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Encephalization in primates has led to the possibility of complex social navigation structures to
emerge (Lefebre, 2012). The issue of the origin of this disproportionate increase in brain-body size
has received considerable debate. While Dunbar (2009) proposed that the large brain of primates was
facilitated by the increasingly complex social systems, as compared to other vertebrates, other
competing theories, notably ecological and cultural explanations, have emerged. These hypotheses
pose that drivers of brain growth were environmental changes – such as extended territory, dietary
transitions – and respectively, learning of technical innovations (tool use). However, cultural
innovation and social complexity co-vary between primate species in analyses of brain correlates
(Reader & Laland, 2002). This essay will thus compare the more distantly related ecological and
social hypotheses of encephalization and suggest their potential roles in the evolution of sociality.

The ecological factor hypothesis initially emerged from the observation that the brain is an
energetically expensive tissue (Aiello & Wheeler, 1995) and consequently posits that species with
better use of resources had higher adaptive fitness. This is measured by the transition to a fruit-based
(frugivory) or omnivorous diet in nonhuman primates (NHP), whose scattered distribution would
require superior cognitive ability, forced by territory mapping and extensive home range (Lefebre,
2012). However, the opposite mechanism is also possible, which will be explored later.

The social brain hypothesis (SBH), in contrast to other types of social intelligence, emphasizes the
role of cognitive abilities in supporting the cohesion of a group, despite ecological problems (Dunbar &
Schultz, 2017). It proposed that coordination in groups requires higher cognitive capacity, therefore
the correlation of group size with cognitive ability has been central in the argument. As MacLean et al.
(2013) show, social intelligence (unlike non-social) correlates with group size in lemur populations.
However, they tested deception (instead of cooperation) in the social intelligence experiment, by
quantifying when the animal would steal food from two human competitors according to body/head
orientation. Thus, competitive ability is predicted to increase with group size (MacLean et al., 2013).

The main support for both hypotheses comes from research of relative brain size. While multiple
studies showed no correlation between whole brain size (or relative brain size) and group size
(Dunbar, 1992; MacLean et al., 2013), neocortical size is more reliably correlated (Dunbar, 1992).
Testing one ecological variable as well (proportionality of fruit in diet), Dunbar found no correlation
with neocortical size or sociality. However, measures of whole (relative) brain size showed the
opposite pattern in a study by DeCasien et al. (2017), where the frugivorous types showed larger
brains than folivores. They found that increases in social complexity (as measured by group size,
social and mating systems) did not correlate with brain size on any of the measures. One difference
between the studies is the breadth of analysis, with 38 genera in Dunbar’s study, and >140 species by
DeCasien et al. The robustness of the latter may explain why other studies did not find a correlation of
diet with brain size (Navarrete et al., 2016), although here they also controlled for group size in their
analysis. Still, a more notable difference is the brain size characteristic (whole or neocortex) used.

The correlation of whole brain with diet may be more relevant in the context of the hypothesis it
defends. As authors suggest, regions outside of the neocortex are linked to intelligence – indeed,

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whole brain size is correlated with general intelligence (Deaner et al., 2007). More recently however,
this was disproven in some species, such as the lemurs. Fichtel et al. (2020) compared the cognitive
skills of lemurs on tests of physical and social domains and found unusually high performance, which
could not be explained by either brain-body ratio or social system. Thus, relative brain size can be
correlated with diet – nevertheless, analyses of behaviourally complexity (such as sociality,
visuospatial ability in the study mentioned) may benefit less from its use. As Healy and Rowe (2007)
review, early analyses of whole brain size later moved to investigating discrete areas, which were
linked to more specific functions (for instance, spatial memory correlated with hippocampus size in
birds). This, they argue, can provide more informative conclusions than general encephalization.

This perspective supports the finding that social complexity is supported by enlargement of specific
areas. For instance, amygdala volume (linked to social cue detection) correlated in humans with the
size of their social network (the number of people with whom one maintains regular contact), similar to
apes (Bickart et al., 2011). The density of amygdala and superior temporal sulcus (STS) was found to
be a better predictor of group size is humans and NHP than its volume (Murray & Fellows, 2022). A
causal link between area and function was also established. Lesioning the amygdala-STS connection
led to impairment in social cognition, while monkeys whose amygdala-orbitofrontal cortex(OFC) was
lesioned were impaired in object valuation (Murray & Fellows, 2022). These data suggest that discrete
brain areas were involved in social complexity in the SBH. While diet can explain whole brain
enlargement, social behaviour has a no less important role in brain development (and, as noted, may
increase grey matter density rather than volume), the causes of which will be explored further.

One of the likely explanations for the development of brain size due to diet is that energy constraints
were removed in brain development. Isler and van Schaik (2009) devised an evolutionary pattern
whereby relative brain size compensated with the energetic input or incurred trade-offs with other
functions (such as digestion, reproduction). For instance, they observe that mammals with large
brains have lower fertility than their small-brained relatives, but not when females display cooperative
care of young. Therefore sociality could have evolved to overcome the energetic constraint.
Nevertheless, this constraint has clear effects on brain development, for example in marsupials living
in highly seasonal areas – with seasonal low resource availability. These display smaller brains than
their relatives in low seasonal areas, despite small social differences (Weisbecker et al., 2015).

An alternative speculation on the evolution of brain size is related to foraging complexity. The
suggestion that developing the skill can induce the effect on brain (Lefebre, 2012) is true if there was
higher fitness of the large brain phenotype among other variants, which is eventually genetically
accommodated if alleles become associated with the phenotype. On these alleles, selection can then
act upon (West-Eberhard, 2005). However, the reverse is possible. Namely, a subpopulation of
primates with larger brain size, with higher energetic needs, may have improved their adaptation by
transitioning to frugivory, case in which the reason for brain development remains unexplained.

Similarly, the development of social behaviour, enabled by brain changes in the SBH, supports both
direct and indirect explanations. According to Dunbar (1992), the enlarged brain allows individuals to

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maintain larger groups. Although group size leads to more frequent conflicts, which would predict that
individuals disperse (group fragmentation), this is not the pattern observed in natural environments.
Instead, stress is resolved by affiliation to coalitions and grooming cliques, both of which have been
correlated in size with neocortex volume (Dunbar & Schultz, 2017). More importantly, fecundity of
females was linked to the size of their cliques, and females with more grooming partners had a better
response to stress, such that they have lower cortisol levels after stressful events (Dunbar & Schultz,
2017). Both fecundity and stress regulation can help improve the fitness of these females, which
provides a proximate mechanism for the selection of the favourable trait – brain size.

The perspective would suggest a direct effect of sociality on brain evolution, linking SBH with the
evolution of sociality. This assumes that sociality is the pressure that induced selection in brain size.
However, because a larger brain accommodated for improved social functioning, does not imply that
other factors were more significant in the process of encephalization. As warned by Gould and
Lewontin (1979), a presently adaptive trait is not necessarily the factor originally responsible for its
selection. In the context of SBH, there are observed markers of social complexity (e.g., cooperation
and deception) which could have gradually appeared in development and enabled group living rather
than representing the selective pressure.

Evidence for this comes from studies investigating the effect of social factors on brain development.
Maternal care appears to have the most significant effect on the adaptive ability of offspring in social
environments. As reviewed by Curley and Champagne (2016), care was correlated with the levels of
brain-derived neurotrophic factor (BDNF), a promoter of neurogenesis, such that reduced care
inactivated the gene in mice, and communal rearing further upregulated it. In primates, low maternal
involvement increased circulating cortisol releasing factor (CRF). Cortisol has an established negative
effect on neurogenesis (Odaka et al., 2017). These findings suggest that increased maternal care is a
likely contributor in the emergence of social brains, as this precedes group living in evolutionary time
(Isler & van Schaik, 2012).

In contrast, there is little evidence linking frugivory with increases in sociality, as previously noted
(Dunbar, 1992). Behaviourally, teaching smart foraging may be related to an improved diet, whose
transmission observed improvements in fitness (Dunbar & Schultz, 2017). The link remains mainly
metabolic – improved food quality enabled better offspring development, as seen in superior effects of
cooperative breeding and care (Isler & van Schaik, 2012). On these, sociality can then enhance
discrete structures (SBH). However, diet alone does not sufficiently explain emergence of sociality.

In sum, sociality was supported by the evolution of larger relative brain sizes in primates. The
mechanisms of this admit an interaction between diet and behavioural selection in brain evolution.
Analyses of both hypotheses (ecological and SBH) reveal different effects on brain development, with
larger brains due to nutritional quality, but larger brain areas related to sociality in more social
animals. Nevertheless, developmental factors related to brain function must be investigated before
establishing the role of the social brain hypothesis in the evolution of sociality.