Nongeniculate afferents to striate cortex in macaques

Horseradish peroxidase (HRP) was injected in relatively massive amounts to cover most, or portions, of opercular striate cortex in four macaques. Absence of transcallosal or circumventricular labelling, plus discrete and consistent retrograde labelling in other areas in the four cases, assured the validity and specificity of the observations. Numerous labelled cells in regions directly bordering striate cortex, however, were excluded from the analysis because of the possibility of uptake consequent to physical diffusion. With this exception, all labelled cells were counted at roughly 2-mm intervals for one case with extensive unilateral injection of HRP. Even excluding the closely circumstriate population, the totals indicate that more than 30% of the afferent input to striate cortex arises from nongeniculate sources. Four areas of neocortex together make up about one-fourth of the total afferents: superior temporal sulcus 17.1%; inferior occipital area, 6.1%; intraparietal sulcus, 0.4%; and parahippocampal gyrus, 0.3%. Other areas projecting to striate cortex include claustrum, pulvinar, nucleus paracentralis, raphé system, locus coeruleus, and the nucleus basalis of Meynert. Cells of the latter were particularly striking with their very heavy uptake of HRP, and, even in cases of minimal effective injection, were scattered throughout an extensive area from the posterior edge of the globus pallidus passing rostrally beyond the chiasm and into the nucleus of the diagonal band. On the basis of their distribution and known cholinergic affinity, it is argued that this group also includes the cells labelled in and around lateral hypothalamus and cerebral peduncle, and that as a whole the group constitutes a cholinergic counterpart of the diffusely projecting monoaminergic systems. It seems possible that the basalis projection at first follows a fornical-subcallosal pathway to reach striate cortex via callosoperforant fibers.