236 Cranial Nerves

Saccadic System
The saccadic system shifts the fovea rapidly from one target to another, moving
the eyes at speeds of up to 900 degrees per second. The high speed of saccadic
movement minimizes the time that the fovea is off target. Signals for saccadic
movement arise in the superior colliculi in the midbrain. The superior colliculi
receive saccade-generating signals from four major sources: the frontal eye fields
of the frontal lobes, which are involved in consciously selecting an object of
interest; the retinae; the somatosensory system; and the auditory system (Figure
13-11).
The saccades generated by the frontal eye fields are the only totally
voluntary eye movements that we make. The saccades generated in response to
signals from the retinae, the somatosensory system, and the auditory system are
visual reflexes. For example, we reflexively orient our eyes to a sudden flash of
light, a loud sound, or an area of the body that is subjected to unexpected
(usually noxious) sensory input.

Gaze Centers
The frontal eye fields and the superior colliculi project to "gaze centers" in the
retic-ular formation of the brain stem. Two gaze centers have been identified: one
for vertical gaze (the rostral interstitial nucleus of the MLF) and one for lateral
gaze (the paramedian pontine reticular formation). These reticular nuclei
project to the nuclei of nerves III, IV, and VI, which, in turn, generate saccades
that redirect the fovea to an appropriate location. They are also responsible for
the saccadic phase of the optokinetic reflex and vestibulo-ocular nystagmus.

Vergence System
The four eye movement systems described previously produce conjugate move -
ments (ie, both eyes move in the same direction by the same amount). In
contrast, the vergence system moves the eyes in opposing directions, usually by
differing amounts. The vergence system is present only in animals with
binocular vision and is part of the accommodation reflex (see Chapter III).
When attention is shifted from one object to another object that is closer
or further away, the eyes converge or diverge until the image occupies the same
relative location on both retinae and only one image is perceived. For the eyes to
converge (see Chapter III, Figure III-8), both medial rectus muscles are
activated and both lateral rectus muscles are inhibited. For divergence, the
opposite set of muscle actions is necessary. Little is known about the precise
location of a "vergence" center in humans. However, it is likely in the
midbrain, close to the oculomotor nucleus.
The vergence system works in conjunction with the smooth pursuit and saccadic
systems. The sensory stimulus for vergence is disparity between the relative
locations of the images on both retinae. This disparity is detected by extrastriate
visual cortex neurons, which presumably project to the vergence