Interactions between natural and electrically evoked saccades. III. Is the nonstationarity the result of an integrator not instantaneously reset?

In the monkey, fixed-vector saccades evoked by superior colliculus (SC) stimulation when the animal fixates can be dramatically modified if the stimulation is applied during or immediately after an initial natural saccade. The vector is then deviated in the direction opposite to the displacement just accomplished as if it were compensating for part of the preceding trajectory. Recently, it was suggested that the amplitude of the compensatory deviation is related to the amplitude of the initial saccade linearly, and that the ratio between the two decreases exponentially as stimulation is applied later. These two findings (spatial linearity and temporal nonstationarity) were invoked as evidence for the noninstantaneous resetting of a feedback integrator. Such an integrator is included in most models of saccade generation for the specific purpose of terminating a saccade when it has reached its intended goal. However, the hypothesis of a feedback integrator in the process of being reset implies that the exponential decay of the compensatory deviation is temporally linked to the end of the initial saccade. We analyzed the time course of this decay in stimulation experiments performed at 24 SC sites in two monkeys. The results show that if the start of the exponential decay of compensation is assumed to be linked to the end of the initial saccade, then the relation between the amount of compensatory deviation and the amplitude of the initial saccade is not linear. On the other hand, it is possible to show a linear relation if the measurements of compensatory deviation are made in terms of delay of stimulation from the saccade beginning. We conclude that stimulating the SC just after a visually guided saccade does not seem to test the properties of a feedback integrator. Whether such an integrator is or is not resettable is not likely to be decided by this approach. Conversely, as the nonstationarity of compensation is linked to the beginning of the saccade, the nonstationarity seems to represent a property of an event occurring at saccade onset. We suggest that this event, close to the input of the oculomotor apparatus, is the summation of the visual signal with a damped signal of eye position or displacement.