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1.  Processing power limits social group size: computational evidence for the cognitive costs of sociality 
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.
doi:10.1098/rspb.2013.1151
PMCID: PMC3712454  PMID: 23804623
social brain hypothesis; behavioural synchrony; social network size
2.  New insights into differences in brain organization between Neanderthals and anatomically modern humans 
Previous research has identified morphological differences between the brains of Neanderthals and anatomically modern humans (AMHs). However, studies using endocasts or the cranium itself are limited to investigating external surface features and the overall size and shape of the brain. A complementary approach uses comparative primate data to estimate the size of internal brain areas. Previous attempts to do this have generally assumed that identical total brain volumes imply identical internal organization. Here, we argue that, in the case of Neanderthals and AMHs, differences in the size of the body and visual system imply differences in organization between the same-sized brains of these two taxa. We show that Neanderthals had significantly larger visual systems than contemporary AMHs (indexed by orbital volume) and that when this, along with their greater body mass, is taken into account, Neanderthals have significantly smaller adjusted endocranial capacities than contemporary AMHs. We discuss possible implications of differing brain organization in terms of social cognition, and consider these in the context of differing abilities to cope with fluctuating resources and cultural maintenance.
doi:10.1098/rspb.2013.0168
PMCID: PMC3619466  PMID: 23486442
Neanderthals; brain's orbits; body mass; social cognition
3.  Social cognition on the Internet: testing constraints on social network size 
The social brain hypothesis (an explanation for the evolution of brain size in primates) predicts that humans typically cannot maintain more than 150 relationships at any one time. The constraint is partly cognitive (ultimately determined by some aspect of brain volume) and partly one of time. Friendships (but not necessarily kin relationships) are maintained by investing time in them, and failure to do so results in an inexorable deterioration in the quality of a relationship. The Internet, and in particular the rise of social networking sites (SNSs), raises the possibility that digital media might allow us to circumvent some or all of these constraints. This allows us to test the importance of these constraints in limiting human sociality. Although the recency of SNSs means that there have been relatively few studies, those that are available suggest that, in general, the ability to broadcast to many individuals at once, and the possibilities this provides in terms of continuously updating our understanding of network members’ behaviour and thoughts, do not allow larger networks to be maintained. This may be because only relatively weak quality relationships can be maintained without face-to-face interaction.
doi:10.1098/rstb.2012.0121
PMCID: PMC3385686  PMID: 22734062
social brain hypothesis; social networks; Internet; social networking site; relationships
4.  Bridging the bonding gap: the transition from primates to humans 
Primate societies are characterized by bonded social relationships of a kind that are rare in other mammal taxa. These bonded relationships, which provide the basis for coalitions, are underpinned by an endorphin mechanism mediated by social grooming. However, bonded relationships of this kind impose constraints on the size of social groups that are possible. When ecological pressures have demanded larger groups, primates have had to evolve new mechanisms to facilitate bonding. This has involved increasing the size of vocal and visual communication repertoires, increasing the time devoted to social interaction and developing a capacity to manage two-tier social relationships (strong and weak ties). I consider the implications of these constraints for the evolution of human social communities and argue that laughter was an early evolutionary innovation that helped bridge the bonding gap between the group sizes characteristic of chimpanzees and australopithecines and those in later hominins.
doi:10.1098/rstb.2011.0217
PMCID: PMC3367699  PMID: 22641822
social complexity; social network; primates; social bonding; laughter
5.  Orbital prefrontal cortex volume predicts social network size: an imaging study of individual differences in humans 
The social brain hypothesis, an explanation for the unusually large brains of primates, posits that the size of social group typical of a species is directly related to the volume of its neocortex. To test whether this hypothesis also applies at the within-species level, we applied the Cavalieri method of stereology in conjunction with point counting on magnetic resonance images to determine the volume of prefrontal cortex (PFC) subfields, including dorsal and orbital regions. Path analysis in a sample of 40 healthy adult humans revealed a significant linear relationship between orbital (but not dorsal) PFC volume and the size of subjects' social networks that was mediated by individual intentionality (mentalizing) competences. The results support the social brain hypothesis by indicating a relationship between PFC volume and social network size that applies within species, and, more importantly, indicates that the relationship is mediated by social cognitive skills.
doi:10.1098/rspb.2011.2574
PMCID: PMC3321718  PMID: 22298855
MRI; prefrontal cortex; stereology; social network
6.  Social laughter is correlated with an elevated pain threshold 
Although laughter forms an important part of human non-verbal communication, it has received rather less attention than it deserves in both the experimental and the observational literatures. Relaxed social (Duchenne) laughter is associated with feelings of wellbeing and heightened affect, a proximate explanation for which might be the release of endorphins. We tested this hypothesis in a series of six experimental studies in both the laboratory (watching videos) and naturalistic contexts (watching stage performances), using change in pain threshold as an assay for endorphin release. The results show that pain thresholds are significantly higher after laughter than in the control condition. This pain-tolerance effect is due to laughter itself and not simply due to a change in positive affect. We suggest that laughter, through an endorphin-mediated opiate effect, may play a crucial role in social bonding.
doi:10.1098/rspb.2011.1373
PMCID: PMC3267132  PMID: 21920973
laughter; positive affect; pain threshold; endorphins; social bonding
7.  Altruism in networks: the effect of connections 
Biology Letters  2011;7(5):651-653.
Why are individuals altruistic to their friends? Theory suggests that individual, relationship and network factors will all influence the levels of altruism; but to date, the effects of social network structure have received relatively little attention. The present study uses a novel correlational design to test the prediction that an individual will be more altruistic to friends who are well-connected to the individual's other friends. The result shows that, as predicted, even when controlling for a range of individual and relationship factors, the network factor (number of connections) makes a significant contribution to altruism, thus showing that individuals are more likely to be altruistic to better-connected members of their social networks. The implications of incorporating network structure into studies of altruism are discussed.
doi:10.1098/rsbl.2010.1202
PMCID: PMC3169038  PMID: 21411451
altruism; network; friendship; cooperation
8.  Rowers' high: behavioural synchrony is correlated with elevated pain thresholds 
Biology Letters  2009;6(1):106-108.
Physical exercise is known to stimulate the release of endorphins, creating a mild sense of euphoria that has rewarding properties. Using pain tolerance (a conventional non-invasive assay for endorphin release), we show that synchronized training in a college rowing crew creates a heightened endorphin surge compared with a similar training regime carried out alone. This heightened effect from synchronized activity may explain the sense of euphoria experienced during other social activities (such as laughter, music-making and dancing) that are involved in social bonding in humans and possibly other vertebrates.
doi:10.1098/rsbl.2009.0670
PMCID: PMC2817271  PMID: 19755532
endorphins; rowing; synchronized performance; euphoria
9.  Network cohesion, group size and neocortex size in female-bonded Old World primates 
Most primates are intensely social and spend a large amount of time servicing social relationships. In this study, we use social network analysis to examine the relationship between primate group size, total brain size, neocortex ratio and several social network metrics concerned with network cohesion. Using female grooming networks from a number of Old World monkey species, we found that neocortex size was a better predictor of network characteristics than endocranial volumes. We further found that when we controlled for group size, neocortex ratio was negatively correlated with network density, connectivity, relative clan size and proportional clan membership, while there was no effect of neocortex ratio on change in connectivity following the removal of the most central female in the network. Thus, in species with larger neocortex ratios, females generally live in more fragmented networks, belong to smaller grooming clans and are members of relatively fewer clans despite living in a closely bonded group. However, even though groups are more fragmented to begin with among species with larger neocortices, the removal of the most central individual does cause groups to fall apart, suggesting that social complexity may ultimately involve the management of highly fragmented social groups while at the same time maintaining overall social cohesion. These results emphasize a need for more detailed brain data on a wider sample of primate species.
doi:10.1098/rspb.2009.1409
PMCID: PMC2817113  PMID: 19793756
primates; social brain; social cohesion; social complexity; neocortex ratio
10.  The evolution of the social brain: anthropoid primates contrast with other vertebrates 
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).
doi:10.1098/rspb.2007.0693
PMCID: PMC2274976  PMID: 17652066
brain size; pairbonding; mammals; birds; primates
11.  Understanding primate brain evolution 
We present a detailed reanalysis of the comparative brain data for primates, and develop a model using path analysis that seeks to present the coevolution of primate brain (neocortex) and sociality within a broader ecological and life-history framework. We show that body size, basal metabolic rate and life history act as constraints on brain evolution and through this influence the coevolution of neocortex size and group size. However, they do not determine either of these variables, which appear to be locked in a tight coevolutionary system. We show that, within primates, this relationship is specific to the neocortex. Nonetheless, there are important constraints on brain evolution; we use path analysis to show that, in order to evolve a large neocortex, a species must first evolve a large brain to support that neocortex and this in turn requires adjustments in diet (to provide the energy needed) and life history (to allow sufficient time both for brain growth and for ‘software’ programming). We review a wider literature demonstrating a tight coevolutionary relationship between brain size and sociality in a range of mammalian taxa, but emphasize that the social brain hypothesis is not about the relationship between brain/neocortex size and group size per se; rather, it is about social complexity and we adduce evidence to support this. Finally, we consider the wider issue of how mammalian (and primate) brains evolve in order to localize the social effects.
doi:10.1098/rstb.2006.2001
PMCID: PMC2346523  PMID: 17301028
brain evolution; life history; neocortex; primate; social brain hypothesis
12.  Chimpanzee and felid diet composition is influenced by prey brain size 
Biology Letters  2006;2(4):505-508.
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.
doi:10.1098/rsbl.2006.0519
PMCID: PMC1833997  PMID: 17148274
prey choice; tropical forests; predation risks; hunting strategies
13.  Both social and ecological factors predict ungulate brain size 
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.
doi:10.1098/rspb.2005.3283
PMCID: PMC1560022  PMID: 16555789
social brain; behavioural flexibility; cognitive capacity; social complexity
14.  A community-level evaluation of the impact of prey behavioural and ecological characteristics on predator diet composition. 
Although predation avoidance is the most commonly invoked explanation for vertebrate social evolution, there is little evidence that individuals in larger groups experience lower predation rates than those in small groups. We compare the morphological and behavioural traits of mammal prey species in the Taï forest, Ivory Coast, with the diet preferences of three of their non-human predators: leopards, chimpanzees and African crowned eagles. Individual predators show marked differences in their predation rates on prey species of different body sizes, but clear patterns with prey behaviour were apparent only when differences in prey habitat use were incorporated into the analyses. Leopard predation rates are highest for terrestrial species living in smaller groups, whereas eagle predation rates are negatively correlated with group size only among arboreal prey. When prey predation rates are summed over all three predators, terrestrial species incur higher predation rates than arboreal species and, within both categories, predation rates decline with increasing prey group size and decreasing density of groups in the habitat. These results reveal that it is necessary to consider anti-predator strategies in the context of a dynamic behavioural interaction between predators and prey.
PMCID: PMC1691645  PMID: 15209106
15.  Discrete hierarchical organization of social group sizes 
The ‘social brain hypothesis’ for the evolution of large brains in primates has led to evidence for the coevolution of neocortical size and social group sizes, suggesting that there is a cognitive constraint on group size that depends, in some way, on the volume of neural material available for processing and synthesizing information on social relationships. More recently, work on both human and non-human primates has suggested that social groups are often hierarchically structured. We combine data on human grouping patterns in a comprehensive and systematic study. Using fractal analysis, we identify, with high statistical confidence, a discrete hierarchy of group sizes with a preferred scaling ratio close to three: rather than a single or a continuous spectrum of group sizes, humans spontaneously form groups of preferred sizes organized in a geometrical series approximating 3–5, 9–15, 30–45, etc. Such discrete scale invariance could be related to that identified in signatures of herding behaviour in financial markets and might reflect a hierarchical processing of social nearness by human brains.
doi:10.1098/rspb.2004.2970
PMCID: PMC1634986  PMID: 15734699
social brain hypothesis; social group size; log-periodicity; fractal analysis
16.  Female-biased reproductive strategies in a Hungarian Gypsy population 
Hungarian Gypsy populations invest more heavily in daughters than in sons compared to co-resident Hungarians, in conformity with the predictions of the Trivers–Willard hypothesis. These effects are shown for four different measures of parental investment (sex ratio at birth, frequency of abortion, duration of breast-feeding and length of education). Opportunities for hypergamy into the wealthier Hungarian population appears to be one factor causing Gypsies to prefer daughters over sons. We show that differential investment by sex of offspring is directly related to the fitness pay-offs that accrue for each population through both sexes of offspring.
doi:10.1098/rspb.1997.0003
PMCID: PMC1688218
Parental Investment Sex Bias Sex Ratio Trivers–Willard Effect Gypsies

Results 1-16 (16)