Encoding of target direction and speed during visual instruction and arm tracking in dorsal premotor and primary motor cortical neurons

The encoding of direction and speed in the discharge of dorsal premotor (PMd) and primary motor (MI) neurons was studied during two-dimensional visually-instructed pursuit arm movements in which eight directions and four constant speeds were independently manipulated. Each trial consisted of equal durations of visual observation of target movement without hand movement (cue) and visual pursuit-tracking of the target with the hand (track). A total of 240 neurons was recorded from PMd and MI in two Macaca mulatta monkeys. Two classes of regression analyses were used to relate neuronal firing during the cue and track periods to direction and speed. First, the average firing from each period was fitted to target direction or speed. Period-averaged firing significantly correlated with direction more frequently in the track than in the cue period. Conversely, correlations with speed (with or without direction) were more common in the cue than in the track period. Secondly, a binwise regression evaluated the temporal evolution of firing correlations with direction and speed. Supporting the period-based results, significant binwise correlations of the discharge with speed occurred preferentially during the cue period when there was no hand movement. Prior to movement, correlations of the firing with direction became significant and continued through the movement. Both analyses demonstrated a distinct tendency for neurons to be modulated by speed information early and by direction information later. This temporal parcellation reflects both the sequential demands of the task and constraints placed on the neural computations. The early representation of target speed is hypothesized to reflect the need to calculate a 'go signal' for the initiation of movement.