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The Journal of Neurophysiology Vol. 84 No. 2 August 2000, pp. 876-891
Copyright ©2000 by the American Physiological Society
Laboratory of Sensorimotor Research, National Eye Institute, Bethesda, Maryland 20892
Krauzlis, Richard J.,
Michele A. Basso, and
Robert H. Wurtz.
Discharge Properties of Neurons in the Rostral Superior
Colliculus of the Monkey During Smooth-Pursuit Eye Movements. J. Neurophysiol. 84: 876-891, 2000. The
intermediate and deep layers of the monkey superior colliculus (SC)
comprise a retinotopically organized map for eye movements. The rostral
end of this map, corresponding to the representation of the fovea,
contains neurons that have been referred to as "fixation cells"
because they discharge tonically during active fixation and pause
during the generation of most saccades. These neurons also possess
movement fields and are most active for targets close to the fixation
point. Because the parafoveal locations encoded by these neurons are
also important for guiding pursuit eye movements, we studied these
neurons in two monkeys as they generated smooth pursuit. We found that
fixation cells exhibit the same directional preferences during pursuit
as during small saccades
they increase their discharge during
movements toward the contralateral side and decrease their discharge
during movements toward the ipsilateral side. This pursuit-related
activity could be observed during saccade-free pursuit and was not
predictive of small saccades that often accompanied pursuit. When we
plotted the discharge rate from individual neurons during pursuit as a
function of the position error associated with the moving target, we
found tuning curves with peaks within a few degrees contralateral of
the fovea. We compared these pursuit-related tuning curves from each
neuron to the tuning curves for a saccade task from which we separately
measured the visual, delay, and peri-saccadic activity. We found the
highest and most consistent correlation with the delay activity
recorded while the monkey viewed parafoveal stimuli during fixation.
The directional preferences exhibited during pursuit can therefore be
attributed to the tuning of these neurons for contralateral locations
near the fovea. These results support the idea that fixation cells are
the rostral extension of the buildup neurons found in the more caudal
colliculus and that their activity conveys information about the size
of the mismatch between a parafoveal stimulus and the currently
foveated location. Because the generation of pursuit requires a break
from fixation, the pursuit-related activity indicates that these
neurons are not strictly involved with maintaining fixation.
Conversely, because activity during the delay period was found for many
neurons even when no eye movement was made, these neurons are also not obligatorily related to the generation of a movement. Thus the tonic
activity of these rostral neurons provides a potential position-error signal rather than a motor commanda principle that may be applicable to buildup neurons elsewhere in the SC.
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