Cerebellum-dependent motor learning: lessons from adaptation of eye movements in primates

In order to ameliorate the consequences of ego motion for vision, human and nonhuman observers generate reflexive, compensatory eye movements based on visual as well as vestibular information, helping to stabilize the images of visual scenes on the retina despite ego motion. And in order to fully exploit the advantages of foveal vision, they make saccades to shift the image of an object onto the fovea and smooth pursuit eye movements to stabilize it there despite continuing object movement relative to the observer. With the exception of slow visually driven eye movements, which can be understood as manifestations of relatively straightforward feedback systems, most eye movements require a direct conversion of sensory input into appropriate motor responses in the absence of immediate sensory feedback. Hence, in order to generate appropriate oculomotor responses, the parameters linking input and output must be chosen suitably. Moreover, as the parameters may change from one manifestation of a movement to the next, for instance because of oculomotor fatigue, the choices should also be quickly modifiable. This chapter will present evidence showing that this fast parametric optimization, understood as a functionally distinct example of motor learning, is an accomplishment of specific parts of the cerebellum devoted to the control of eye movements. It will also discuss recent electrophysiological results suggesting how this specific form of motor learning may emerge from information processing in cerebellar circuits.