Properties of saccade-related neurons in the cat superior colliculus: patterns of movement fields and discharge timing

The discharge characteristics of saccade-related neurons in the superior colliculi (SC) of trained cats were investigated in order to understand the involvement of the SC in dynamic control of saccades. In particular, two aspects of saccade-related collicular activity in relation to visually guided saccades were quantitatively analyzed: the bounded pattern of amplitude tuning and the timing of neural activity in relation to saccade offset. In order to determine whether the amplitude tuning of SC cells is bounded or unbounded, we characterized it with a quantitative index, the unbounded-field (UF) index. Analysis of the characteristics of the distribution of this index, including the absence of unbounded fields for cells within the central 10 degrees, suggested that all cells form a single class and that apparent unbounded amplitude tuning is an artifact occurring when the range of gaze shift is truncated by restraining the head. This interpretation is supported by the results of a computer simulation based on truncation of gaze-shift range that closely replicated the experimental data. Analyses of relative location of peak motor activity in the spatial and temporal dimensions revealed that the peak discharge was more closely linked to the midpoint of the saccade than to the offset of saccade, regardless of the pattern of amplitude tuning. These results do not support the proposal advanced by the moving-hill hypothesis: the bounded pattern of amplitude tuning is inconsistent with rostral migration of the activity locus during execution of a saccade, and the relative location of the peak motor activity is inconsistent with the idea that the SC activity encodes dynamic motor error (the difference between desired and current gaze directions).