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1.  Dolphin insula reflects minicolumnar organization of mammalian isocortex 
Translational neuroscience  2010;1(1):37-42.
The brain of the bottlenose dolphin exhibits patterns of isocortical parcellation and cytoarchitecture distinct from those seen in primates, yet cell clusters in anterior insula are comparable in scale to module-like cell arrangements found throughout isocortex in other placental mammalian species with long divergent evolutionary histories. This similarity may be due to common ancestry, or to convergence as a result of selective constraints on organization of connections within such modules. Differences reflect alternate arrangements of minicolumns, an elemental cytoarchitectonic motif of isocortex defined by radially oriented pyramidal cell arrays. In contrast with larger modular structures incorporating them, minicolumns have been highly conserved in mammalian evolution. In this study a previously validated imaging method was employed to assess verticality, D, a parameter indicating radial bias of isocortex. Photomicrographs of coronal Nissl-stained sections of dolphin anterior insular cortex were compared with sections from human brains of putatively homologous areas as well as other isocortical areas differing in modular organization. Dolphin insula exhibited a high degree of verticality consistent with conserved minicolumnar organization. Our findings indicate that a basic structural motif of isocortex is synapomorphic in a species of marine mammal exhibiting unique phylogenetically derived isocortical characteristics.
doi:10.2478/v10134-010-0010-2
PMCID: PMC3332127  PMID: 22532930
Cetaceans; Neocortex; Minicolumns; Pyramidal cells; Tursiops truncatus
2.  Social intelligence in the spotted hyena (Crocuta crocuta) 
If the large brains and great intelligence characteristic of primates were favoured by selection pressures associated with life in complex societies, then cognitive abilities and nervous systems with primate-like attributes should have evolved convergently in non-primate mammals living in large, elaborate societies in which social dexterity enhances individual fitness. The societies of spotted hyenas are remarkably like those of cercopithecine primates with respect to size, structure and patterns of competition and cooperation. These similarities set an ideal stage for comparative analysis of social intelligence and nervous system organization. As in cercopithecine primates, spotted hyenas use multiple sensory modalities to recognize their kin and other conspecifics as individuals, they recognize third-party kin and rank relationships among their clan mates, and they use this knowledge adaptively during social decision making. However, hyenas appear to rely more intensively than primates on social facilitation and simple rules of thumb in social decision making. No evidence to date suggests that hyenas are capable of true imitation. Finally, it appears that the gross anatomy of the brain in spotted hyenas might resemble that in primates with respect to expansion of frontal cortex, presumed to be involved in the mediation of social behaviour.
doi:10.1098/rstb.2006.1993
PMCID: PMC2346515  PMID: 17289649
hyena; hyaena; intelligence; social cognition; brain morphology
3.  An Exploration of the Social Brain Hypothesis in Insects 
The “social brain hypothesis” posits that the cognitive demands of sociality have driven the evolution of substantially enlarged brains in primates and some other mammals. Whether such reasoning can apply to all social animals is an open question. Here we examine the evolutionary relationships between sociality, cognition, and brain size in insects, a taxonomic group characterized by an extreme sophistication of social behaviors and relatively simple nervous systems. We discuss the application of the social brain hypothesis in this group, based on comparative studies of brain volumes across species exhibiting various levels of social complexity. We illustrate how some of the major behavioral innovations of social insects may in fact require little information-processing and minor adjustments of neural circuitry, thus potentially selecting for more specialized rather than bigger brains. We argue that future work aiming to understand how animal behavior, cognition, and brains are shaped by the environment (including social interactions) should focus on brain functions and identify neural circuitry correlates of social tasks, not only brain sizes.
doi:10.3389/fphys.2012.00442
PMCID: PMC3506958  PMID: 23205013
cognition; insects; mushroom bodies; sociality; social brain hypothesis
4.  Social brains, simple minds: does social complexity really require cognitive complexity? 
The social brain hypothesis is a well-accepted and well-supported evolutionary theory of enlarged brain size in the non-human primates. Nevertheless, it tends to emphasize an anthropocentric view of social life and cognition. This often leads to confusion between ultimate and proximate mechanisms, and an over-reliance on a Cartesian, narratively structured view of the mind and social life, which in turn lead to views of social complexity that are congenial to our views of ourselves, rather than necessarily representative of primate social worlds. In this paper, we argue for greater attention to embodied and distributed theories of cognition, which get us away from current fixations on ‘theory of mind’ and other high-level anthropocentric constructions, and allow for the generation of testable hypotheses that combine neurobiology, psychology and behaviour in a mutually reinforcing manner.
doi:10.1098/rstb.2006.1995
PMCID: PMC2346517  PMID: 17255006
social complexity; primates; anthropocentrism; social brain; distributed cognition; embodied cognition
5.  Group Size Predicts Social but Not Nonsocial Cognition in Lemurs 
PLoS ONE  2013;8(6):e66359.
The social intelligence hypothesis suggests that living in large social networks was the primary selective pressure for the evolution of complex cognition in primates. This hypothesis is supported by comparative studies demonstrating a positive relationship between social group size and relative brain size across primates. However, the relationship between brain size and cognition remains equivocal. Moreover, there have been no experimental studies directly testing the association between group size and cognition across primates. We tested the social intelligence hypothesis by comparing 6 primate species (total N = 96) characterized by different group sizes on two cognitive tasks. Here, we show that a species’ typical social group size predicts performance on cognitive measures of social cognition, but not a nonsocial measure of inhibitory control. We also show that a species’ mean brain size (in absolute or relative terms) does not predict performance on either task in these species. These data provide evidence for a relationship between group size and social cognition in primates, and reveal the potential for cognitive evolution without concomitant changes in brain size. Furthermore our results underscore the need for more empirical studies of animal cognition, which have the power to reveal species differences in cognition not detectable by proxy variables, such as brain size.
doi:10.1371/journal.pone.0066359
PMCID: PMC3694165  PMID: 23840450
6.  The social nature of primate cognition 
The hypothesis that the enlarged brain size of the primates was selected for by social, rather than purely ecological, factors has been strongly influential in studies of primate cognition and behaviour over the past two decades. However, the Machiavellian intelligence hypothesis, also known as the social brain hypothesis, tends to emphasize certain traits and behaviours, like exploitation and deception, at the expense of others, such as tolerance and behavioural coordination, and therefore presents only one view of how social life may shape cognition. This review outlines work from other relevant disciplines, including evolutionary economics, cognitive science and neurophysiology, to illustrate how these can be used to build a more general theoretical framework, incorporating notions of embodied and distributed cognition, in which to situate questions concerning the evolution of primate social cognition.
doi:10.1098/rspb.2005.3200
PMCID: PMC1559889  PMID: 16191591
primate; social brain; embodied cognition; distributed cognition; mirror neurons
7.  Evolution, development, and plasticity of the human brain: from molecules to bones 
Neuroanatomical, molecular, and paleontological evidence is examined in light of human brain evolution. The brain of extant humans differs from the brains of other primates in its overall size and organization, and differences in size and organization of specific cortical areas and subcortical structures implicated into complex cognition and social and emotional processing. The human brain is also characterized by functional lateralizations, reflecting specializations of the cerebral hemispheres in humans for different types of processing, facilitating fast and reliable communication between neural cells in an enlarged brain. The features observed in the adult brain reflect human-specific patterns of brain development. Compared to the brains of other primates, the human brain takes longer to mature, promoting an extended period for establishing cortical microcircuitry and its modifications. Together, these features may underlie the prolonged period of learning and acquisition of technical and social skills necessary for survival, creating a unique cognitive and behavioral niche typical of our species. The neuroanatomical findings are in concordance with molecular analyses, which suggest a trend toward heterochrony in the expression of genes implicated in different functions. These include synaptogenesis, neuronal maturation, and plasticity in humans, mutations in genes implicated in neurite outgrowth and plasticity, and an increased role of regulatory mechanisms, potentially promoting fast modification of neuronal morphologies in response to new computational demands. At the same time, endocranial casts of fossil hominins provide an insight into the timing of the emergence of uniquely human features in the course of evolution. We conclude by proposing several ways of combining comparative neuroanatomy, molecular biology and insights gained from fossil endocasts in future research.
doi:10.3389/fnhum.2013.00707
PMCID: PMC3812990  PMID: 24194709
pyramidal neurons; plasticity; neuropsin; brain evolution; development; amygdala; endocast; human evolution
8.  Sexual Selection and the Evolution of Brain Size in Primates 
PLoS ONE  2006;1(1):e62.
Reproductive competition among males has long been considered a powerful force in the evolution of primates. The evolution of brain size and complexity in the Order Primates has been widely regarded as the hallmark of primate evolutionary history. Despite their importance to our understanding of primate evolution, the relationship between sexual selection and the evolutionary development of brain size is not well studied. The present research examines the evolutionary relationship between brain size and two components of primate sexual selection, sperm competition and male competition for mates. Results indicate that there is not a significant relationship between relative brain size and sperm competition as measured by relative testis size in primates, suggesting sperm competition has not played an important role in the evolution of brain size in the primate order. There is, however, a significant negative evolutionary relationship between relative brain size and the level of male competition for mates. The present study shows that the largest relative brain sizes among primate species are associated with monogamous mating systems, suggesting primate monogamy may require greater social acuity and abilities of deception.
doi:10.1371/journal.pone.0000062
PMCID: PMC1762360  PMID: 17183693
9.  Regional Patterns of Cerebral Cortical Differentiation Determined by Diffusion Tensor MRI 
Cerebral Cortex (New York, NY)  2009;19(12):2916-2929.
The morphology of axonal and dendritic arbors in the immature cerebral cortex influences the degree of anisotropy in water diffusion. This enables cortical maturation to be monitored by the noninvasive technique of diffusion tensor magnetic resonance imaging (DTI). Herein, we utilized DTI of postmortem ferret brain to quantify regional and temporal patterns in cortical maturation. We found that diffusion anisotropy within the isocortex decreases over the first month of life, coinciding closely in time with expansion of axonal and dendritic cellular processes of pyramidal neurons. Regional patterns consist of differences between allocortex and isocortex, a regional anisotropy gradient that closely parallels the transverse neurogenetic gradient, and differences between primary and nonprimary isocortical areas. By combining the temporal and regional factors, the isocortical developmental gradient magnitude corresponds to a 5-day difference in maturity between relatively developed rostral/caudal isocortex at the gradient source and less mature isocortex at the occipital pole. Additionally, the developmental trajectory of primary areas precedes nonprimary areas by 2.7 days. These quantitative estimates coincide with previous histological studies of ferret development. Similarities in cerebral cortical diffusion anisotropy observed between ferret and other species suggest the framework developed here is of general potential relevance.
doi:10.1093/cercor/bhp061
PMCID: PMC2774393  PMID: 19363145
brain; development; DTI; dendritic branching; ferret
10.  Cognitive adaptations of social bonding in birds 
The ‘social intelligence hypothesis’ was originally conceived to explain how primates may have evolved their superior intellect and large brains when compared with other animals. Although some birds such as corvids may be intellectually comparable to apes, the same relationship between sociality and brain size seen in primates has not been found for birds, possibly suggesting a role for other non-social factors. But bird sociality is different from primate sociality. Most monkeys and apes form stable groups, whereas most birds are monogamous, and only form large flocks outside of the breeding season. Some birds form lifelong pair bonds and these species tend to have the largest brains relative to body size. Some of these species are known for their intellectual abilities (e.g. corvids and parrots), while others are not (e.g. geese and albatrosses). Although socio-ecological factors may explain some of the differences in brain size and intelligence between corvids/parrots and geese/albatrosses, we predict that the type and quality of the bonded relationship is also critical. Indeed, we present empirical evidence that rook and jackdaw partnerships resemble primate and dolphin alliances. Although social interactions within a pair may seem simple on the surface, we argue that cognition may play an important role in the maintenance of long-term relationships, something we name as ‘relationship intelligence’.
doi:10.1098/rstb.2006.1991
PMCID: PMC2346513  PMID: 17255008
avian brain; jackdaw; monogamy; pair bonding; rook; social intelligence
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.  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
13.  Neocortex size predicts deception rate in primates. 
Human brain organization is built upon a more ancient adaptation, the large brain of simian primates: on average, monkeys and apes have brains twice as large as expected for mammals of their size, principally as a result of neocortical enlargement. Testing the adaptive benefit of this evolutionary specialization depends on finding an association between brain size and function in primates. However, most cognitive capacities have been assessed in only a restricted range of species under laboratory conditions. Deception of conspecifics in social circumstances is an exception, because a corpus of field data is available that encompasses all major lines of the primate radiation. We show that the use of deception within the primates is well predicted by the neocortical volume, when observer effort is controlled for; by contrast, neither the size of the rest of the brain nor the group size exert significant effects. These findings are consistent with the hypothesis that neocortical expansion has been driven by social challenges among the primates. Complex social manipulations such as deception are thought to be based upon rapid learning and extensive social knowledge; thus, learning in social contexts may be constrained by neocortical size.
doi:10.1098/rspb.2004.2780
PMCID: PMC1691785  PMID: 15306289
14.  Visual specialization and brain evolution in primates. 
Several theories have been proposed to explain the evolution of species differences in brain size, but no consensus has emerged. One unresolved question is whether brain size differences are a result of neural specializations or of biological constraints affecting the whole brain. Here I show that, among primates, brain size variation is associated with visual specialization. Primates with large brains for their body size have relatively expanded visual brain areas, including the primary visual cortex and lateral geniculate nucleus. Within the visual system, it is, in particular, one functionally specialized pathway upon which selection has acted: evolutionary changes in the number of neurons in parvocellular, but not magnocellular, layers of the lateral geniculate nucleus are correlated with changes in both brain size and ecological variables (diet and social group size). Given the known functions of the parvocellular pathway, these results suggest that the relatively large brains of frugivorous species are products of selection on the ability to perceive and select fruits using specific visual cues such as colour. The separate correlation between group size and visual brain evolution, on the other hand, may indicate the visual basis of social information processing in the primate brain.
PMCID: PMC1689478  PMID: 9821360
15.  Cognitive Consilience: Primate Non-Primary Neuroanatomical Circuits Underlying Cognition 
Interactions between the cerebral cortex, thalamus, and basal ganglia form the basis of cognitive information processing in the mammalian brain. Understanding the principles of neuroanatomical organization in these structures is critical to understanding the functions they perform and ultimately how the human brain works. We have manually distilled and synthesized hundreds of primate neuroanatomy facts into a single interactive visualization. The resulting picture represents the fundamental neuroanatomical blueprint upon which cognitive functions must be implemented. Within this framework we hypothesize and detail 7 functional circuits corresponding to psychological perspectives on the brain: consolidated long-term declarative memory, short-term declarative memory, working memory/information processing, behavioral memory selection, behavioral memory output, cognitive control, and cortical information flow regulation. Each circuit is described in terms of distinguishable neuronal groups including the cerebral isocortex (9 pyramidal neuronal groups), parahippocampal gyrus and hippocampus, thalamus (4 neuronal groups), basal ganglia (7 neuronal groups), metencephalon, basal forebrain, and other subcortical nuclei. We focus on neuroanatomy related to primate non-primary cortical systems to elucidate the basis underlying the distinct homotypical cognitive architecture. To display the breadth of this review, we introduce a novel method of integrating and presenting data in multiple independent visualizations: an interactive website (http://www.frontiersin.org/files/cognitiveconsilience/index.html) and standalone iPhone and iPad applications. With these tools we present a unique, annotated view of neuroanatomical consilience (integration of knowledge).
doi:10.3389/fnana.2011.00065
PMCID: PMC3243081  PMID: 22194717
cerebral cortex; thalamus; basal ganglia; circuitry; consilience; isocortex; cognition
16.  The costs and benefits of flexibility as an expression of behavioural plasticity: a primate perspective 
Traditional neo-Darwinism ascribes geographical variation in morphology or in behaviour to varying selection on local genotypes. However, mobile and long-lived organisms cannot achieve local adaptation this way, leading to a renewed interest in plasticity. I examined geographical variation in orang-utan subsistence and social behaviour, and found this to be largely owing to behavioural plasticity, here called flexibility, both in the form of flexible individual decisions and of socially transmitted (cultural) innovations. Although comparison with other species is difficult, the extent of such flexibility is almost certainly limited by brain size. It is shown that brains can only increase relative to body size where the cognitive benefits they produce are reliably translated into improved survival rate. This means that organisms that are very small, face many predators, live in highly seasonal environments, or lack opportunities for social learning cannot evolve greater flexibility, and must achieve local adaptation through selection on specific genotypes. On the other hand, as body and brain size increase, local adaptation is increasingly achieved through selection on plasticity. The species involved are also generally those that most need it, being more mobile and longer-lived. Although high plasticity buffers against environmental change, the most flexible organisms face a clear limit because they respond slowly to selection. Thus, paradoxically, the largest-brained animals may actually be vulnerable to the more drastic forms of environmental change, such as those induced by human actions.
doi:10.1098/rstb.2012.0339
PMCID: PMC3638442  PMID: 23569287
geographical variation; local adaptation; flexibility
17.  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
18.  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
19.  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
20.  Expensive Brains: “Brainy” Rodents have Higher Metabolic Rate 
Brains are the centers of the nervous system of animals, controlling the organ systems of the body and coordinating responses to changes in the ecological and social environment. The evolution of traits that correlate with cognitive ability, such as relative brain size is thus of broad interest. Brain mass relative to body mass (BM) varies among mammals, and diverse factors have been proposed to explain this variation. A recent study provided evidence that energetics play an important role in brain evolution (Isler and van Schaik, 2006). Using composite phylogenies and data drawn from multiple sources, these authors showed that basal metabolic rate (BMR) correlates with brain mass across mammals. However, no such relationship was found within rodents. Here we re-examined the relationship between BMR and brain mass within Rodentia using a novel species-level phylogeny. Our results are sensitive to parameter evaluation; in particular how species mass is estimated. We detect no pattern when applying an approach used by previous studies, where each species BM is represented by two different numbers, one being the individual that happened to be used for BMR estimates of that species. However, this approach may compromise the analysis. When using a single value of BM for each species, whether representing a single individual, or available species mean, our findings provide evidence that brain mass (independent of BM) and BMR are correlated. These findings are thus consistent with the hypothesis that large brains evolve when the payoff for increased brain mass is greater than the energetic cost they incur.
doi:10.3389/fnevo.2011.00002
PMCID: PMC3141350  PMID: 21811456
Rodentia; Bayesian inference; correlated evolution
21.  Phylogenetic signal in primate behaviour, ecology and life history 
Examining biological diversity in an explicitly evolutionary context has been the subject of research for several decades, yet relatively recent advances in analytical techniques and the increasing availability of species-level phylogenies, have enabled scientists to ask new questions. One such approach is to quantify phylogenetic signal to determine how trait variation is correlated with the phylogenetic relatedness of species. When phylogenetic signal is high, closely related species exhibit similar traits, and this biological similarity decreases as the evolutionary distance between species increases. Here, we first review the concept of phylogenetic signal and suggest how to measure and interpret phylogenetic signal in species traits. Second, we quantified phylogenetic signal in primates for 31 variables, including body mass, brain size, life-history, sexual selection, social organization, diet, activity budget, ranging patterns and climatic variables. We found that phylogenetic signal varies extensively across and even within trait categories. The highest values are exhibited by brain size and body mass, moderate values are found in the degree of territoriality and canine size dimorphism, while low values are displayed by most of the remaining variables. Our results have important implications for the evolution of behaviour and ecology in primates and other vertebrates.
doi:10.1098/rstb.2012.0341
PMCID: PMC3638444  PMID: 23569289
macroevolution; phylogenetic comparative methods; trait variation; Brownian motion; mammal
22.  Fractionation of social brain circuits in autism spectrum disorders 
Brain  2012;135(9):2711-2725.
Autism spectrum disorders are developmental disorders characterized by impairments in social and communication abilities and repetitive behaviours. Converging neuroscientific evidence has suggested that the neuropathology of autism spectrum disorders is widely distributed, involving impaired connectivity throughout the brain. Here, we evaluate the hypothesis that decreased connectivity in high-functioning adolescents with an autism spectrum disorder relative to typically developing adolescents is concentrated within domain-specific circuits that are specialized for social processing. Using a novel whole-brain connectivity approach in functional magnetic resonance imaging, we found that not only are decreases in connectivity most pronounced between regions of the social brain but also they are selective to connections between limbic-related brain regions involved in affective aspects of social processing from other parts of the social brain that support language and sensorimotor processes. This selective pattern was independently obtained for correlations with measures of social symptom severity, implying a fractionation of the social brain in autism spectrum disorders at the level of whole circuits.
doi:10.1093/brain/aws160
PMCID: PMC3437021  PMID: 22791801
autism spectrum disorders; functional connectivity; resting state functional MRI; limbic system; cluster analysis
23.  Age-Related Differences in Corpus Callosum Area of Capuchin Monkeys 
Neuroscience  2011;202:202-208.
Capuchin monkeys (Cebus apella) are New World primates with relatively large brains for their body size. The developmental trajectories of several brain regions – including cortical white matter, frontal lobe white matter, and basal ganglia nuclei – are similar to humans. Additionally, capuchins have independently evolved several behavioral and anatomical characteristics in common with humans and chimpanzees – including complex manipulative abilities, use of tools, and the use of precision grips – making them interesting species for studies of comparative brain morphology and organization. Here we report the first investigation into the development of the corpus callosum and its regional subdivisions in capuchins. Corpus callosum development was quantified using high-resolution structural MRI images from 39 socially reared subjects (male n = 22; female n = 18) ranging in age from 4 days (infancy) – 20 years (middle adulthood). The total area of the corpus callosum and the subdivisions of the genu, rostral midbody, medial midbody, caudal midbody, and splenium were traced from the midsagittal section. Total corpus callosum area displayed significant differences across this time span and was best explained by quadratic growth. Sustained linear growth was observed in the subdivisions of the genu, rostral midbody, and splenium; sustained quadratic growth was seen in the subdivision of the medial midbody. Differences in growth were not detected in the subdivision of the caudal midbody. Females had a larger raw area of the total CC and of the medial midbody and caudal midbody throughout the lifespan. Our results indicate that capuchins show continued white matter development beyond adolescence in regions related to cognitive and motor development.
doi:10.1016/j.neuroscience.2011.11.074
PMCID: PMC3293371  PMID: 22173013
brain development; Cebus; corpus callosum
24.  The Dorsal Pallium in Zebrafish, Danio rerio (Cyprinidae, Teleostei) 
Brain research  2011;1381:95-105.
Zebrafish as a neurogenetic model system depends on the correct neuroanatomical understanding of its brain organization. Here, we address the unresolved question regarding a possible zebrafish homologue of the dorsal pallial division, the region that in mammals gives rise to the isocortex. Analyzing the distributions of nicotine adenine dinucleotide phosphate diphorase (NADPHd) activity and parvalbumin in the anterior zebrafish telencephalon, we show that against previous assumptions the central (Dc) zone possesses its own germinative region in the dorsal proliferative zone. We define the central (Dc) zone as topologically corresponding to the dorsal pallial division of other vertebrates (mammalian isocortex). In addition, we confirm through BrdU-labeling experiments that the posterior (Dp) zone is formed by radial migration and homologous to the mammalian piriform cortex. Based on our results, we propose a new developmental and organizational model of the zebrafish pallium—one which is the result of a complex outward-inward folding.
doi:10.1016/j.brainres.2010.12.089
PMCID: PMC3052766  PMID: 21219890
amygdala; cortex; eversion; evagination; forebrain; migration; ray-finned fish; teleost
25.  Maternal-fetal unit interactions and eutherian neocortical development and evolution 
The conserved brain design that primates inherited from early mammals differs from the variable adult brain size and species-specific brain dominances observed across mammals. This variability relies on the emergence of specialized cerebral cortical regions and sub-compartments, triggering an increase in brain size, areal interconnectivity and histological complexity that ultimately lies on the activation of developmental programs. Structural placental features are not well correlated with brain enlargement; however, several endocrine pathways could be tuned with the activation of neuronal progenitors in the proliferative neocortical compartments. In this article, we reviewed some mechanisms of eutherians maternal–fetal unit interactions associated with brain development and evolution. We propose a hypothesis of brain evolution where proliferative compartments in primates become activated by “non-classical” endocrine placental signals participating in different steps of corticogenesis. Changes in the inner placental structure, along with placenta endocrine stimuli over the cortical proliferative activity would allow mammalian brain enlargement with a concomitant shorter gestation span, as an evolutionary strategy to escape from parent-offspring conflict.
doi:10.3389/fnana.2013.00022
PMCID: PMC3715729  PMID: 23882189
cerebral cortex development; placenta; maternal–fetal unit; evolution; serotonin; eutherians; transcriptome

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