The Saccadic System

Saccades are fast conjugate eye movements that move both eyes quickly in the same direction, so that the image of an object of interest is brought on the foveae. Saccades can be made not only toward visual targets, but also toward auditory and tactile stimuli, as well as toward memorized targets. Saccades can be generated reflexively, and they are responsible for resetting the eyes back to the mid-orbital position during vestibulo-ocular or optokinetic stimulation. Saccades need to be fast to get the eyes on the target as soon as possible. They also need to be fast because our eyes act like cameras with slow shutters—vision is so blurred during saccades that the eyes have to move quickly to minimize the time during which no clear image is captured on the foveae. Indeed, saccades are the fastest type of eye movements, and they are among the fastest movements that the body can make. Saccade speed is not under voluntary control but depends on the size of the movement, with larger saccades attaining higher peak velocities. It has been estimated that we make more than 100,000 saccades per day. The burst neuron circuits in the brainstem provide the necessary motor signals to the extraocular muscles for the generation of saccades. There is a division of labor between the pons and the midbrain, with the pons primarily involved in generating horizontal saccades and the midbrain primarily involved in generating vertical and torsional saccades. However, because eye movements are a component of cognitive and purposeful behaviors in higher mammals, the process of deciding when and where to make a saccade occurs in the cerebral cortex. Not only does the cortex exert control over saccades through direct projections to the burst neuron circuits, it also acts via the superior colliculus. The superior colliculus is located in the midbrain and consists of seven layers: three superficial layers and four intermediate/ deep layers. The three superficial layers receive direct inputs from both the retina and striate cortex, and they contain a retinotopic representation of the contralateral visual hemifield.