Experimental test of two models for the generation of oblique saccades.

In this paper, we report a detailed study of the dynamic properties of horizontal, vertical and oblique saccades. These eye movements were measured with an improved version of the double-magnetic induction method in two rhesus monkeys. We found that onsets of orthogonal components of oblique saccades are so well synchronized in the monkey that a common initiation system seems likely. Saccade vectors obeyed a nonlinear peak-velocity/amplitude relationship in all directions. The peak-velocity/duration/amplitude relationship for components was not fixed, but depended on the relative size of the orthogonal component: for a component with a given size, its duration increased and its peak velocity decreased, as the saccade vector to which it contributed turned away from the component direction under consideration. This stretching effect, which reflects a nonlinearity in the system, was negligible for small saccade vectors but became very pronounced in large oblique saccades. These experimental data were confronted with quantitative predictions derived from two different models for the generation of saccades in two dimensions. It appears that a model which assumes the existence of synchronized, but otherwise independent, pulse generators for horizontal and vertical components must be rejected. An alternative model, featuring a nonlinear vectorial pulse generator followed by a decomposition stage which generates component velocity command signals from the vectorial eye velocity signal, provides good fit with the data. According to this common-source model, the two nonlinear phenomena observed, viz., the curvilinear peak-velocity/amplitude relationship of saccades in all directions and component stretching in large oblique saccades, are due to a single nonlinearity in the proposed vectorial pulse generator. A possible neural basis for the common-source model is discussed.