Cortico-cortical networks and cortico-subcortical loops for the higher control of eye movements.

There are multiple distinct regions, or eye fields, in the cerebral cortex that contribute directly to the initiation and control of voluntary eye movements. We concentrate on six of these: the frontal eye field, parietal eye field, supplementary eye field, middle superior temporal area, prefrontal eye field, and area 7 m (precuneus in humans). In each of these regions: (1) there is neural activity closely related to eye movements; (2) electrical microstimulation produces or modifies eye movements; (3) surgical lesions or chemical inactivation impairs eye movements; (4) there are direct neural projections to major structures in the brainstem oculomotor system; and (5) increased activity is observed during eye movement tasks in functional magnetic resonance imaging or positron emission tomography experiments in humans. Each of these eye fields is reciprocally connected with the other eye fields, and each receives visual information directly from visual association cortex. Each eye field has distinct subregions that are concerned with either saccadic or pursuit eye movements. The saccadic subregions are preferentially interconnected with other saccade subregions and the pursuit subregions are preferentially interconnected with other pursuit subregions. Current evidence strongly supports the proposal that there are parallel cortico-cortical networks that control purposeful saccadic and pursuit eye movements, and that the activity in those networks is modulated by feedback information, via the thalamus, from the superior colliculus, basal ganglia, and cerebellum.