Neural changes that occurred during human evolution to support language are poorly understood. As a basis of comparison to humans, we used design-based stereological methods to estimate volumes, total neuron numbers, and neuron densities in Brodmann's areas 44 and 45 in both cerebral hemispheres of 12 chimpanzees (Pan troglodytes), one of our species' closest living relatives. We found that the degree of interindividual variation in the topographic location and quantitative cytoarchitecture of areas 44 and 45 in chimpanzees was comparable to that seen in humans from previous studies. However, in contrast to the documented asymmetries in humans, we did not find significant population-level hemispheric asymmetry for any measures of areas 44 and 45 in chimpanzees. Furthermore, there was no relationship between asymmetries of stereological data and magnetic resonance imaging-based measures of inferior frontal gyrus morphology or hand preference on 2 different behavioral tasks. These findings suggest that Broca's area in the left hemisphere expanded in relative size during human evolution, possibly as an adaptation for our species' language abilities.

Human language is distinctive compared with the communication systems of other species. Yet, several questions concerning its emergence and evolution remain unresolved. As a means of evaluating the neuroanatomical changes relevant to language that accompanied divergence from the last common ancestor of chimpanzees, bonobos and humans, we defined the cytoarchitectonic boundaries of area Tpt, a component of Wernicke's area, in 12 common chimpanzee brains and used designbased stereologic methods to estimate regional volumes, total neuron number and neuron density. In addition, we created a probabilistic map of the location of area Tpt in a template chimpanzee brain coordinate space. Our results show that chimpanzees display significant population-level leftward asymmetry of area Tpt in terms of neuron number, with volume asymmetry approaching significance. Furthermore, asymmetry in the number of neurons in area Tpt was positively correlated with asymmetry of neuron numbers in Brodmann's area 45, a component of Broca's frontal language region. Our findings support the conclusion that leftward asymmetry of Wernicke's area originated prior to the appearance of modern human language and before our divergence from the last common ancestor. Moreover, this study provides the first evidence of covariance between asymmetry of anterior and posterior cortical regions that in humans are important to language and other higher order cognitive functions.

Although behavioral lateralization is known to correlate with certain aspects of brain asymmetry in primates, there are limited data concerning hemispheric biases in the microstructure of the neocortex. In the present study, we investigated whether there is asymmetry in synaptophysin-immunoreactive puncta density and protein expression levels in the region of hand representation of the primary motor cortex in chimpanzees (Pan troglodytes). Synaptophysin is a presynaptic vesicle-associated protein found in nearly all synapses of the central nervous system. We also tested whether there is a relationship between hand preference on a coordinated bimanual task and the interhemispheric distribution of synaptophysin as measured by both stereologic counts of immunoreactive puncta and by Western blotting. Our results demonstrated that synaptophysin-immunoreactive puncta density is not asymmetric at the population level, whereas synaptophysin protein expression levels are significantly higher in the right hemisphere. Handedness was correlated with interindividual variation in synaptophysin-immunoreactive puncta density. As a group, left-handed and ambidextrous chimpanzees showed a rightward bias in puncta density. In contrast, puncta densities were symmetrical in right-handed chimpanzees. These findings support the conclusion that synapse asymmetry is modulated by lateralization of skilled motor behavior in chimpanzees.