Neuronal specification of direction and distance during reaching movements in the superior precentral premotor area and primary motor cortex of monkeys

Q. G. Fu, J. I. Suarez and T. J. Ebner Department of Neurosurgery, University of Minnesota, Minneapolis 55455. 1. Single-unit neuronal activity was recorded in the primary motor and superior precentral premotor areas of two rhesus monkeys during an arm reaching task. The task involved moving a cursor displayed on a video terminal using a draftsman's arm-type manipulandum. From a centrally located start box the animal was required to move to 1 of 48 target boxes at eight different directions (0-360 degrees in 45 degrees intervals) and six distances (1.4-5.4 cm in 0.8-cm increments). Both direction and distance for the upcoming movement were unpredictable. 2. The activity of 197 arm movement-related cells was recorded and evaluated for each of the 48 targets. Histological examination showed the cells to be primarily in the primary motor cortex or in the premotor area around the superior precentral sulcus. Each cell's discharge was aligned on movement onset and averaged over five trials for each target. Movement kinematics including hand path velocity were also determined. The task time was divided into three epochs, a premovement period (PT), a movement period (MT), and total time (TT = PT+MT). For each epoch the average firing was correlated with the direction and distance of the movement using various regression procedures. 3. An analysis of variance (ANOVA) showed that the majority of neurons were modulated significantly by movement direction in each of the three time periods, PT (73.7%), MT (68.3%), and TT (78.5%). The relationship of the firing to direction was fit to a cosine tuning function for each significantly modulated cell. In 86.3% of the cells the firing was correlated significantly with a cosine function of movement direction in TT. A cell's preferred direction varied little for different movement distances. The mean difference in preferred direction for the smallest possible change in distance (0.8 cm) was 12.8 +/- 11.4 degrees (SD) and 17.1 +/- 14.7 degrees for the largest change in distance (4.0 cm). 4. Correlation analysis revealed that the activity of the majority of cells was modulated significantly by distance along at least one direction in each of the three time periods, PT (46.8%), MT (68.8%), and TT (67.7%). Subsequently, a univariate linear regression model was used to quantify a cell's discharge as a function of distance. For the regressions of firing with distance with a statistically significant correlation (r > 0.8), the mean s