Connections of area 8 with area 6 in the brain of the macaque monkey

The corticocortical connections between the arcuate area (Walker's areas 8A and 45 or Brodmann's area 8) and the premotor and supplementary motor areas (Vogts' area 6) in the brain of the macaque monkey were studied microscopically with wheat germ agglutinin-conjugated horseradish peroxidase (WGA-HRP), which was injected into Brodmann's area 8 primarily to elucidate the projections of area 8 into area 6. In addition, in the same material, the pattern of connections between the arcuate area and area 46 of the prefrontal cortex was studied. On the basis of the findings of the present study, an anatomical cortical hierarchy for areas 8, 6, and 46 is discussed. Anterogradely labeled axon terminals and retrogradely labeled cells appeared in the premotor area, in the supplementary motor area, and in area 46 on the side of injection; sites containing labeled axon terminals also contained labeled cells. In other words, the examined connections were reciprocal. In the labeled areas, labeled terminals and cells coexisted, and most formed radial columns. However, no labeling of cells and terminals was seen in the motor area (Brodmann's area 4). After injection of WGA-HRP into area 8A, labeled terminals and cells appeared predominantly in the superior premotor area (a region of the premotor area above the arcuate spur, Vogt and Vogt's upper areas 6a alpha and 6a beta), forming one, two, or three bands of label in the anteroposterior direction, whereas labeling occurred to a lesser extent in the inferior premotor area (a region of the premotor area below the arcuate spur, Vogt and Vogt's areas 4c, lower 6a alpha and 6b). In contrast, injection of WGA-HRP into area 45 resulted in the predominant labeling of the inferior premotor area with scant labeling in the superior premotor area. In the premotor area, labeled terminals were distributed over the entire cortical depth, although few were found in the lower half of layer III, and labeled cells were distributed mainly in layer III. In area 46 of the banks of the principal sulcus in the prefrontal cortex, labeled terminals were distributed in all cortical layers or over the entire cortical depth with a lower concentration in layer IV; labeled cells were found mostly in layers III and V, with a relatively high density in layer V.