and processed may limit the size and/or complexity of social groups that a species can maintain. We use an agent-based model to test the hypothesis that the complexity of information processed influences the computational demands involved. We show that successive increases in the kinds of information processed allow organisms to break through the glass ceilings that otherwise limit the size of social groups: larger groups can only be achieved at the cost of more sophisticated kinds of information processing that are disadvantageous when optimal group size is small. These results simultaneously support both the social brain and the social complexity hypotheses.

Psychological studies of relationships tend to focus on specific types of close personal relationships (romantic, parent-offspring, friendship) and examine characteristics of both the individuals and the dyad. This paper looks more broadly at the wider range of relationships that constitute an individual's personal social world. Recent work on the composition of personal social networks suggests that they consist of a series of layers that differ in the quality and quantity of relationships involved. Each layer increases relationship numbers by an approximate multiple of 3 (5-15-50-150) but decreasing levels of intimacy (strong, medium, and weak ties) and frequency of interaction. To account for these regularities, we draw on both social and evolutionary psychology to argue that relationships at different layers serve different functions and have different cost-benefit profiles. At each layer, the benefits are asymptotic but the costs of maintaining a relationship at that level (most obviously, the time that has to be invested in servicing it) are roughly linear with the number of relationships. The trade-off between costs and benefits at a given level, and across the different types of demands and resources typical of different levels, gives rise to a distribution of social effort that generates and maintains a hierarchy of layered sets of relationships within social networks. We suggest that, psychologically, these trade-offs are related to the level of trust in a relationship, and that this is itself a function of the time invested in the relationship., (©2011 The British Psychological Society.)

Evolutionary encephalization, or increasing brain size relative to body size, is assumed to be a general phenomenon in mammals. However, despite extensive evidence for variation in both absolute and relative brain size in extant species, there have been no explicit tests of patterns of brain size change over evolutionary time. Instead, allometric relationships between brain size and body size have been used as a proxy for evolutionary change, despite the validity of this approach being widely questioned. Here we relate brain size to appearance time for 511 fossil and extant mammalian species to test for temporal changes in relative brain size over time. We show that there is wide variation across groups in encephalization slopes across groups and that encephalization is not universal in mammals. We also find that temporal changes in brain size are not associated with allometric relationships between brain and body size. Furthermore, encephalization trends are associated with sociality in extant species. These findings test a major underlying assumption about the pattern and process of mammalian brain evolution and highlight the role sociality may play in driving the evolution of large brains.

Grooming is a widespread activity throughout the animal kingdom, but in primates (including humans) social grooming, or allo-grooming (the grooming of others), plays a particularly important role in social bonding which, in turn, has a major impact on an individual's lifetime reproductive fitness. New evidence from comparative brain analyses suggests that primates have social relationships of a qualitatively different kind to those found in other animal species, and I suggest that, in primates, social grooming has acquired a new function of supporting these. I review the evidence for a neuropeptide basis for social bonding, and draw attention to the fact that the neuroendrocrine pathways involved are quite unresolved. Despite recent claims for the central importance of oxytocin, there is equally good, but invariably ignored, evidence for a role for endorphins. I suggest that these two neuropeptide families may play different roles in the processes of social bonding in primates and non-primates, and that more experimental work will be needed to tease them apart.

The social brain hypothesis argues that large brains have arisen over evolutionary time as a response to the social and ecological conflicts inherent in group living. We test predictions arising from the hypothesis using comparative data from birds and four mammalian orders (Carnivora, Artiodactyla, Chiroptera and Primates) and show that, across all non-primate taxa, relative brain size is principally related to pairbonding, but with enduring stable relationships in primates. We argue that this reflects the cognitive demands of the behavioural coordination and synchrony that is necessary to maintain stable pairbonded relationships. However, primates differ from the other taxa in that they also exhibit a strong effect of group size on brain size. We use data from two behavioural indices of social intensity (enduring bonds between group members and time devoted to social activities) to show that primate relationships differ significantly from those of other taxa. We suggest that, among vertebrates in general, pairbonding represents a qualitative shift from loose aggregations of individuals to complex negotiated relationships, and that these bonded relationships have been generalized to all social partners in only a few taxa (such as anthropoid primates).

The evolution of unusually large brains in some groups of animals, notably primates, has long been a puzzle. Although early explanations tended to emphasize the brain's role in sensory or technical competence (foraging skills, innovations, and way-finding), the balance of evidence now clearly favors the suggestion that it was the computational demands of living in large, complex societies that selected for large brains. However, recent analyses suggest that it may have been the particular demands of the more intense forms of pairbonding that was the critical factor that triggered this evolutionary development. This may explain why primate sociality seems to be so different from that found in most other birds and mammals: Primate sociality is based on bonded relationships of a kind that are found only in pairbonds in other taxa.

Prey use a wide variety of anti-predator defence strategies, including morphological and chemical defences as well as behavioural traits (risk-modulated habitat use, changes in activity patterns, foraging decisions and group living). The critical test of how effective anti-predator strategies are is to relate them to relative indices of mortality across predators. Here, we compare biases in predator diet composition with prey characteristics and show that chimpanzee (Pan troglodytes) and felid show the strongest and the most consistent predator bias towards small-brained prey. We propose that large-brained prey are likely to be more effective at evading predators because they can effectively alter their behavioural responses to specific predator encounters. Thus, we provide evidence for the hypothesis that brain size evolution is potentially driven by selection for more sophisticated and behaviourally flexible anti-predator strategies.

Among mammals, the members of some Orders have relatively large brains. Alternative explanations for this have emphasized either social or ecological selection pressures favouring greater information-processing capacities, including large group size, greater foraging efficiency, higher innovation rates, better invasion success and complex problem solving. However, the focal taxa for these analyses (primates, carnivores and birds) often show both varied ecological competence and social complexity. Here, we focus on the specific relationship between social complexity and brain size in ungulates, a group with relatively simple patterns of resource use, but extremely varied social behaviours. The statistical approach we used, phylogenetic generalized least squares, showed that relative brain size was independently associated with sociality and social complexity as well as with habitat use, while relative neocortex size is associated with social but not ecological factors. A simple index of sociality was a better predictor of both total brain and neocortex size than group size, which may indicate that the cognitive demands of sociality depend on the nature of social relationships as well as the total number of individuals in a group.

Social group size has been shown to correlate with neocortex size in primates. Here we use comparative analyses to show that social group size is independently correlated with the size of non-V1 neocortical areas, but not with other more proximate components of the visual system or with brain systems associated with emotional cueing (e.g. the amygdala). We argue that visual brain components serve as a social information 'input device' for socio-visual stimuli such as facial expressions, bodily gestures and visual status markers, while the non-visual neocortex serves as a 'processing device' whereby these social cues are encoded, interpreted and associated with stored information. However, the second appears to have greater overall importance because the size of the V1 visual area appears to reach an asymptotic size beyond which visual acuity and pattern recognition may not improve significantly. This is especially true of the great ape clade (including humans), that is known to use more sophisticated social cognitive strategies.

The social brain hypothesis predicts that humans have an average of about 150 relationships at any given time. Within this 150, there are layers of friends of an ego, where the number of friends in a layer increases as the emotional closeness decreases. Here we analyse a mobile phone dataset, firstly, to ascertain whether layers of friends can be identified based on call frequency. We then apply different clustering algorithms to break the call frequency of egos into clusters and compare the number of alters in each cluster with the layer size predicted by the social brain hypothesis. In this dataset we find strong evidence for the existence of a layered structure. The clustering yields results that match well with previous studies for the innermost and outermost layers, but for layers in between we observe large variability. [ABSTRACT FROM AUTHOR]

The social brain (or Machiavellian Intelligence) hypothesis was proposed to explain primates' unusually large brains: It argues that the cognitive demands of living in complexly bonded social groups selected for increases in executive brain (principally neocortex). The evidence for this and alternative hypotheses is reviewed. Although there remain difficulties of interpretation, the bulk of the evidence comes down in favor of the social brain hypothesis. The extent to which the cognitive demands of bonding large intensely social groups involve aspects of social cognition, such as theory of mind, is explored. These findings are then related to the evolution of social group size, language, and culture within the hominid lineage. [ABSTRACT FROM AUTHOR]

Striedter's account of human brain evolution fails on two key counts. First, he confuses developmental constraints with selection explanations in the initial jump in hominid brain size around two MYA. Second, he misunderstands the Machiavellian Intelligence explanation. [ABSTRACT FROM AUTHOR]