Architectionis, somatotopic organization, and ipsilateral cortical connections of the primary motor area (M1) of owl monkeys

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Abstract
The ipsilateral cortical connections of primary motor cortex (M1) of owl monkeys were revealed by injecting WGA‐HRP and fluorescent tracers into M1 sites identified by intracortical microstimulation. In some of the same animals, the extent and somatotopic organization of M1 was determined by making detailed microstimulation movement maps and relating the results to cortical architectonics. Thus, delineation of M1 was based on a combination of physiological and anatomical characteristics. M1 comprised most, but not all, of the cortex rostral to area 3a where movements were evoked at low levels of current (40 μA or less). Analysis of somatotopic patterns and architectonics placed some of the low‐threshold sites in a ventral premotor field (PMV) and the dorsomedially situated supplementary motor area (SMA). Movements were also reliably elicited from a dorsal premotor area (PMD) at higher currents. M1 was characterized by a somatotopic global organization, representing hindlimb, trunk, forelimb, and face movements in a mediolateral sequence, and a mosaic local organization, with a given movement typically represented at several different sites. Architectionically, M1 was characterized by the absence of a granular layer IV and the presence of very large layer V pyramidal cells. However, M1 was not uniform in structure: pyramidal cells were larger caudally than rostrally, a feature we used to distinguish caudal (M1c) and rostral (M1r) subdivisions of the area. M1 resembles Brodmann's area 4, although the rostral subdivision has probably been considered as part of area 6 by some workers. Tracer injections of M1 revealed somatotopically distributed connections with motor areas PMD, PMV, and SMA, as well as in somatosensory areas 3a, 1, 2, and S2. Weaker connections were with area 3b, posterior parietal cortex, the parietal ventral area (PV), and cingulate cortex. M1r and M1c differed connectionally as well as architectonically, M1c being connected primarily with somatosensory areas, while M1r was strongly connected with both non‐primary motor cortex and somatosensory cortex. These results indicate that M1 interacts directly with at least three non‐primary motor areas and at least six somatosensory areas.