Synaptic linkages between corticomotoneuronal cells affecting forelimb muscles in behaving primates

To elucidate the cortical circuitry controlling primate forelimb muscles we investigated the synaptic interactions between neighboring motor cortex cells that had postspike output effects in target muscles. In monkeys generating isometric ramp-and-hold wrist torques, pairs of cortical cells were recorded simultaneously with independent electrodes and corticomotoneuronal ("CM") cells were identified by their postspike effects on target forelimb muscles in spike-triggered averages (SpTAs) of electromyographs (EMGs). The response patterns of the cells were determined in response-aligned averages and their synaptic interactions were identified by cross-correlograms of action potentials. The possibility that synchronized firing between cortical cells could mediate spike-correlated effects in the SpTA of EMG was examined in several ways. Sixty-two pairs consisted of one CM cell and a non-CM cell; 15 of these had correlogram peaks of the same magnitude as that of other pairs, but the synchrony peaks did not mediate any postspike effect from the non-CM cell. Twelve pairs of simultaneously recorded CM cells were cross-correlated. Half had features (usually synchrony peaks) in their cross-correlograms and the cells of these pairs also shared some target muscles in common. The other half had flat correlograms and, in most of these pairs, the CM cells affected different muscles. The latter group included pairs of CM cells that facilitated synergistic muscles. These results indicate that common synaptic input specifically affects CM cells that have overlapping muscle fields. Reconstruction of the cortical locations of CM cells affecting 12 different muscles showed a wide and overlapping distribution of cortical colonies of forelimb muscles.