The role of the frontal pursuit area in learning in smooth pursuit eye movements

The frontal pursuit area (FPA) in the cerebral cortex is part of the circuit for smooth pursuit eye movements. The present paper asks whether the FPA is upstream, downstream, or at the site of learning in pursuit eye movements. Learning was induced by having monkeys repeatedly pursue targets that moved at one speed for 150 msec before changing speed. Single-cell recording showed no consistent correlate of pursuit learning in the responses of FPA neurons. Some neurons showed changes in firing in the same direction as the learning, others showed changes in the opposite direction, and many showed no changes at all. In contrast, the eye movements evoked by electrical stimulation of the FPA showed clear correlates of learning. Learning effects were observed when microstimulation was delivered during the initiation of pursuit and during fixation of a stationary target. In addition, learning caused changes in the degree to which stimulation of the FPA enhanced the eye velocity evoked by brief perturbations of a stationary target. The magnitude of the change in the stimulation-evoked eye movement in each tracking condition was proportional to the size of the eye movement evoked under that condition before learning. We conclude that learning occurs downstream from the FPA, possibly within the cerebellum, and that learning may be related to mechanisms that also control the gain of visual-motor responses on a rapid time scale.