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The Journal of Neurophysiology Vol. 86 No. 5 November 2001, pp. 2527-2542
Copyright ©2001 by the American Physiological Society
Howard Hughes Medical Institute and Department of Neurobiology, Stanford University School of Medicine, Stanford, California 94305
Horwitz, Gregory D. and
William T. Newsome.
Target Selection for Saccadic Eye Movements: Direction-Selective
Visual Responses in the Superior Colliculus. J. Neurophysiol. 86: 2527-2542, 2001. We investigated the role
of the superior colliculus (SC) in saccade target selection in rhesus
monkeys who were trained to perform a direction-discrimination task. In
this task, the monkey discriminated between opposed directions of
visual motion and indicated its judgment by making a saccadic eye
movement to one of two visual targets that were spatially aligned with
the two possible directions of motion in the display. Thus the neural circuits that implement target selection in this task are likely to
receive directionally selective visual inputs and be closely linked to
the saccadic system. We therefore studied prelude neurons in the
intermediate and deep layers of the SC that can discharge up to several
seconds before an impending saccade, indicating a relatively high-level
role in saccade planning. We used the direction-discrimination task to
identify neurons whose prelude activity "predicted" the impending
perceptual report several seconds before the animal actually executed
the operant eye movement; these "choice predicting" cells comprised
~30% of the neurons we encountered in the intermediate and deep
layers of the SC. Surprisingly, about half of these prelude cells
yielded direction-selective responses to our motion stimulus during a
passive fixation task. In general, these neurons responded to motion
stimuli in many locations around the visual field including the center
of gaze where the visual discriminanda were positioned during the
direction-discrimination task. Preferred directions generally pointed
toward the location of the movement field of the SC neuron in
accordance with the sensorimotor demands of the discrimination task.
Control experiments indicate that the directional responses do not
simply reflect covertly planned saccades. Our results indicate that a
small population of SC prelude neurons exhibits properties appropriate
for linking stimulus cues to saccade target selection in the context of
a visual discrimination task.
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