Anatomical organization of forebrain circuits in the primate.

The primate forebrain is a complex structure. Thousands of connections have been identified between cortical areas, and between cortical and sub-cortical areas. Previous work, however, has suggested that a number of principles can be used to reduce this complexity. Here, we integrate four principles that have been put forth previously, including a nested model of neocortical connectivity, gradients of connectivity between frontal cortical areas and the striatum and thalamus, shared patterns of sub-cortical connectivity between connected posterior and frontal cortical areas, and topographic organization of cortical-striatal-pallidal-thalamocortical circuits. We integrate these principles into a single model that accounts for a substantial amount of connectivity in the forebrain. We then suggest that studies in evolution and development can account for these four principles, by assuming that the ancestral vertebrate pallium was dominated by medial, hippocampal and ventral-lateral, pyriform areas, and at most a small dorsal pallium. The small dorsal pallium expanded massively in the lineage leading to primates. During this expansion, topological, adjacency relationships were maintained between pallial and sub-pallial areas. This maintained topology led to the connectivity gradients seen between cortex, striatum, pallidum, and thalamus.