SELECTIVE OUTPUT-DISCRIMINATIVE SIGNALS IN THE MOTOR CORTEX OF WAKING MONKEYS

Monkeys and humans have similar capacities to discriminate between the frequencies of mechanical sinusoids delivered to the glabrous skin of their hands. Combined psychophysical-electrophysiological experiments in monkeys discriminating in the range of flutter provided evidence that this capacity depends upon differences in the cycle lengths in the sets of periodically entrained activity, evoked by the stimuli discriminated, in neurons of areas 3b and 1 of the (sensory) hemisphere opposite the stimulated hand. Identical experiments have now been made, in similarly trained and discriminating monkeys, in the motor cortex (area 4) of the hemisphere opposite the arm projecting selectively to one of two targets, to indicate discrimination (five hemispheres, 1137 neurons studied). We observed a selective signal of the upcoming correct discrimination in about 25% of the neurons of area 4 active in the task. The neuronal discharge occurs selectively for stimuli either lower or higher in frequency than that of the base stimulus, and commonly begins within 200-300 msec after onset of the comparison stimulus. These neuronal discharges are aperiodic, with no sign of the stimulus frequencies. EMG recording during performance of the discrimination showed that the muscles of the arm opposite the side of recording were silent during the period of stimulus presentations. Recordings during trials in which the animal made errors showed most commonly that the output of the discrimination operation was itself in error, followed by an appropriate arm projection to the wrong target. We interpret the selective response during the comparison stimulus to be a postdiscrimination signal projected transcallosally from the sensory hemisphere to the motor area of the hemisphere controlling the responding arm. We obtained no evidence that the discrimination operation is localized to any particular area, and we surmise it to occur in the dynamic activity within the distributed system linking the sensory cortex of one hemisphere and the motor cortex of the other. One-third of the neurons of the motor cortex responded to indentation of the skin of the ipsilateral hand, at trial onset. These responses varied from those closely linked to that sensory stimulus to those linked to the upcoming movement of the contralateral hand. These onset responses did not occur when similar sequences of mechanical stimuli were delivered to alert but idling monkeys.