A reticular pattern of intrinsic connections in primate area V2 (area 18)

A system of periodic intrinsic connections is demonstrated in area V2 (area 18) of squirrel and macaque monkeys by large injections of tritiated amino acids, horseradish peroxidase (HRP), and fluorescent latex beads. These connections originate from pyramidal neurons concentrated in layers 3 and 5. Terminations occur in all cortical layers, largely coextensive with labeled neurons but more restricted in layer 4. This multilaminar distribution contrasts with the mainly supragranular localization of periodic intrinsic connections in V1 (area 17), and may imply a close interaction, in V2, of periodic intrinsic connections with pulvinocortical, as well as with corticocortical terminations (concentrated, respectively, in layers 3 and 5, and in lower 3 and 4). As in V1, the tangential configuration of these connections in V2 is reticular or latticelike, and is detectable for 2.5-3.0 mm from an injection site of HRP, 3H amino acids, or latex beads. Cross-sectional widths of labeled regions vary from 250 to 800 micron in squirrel monkey and from 400 to 1,000 micron in macaque, depending on which portion of the lattice is measured. When periodic intrinsic connections are compared with stripes labeled histochemically by cytochrome oxidase (CO), no clear relationship is obvious between the two systems. This result contrasts with the orderly tangential alignment reported between CO-reactive zones in V2 and certain extrinsic connections; namely, pulvinocortical terminations (Livingstone and Hubel, '82) and clusters of neurons projecting to area V4 (DeYoe and Van Essen, '84). Other extrinsic connections, however, such as backgoing connections from V2 to V1, do not seem to have a periodic distribution. Thus, although some discontinuous cortical connections relate to each other in a precise mosaic fashion, intrinsic and some extrinsic connections may observe different modes of organization.