234 Cranial Nerves

counter-rotate smoothly and then snap back to a neutral position in the orbit
and begin a new smooth pursuit movement in the same direction as the
original one. This alternating smooth pursuit-saccadic movement is called
vestibular nystagmus. The VOR adapts rapidly. When head rotation is
sustained, the fluid in the semicircular canals accelerates to match the speed of
the head, and the sensory signal degrades. In these conditions, a second system
of maintaining visual fixation, the optokinetic reflex, becomes important.

Optokinetic Reflex
The OKR is similar to the vestibulo-ocular reflex in that it generates a smooth
pursuit movement that is equal in velocity but opposite in direction to the
movement of the head. A smooth pursuit movement tracks the passing visual
field for a distance and then a saccade brings the eyes back to a neutral
position, and a new smooth pursuit movement is generated to follow the visual
field again. This oscillating eye movement is called optokinetic nystagmus
(OKN). A familiar example of optokinetic nystagmus is the movements of the
eyes as they follow the passing scenery from a moving vehicle.
The sensory signals that drive the OKR arise in the retina. Because
visual processing in the retina is slow relative to signal transduction in the
semicircular canals, the OKR responds slowly to changes in the movement of
the visual fields across the retina. However, unlike the VOR, it does not adapt,
which means that the sensory signals that drive the reflex do not degrade with
time. Retinal ganglion cell axons enter the optic nerve and project caudally to
the pretectal region of the midbrain, which also receives signals from the visual
association areas of the occipital lobes. In turn, the pretectal region projects to
the vestibular nucleus (mainly the medial subnucleus) in the medulla by
presently unknown pathways. The medial subnucleus projects to the lower
motor neurons in the abducens, trochlear, and oculomotor nuclei and also to the
internuclear neurons in the abducens nucleus (Figure 13-10).

Pursuit System
The pursuit system generates the eye movements involved in following a
moving object against a stationary background (ie, following a butterfly in a
garden). Since this system acts to keep moving images centered on the fovea, it
follows that only animals with a fovea have a smooth pursuit system.
Signals for voluntary pursuit arise in the extrastriate visual cortex of the
temporal lobe. The precise route by which these signals reach the lower motor
neurons is not known with certainty. However, it seems likely to follow a
progression where the extrastriate cortex signals the dorsolateral pontine nuclei
in the pons, which, in turn, signal the flocculus and posterior vermis of the
cerebellum, which signal the vestibular nucleus (mainly the medial subnucleus)
that projects via the MLF to the nuclei of cranial nerves III, IV, and VI.