Barrels and septa: separate circuits in rat barrels field cortex.

The neural circuitry within sensory cortex determines its functional properties, and different solutions have evolved for integrating the activity that arises from an array of sensory inputs to cortex. In rodent, circumscribed receptors, such as whiskers, are represented in somatic sensory (S-I) cortex in islands of cells in layer IV called "barrels" surrounded by narrow channels that separate barrels called "septa." These two cortical domains were previously shown to receive sensory inputs through parallel subcortical pathways. Here, by using small biocytin injections, we demonstrate that distinct intrinsic and corticocortical circuitries arise from barrel and septal columns. The intracortical S-I projections originating from barrel columns are rather short-ranged, terminating for the most part within the far boundaries of the most immediate neighboring barrel columns, whereas corticocortical projections reach the second somatosensory (S-II) cortex. In contrast, the intrinsic projections arising from septal columns extend two to three barrels' distance along the row of whisker representation, producing terminals preferentially in other septal columns. Septal corticocortical projections terminate in the dysgranular cortex anterior to E-row barrels and in the posteromedial parietal cortex in addition to S-II. Whereas layer IV barrels are largely isolated from lateral connections, septa are the main conduits of intracortical projections arising from neighboring barrel and septal columns. These results indicate that the two subcortical pathways from whiskers to cortex continue as two distinct partially segregated pathways in cortex.