The organization of pyramidal cells in area 18 of the rhesus monkey.

The aim of this study was to investigate the vertical organization of axons and pyramidal cells in area 18, and to compare it with that in area 17. In area 18 there are regularly spaced vertical bundles of myelinated axons that have an average center-to-center spacing of 21 microns. This arrangement of axons resembles that in area 17. Pyramidal cells in area 18 and their apical dendrites are less regularly arranged. The apical dendrites of the pyramidal cells of layer 6A aggregate with those from layer 5 pyramids to form swathes of apical dendrites that pass into layer 4. There they are joined by the apical dendrites of the small layer 4 pyramids, so that much of the neuropil of layer 4 is occupied by apical dendrites. Most of these apical dendrites form their terminal tufts in layer 3. Very few of them reach layer 1, which is dominated by the apical dendrites of layer 2/3 pyramids. Thus, there are two tiers of apical dendrites and their apical tufts, a deep one formed by the layer 4, 5 and 6 apical dendrites that terminate in layer 3, and a second one formed by the apical dendrites of layer 2/3 pyramids that terminate in layer 1. In contrast, in area 17 the apical dendrites of layer 5 pyramids form discrete clusters that have a center-to-center spacing of 23 microns. These clusters are joined by the apical dendrites of the layer 2/3 pyramids and all of these apical dendrites form their apical tufts in layer 1. Based upon the dispositions of the apical dendrites of the pyramidal cells in area 17 and 18, we speculate that the influences of, and the interactions between, the feed-forward and feed-back signals in the two areas are quite different, because in the two areas different postsynaptic targets are available to these afferents.