The neuronal composition of area 17 of rat visual cortex. IV. The organization of pyramidal cells

In area 17 of the rat visual cortex, most of the apical dendrites of the large layer V pyramidal cells aggregate into oriented clusters, each containing three or more such dendrites. These clusters are not randomly distributed, but have a basic hexagonal packing distribution in which the mean center-to-center spacing is 55-60 microns. The majority of medium-size pyramidal cells of layer V also add their apical dendrites to the clusters. As these clusters pass through layer IV they remain intact, and successively layer III and finally layer II pyramids add their apical dendrites to them. Perhaps because the pyramidal cells in layer II/III are so numerous, some of their apical dendrites form independent groups. Apical dendrites of the small pyramidal neurons in layers VIa and IV seem not to specifically add to the clusters. Instead, apical dendrites of layer VIa pyramids form into contiguous fascicles and sheets, which pass around the groups of layer V pyramidal cell bodies to ascend to layer IV, where most of them form their apical tufts. Layer IV pyramidal cell apical dendrites behave somewhat similarly. These apical dendrites have to pass between the cell bodies of the lower layer III pyramidal cells. To do this, some join the clusters, but others form independent bundles. It is suggested that the pyramidal cells whose apical dendrites are clustered represent vertically oriented neuronal modules whose activity is synchronized, and that different combinations of these modules are excited by afferents to the cortex to provide the bases for the various kinds of functional columns.