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The Journal of Neurophysiology Vol. 82 No. 2 August 1999, pp. 999-1018
Copyright ©1999 by the American Physiological Society
1Laboratory of Sensorimotor Research, National Eye Institute, Bethesda, Maryland 20892-4435; 2Dipartimento di Elettronica, Elettrotecnica ed Informatica, Università degli Studi di Trieste, 34100 Trieste, Italy; 3Laboratory of Neurophysiology and Center for Systems Engineering and Applied Mechanics, Universitè Catholique de Louvain, B-1348 Louvain-La-Neuve, Belgium
Quaia, Christian,
Philippe Lefèvre, and
Lance M. Optican.
Model of the Control of Saccades by Superior Colliculus and
Cerebellum. J. Neurophysiol. 82: 999-1018, 1999. Experimental evidence indicates that the superior colliculus
(SC) is important but neither necessary nor sufficient to produce accurate saccadic eye movements. Furthermore both clinical and experimental evidence points to the cerebellum as an indispensable component of the saccadic system. Accordingly, we have devised a new
model of the saccadic system in which the characteristics of saccades
are determined by the cooperation of two pathways, one through the SC
and the other through the cerebellum. Both pathways are influenced by
feedback information: the feedback determines the decay of activity for
collicular neurons and the timing of the activation for cerebellar
neurons. We have modeled three types of cells (burst, buildup, and
fixation neurons) found in the intermediate layers of the superior
colliculus. We propose that, from the point of view of motor execution,
the burst neurons and the buildup neurons are not functionally distinct
with both providing a directional drive to the brain stem circuitry.
The fixation neurons determine the onset of the saccade by
disfacilitating the omnipause neurons in the brain stem. Excluding
noise-related variations, the ratio of the horizontal to the vertical
components of the collicular drive is fixed throughout the saccade
(i.e., its direction is fixed); the duration of the drive is such that it always would produce hypermetric movements. The cerebellum plays
three roles: first, it provides an additional directional drive, which
improves the acceleration of the eyes; second, it keeps track of the
progress of the saccade toward the target; and third, it ends the
saccade by choking off the collicular drive. The drive provided by the
cerebellum can be adjusted in direction to exert a directional control
over the saccadic trajectory. We propose here a control mechanism that
incorporates a spatial displacement integrator in the cerebellum; under
such conditions, we show that a partial directional control arises
automatically. Our scheme preserves the advantages of several previous
models of the saccadic system (e.g., the lack of a spatial-to-temporal
transformation between the SC and the brain stem; the use of efference
copy feedback to control the saccade), without incurring many of their
drawbacks, and it accounts for a large amount of experimental data.
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