The sizes of Brodmann's areas 44 and 45 (Broca's speech region) and their extent in relation to macroscopic landmarks and surrounding areas differ considerably among the available cytoarchitectonic maps. Such variability may be due to intersubject differences in anatomy, observer‐dependent discrepancies in cytoarchitectonic mapping, or both. Because a reliable definition of cytoarchitectonic borders is important for interpreting functional imaging data, we mapped areas 44 and 45 by means of an observer‐independent technique. In 10 human brains, the laminar distributions of cell densities were measured vertical to the cortical surface in serial coronal sections stained for perikarya. Thousands of density profiles were obtained. Cytoarchitectonic borders were defined as statistically significant changes in laminar patterns. The analysis of the three‐dimensional reconstructed brains and the two areas showed that cytoarchitectonic borders did not consistently coincide with sulcal contours. Therefore, macroscopic features are not reliable landmarks of cytoarchitectonic borders. Intersubject variability in the cytoarchitecture of areas 44 and 45 was significantly greater than cytoarchitectonic differences between these areas in individual brains. Although the volumes of area 44 differed across subjects by up to a factor of 10, area 44 but not area 45 was left‐over‐right asymmetrical in all brains. All five male but only three of five female brains had significantly higher cell densities on the left than on the right side. Such hemispheric and gender differences were not detected in area 45. These morphologic asymmetries of area 44 provide a putative correlate of the functional lateralization of speech production. J. Comp. Neurol. 412:319–341, 1999. © 1999 Wiley‐Liss, Inc.

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The degree of cortical folding found in adult human brains has been analyzed using a gyrification index (GI). This parameter permits the description of a mean value for the whole brain, but also a local specific analysis of different brain regions. Correlation analyses of the GI with age, body weight, body length, brain weight and volume of the prosencephalon and the cortex show no significant results. GI values do not differ significantly between male and female brains, right and left hemispheres or right and left sides of the superior temporal plane. The GI shows maximal values over the prefrontal and the parieto-temporo-occipital association cortex. A comparison between the rostro-caudal GI patterns of human brains and those of prosimians and Old World monkeys shows the largest difference over the prefrontal cortex. The mean GI increases from prosimians to human brains with the highest values for non-human primates being in the pongid group.

During development the human cortex changes from a smooth lissencephalic structure to one that is highly convoluted. Increases in the degree of cortical folding are associated with brain size only for the first part of brain growth; during the second half, differences in cortical folding match those of brain size, resulting in no change in the degree of folding. When the degree of cortical folding is studied as a function of age, a brief postnatal overshoot, an effect of brain size, is observed. The analysis suggests that the mechanical hypothesis of cortical buckling can best explain the degree of cortical folding, but that other hypotheses, like gyrogenesis, are required to explain the placement and orientation of sulci. The adult asymptote in degree of cortical folding is associated with the onset and disappearance of single subplate lamina, suggesting that subplate:cortical plate associations should be examined as causal for gyrification. Areas whose sulci differ in length between the two hemispheres have similar degrees of convolutedness, supporting interpretations that the sizes of gyri are asymmetric in the two hemispheres. The ontogenetic data support the thesis that human cortical proportions evolved when the brain enlarged in size and that the process was not one of neoteny.