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The Journal of Neurophysiology Vol. 87 No. 2 February 2002, pp. 679-695
Copyright ©2002 by the American Physiological Society
1Department of Bioengineering, 2Department of Physiology and Biophysics, and 3Regional Primate Research Center, University of Washington, Seattle, Washington 98195
Soetedjo, Robijanto,
Chris R. S. Kaneko, and
Albert F. Fuchs.
Evidence That the Superior Colliculus Participates in the
Feedback Control of Saccadic Eye Movements. J. Neurophysiol. 87: 679-695, 2002. There is general
agreement that saccades are guided to their targets by means of a motor
error signal, which is produced by a local feedback circuit that
calculates the difference between desired saccadic amplitude and an
internal copy of actual saccadic amplitude. Although the superior
colliculus (SC) is thought to provide the desired saccadic amplitude
signal, it is unclear whether the SC resides in the feedback loop. To
test this possibility, we injected muscimol into the brain stem region
containing omnipause neurons (OPNs) to slow saccades and then
determined whether the firing of neurons at different sites in the SC
was altered. In 14 experiments, we produced saccadic slowing while
simultaneously recording the activity of a single SC neuron. Eleven of
the 14 neurons were saccade-related burst neurons (SRBNs), which
discharged their most vigorous burst for saccades with an optimal
amplitude and direction (optimal vector). The optimal directions for
the 11 SRBNs ranged from nearly horizontal to nearly vertical, with optimal amplitudes between 4 and 17°. Although muscimol injections into the OPN region produced little change in the optimal vector, they
did increase mean saccade duration by 25 to 192.8% and decrease mean
saccade peak velocity by 20.5 to 69.8%. For optimal vector saccades,
both the acceleration and deceleration phases increased in duration.
However, during 10 of 14 experiments, the duration of deceleration
increased as fast as or faster than that of acceleration as saccade
duration increased, indicating that most of the increase in duration
occurred during the deceleration phase. SRBNs in the SC changed their
burst duration and firing rate concomitantly with changes in
saccadic duration and velocity, respectively. All SRBNs showed a robust
increase in burst duration as saccadic duration increased. Five of 11 SRBNs also exhibited a decrease in burst peak firing rate as saccadic
velocity decreased. On average across the neurons, the number of spikes
in the burst was constant. There was no consistent change in the
discharge of the three SC neurons that did not exhibit bursts with
saccades. Our data show that the SC receives feedback from downstream
saccade-related neurons about the ongoing saccades. However, the
changes in SC firing produced in our study do not suggest that the
feedback is involved with producing motor error. Instead, the feedback seems to be involved with regulating the duration of the discharge of
SRBNs so that the desired saccadic amplitude signal remains present
throughout the saccade.
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