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The Journal of Neurophysiology Vol. 84 No. 5 November 2000, pp. 2605-2621
Copyright ©2000 by the American Physiological Society
Section of Neurobiology, Yale University School of Medicine, New Haven, Connecticut 06520-8001
Russo, Gary S. and
Charles J. Bruce.
Supplementary Eye Field: Representation of Saccades and
Relationship Between Neural Response Fields and Elicited Eye
Movements. J. Neurophysiol. 84: 2605-2621, 2000. The
functional organization of the low-threshold supplementary eye field
(SEF) was studied by analyzing presaccadic activity, electrically
elicited saccades, and the relationship between them. Response-field
optimal vectors, defined as the visual field coordinates or saccadic
eye-movement dimensions evoking the highest neural discharge, were
quantitatively estimated for 160 SEF neurons by systematically varying
peripheral target location relative to a central fixation point and
then fitting the responses to Gaussian functions. Saccades were
electrically elicited at 109 SEF sites by microstimulation (70 ms,
10-100 µA) during central fixation. The distribution of response
fields and elicited saccades indicated a complete representation of all
contralateral saccades in SEF. Elicited saccade polar directions ranged
between 97 and 262° (data from left hemispheres were transformed to a
right-hemisphere convention), and amplitudes ranged between 1.8 and
26.9°. Response-field optimal vectors (right hemisphere transformed)
were nearly all contralateral as well; the directions of 115/119 visual
response fields and 80/84 movement response fields ranged between 90 and 279°, and response-field eccentricities ranged between 5 and
50°. Response-field directions for the visual and movement activity
of visuomovement neurons were strongly correlated (r = 0.95). When neural activity and elicited saccades obtained at exactly
the same sites were compared, response fields were highly predictive of
elicited saccade dimensions. Response-field direction was highly
correlated with the direction of saccades elicited at the recording
site (r = 0.92, n = 77). Similarly,
response-field eccentricity predicted the size of subsequent
electrically elicited saccades (r = 0.49, n = 60). However, elicited saccades were generally
smaller than response-field eccentricities and consistently more
horizontal when response fields were nearly vertical. The polar
direction of response fields and elicited saccades remained constant
perpendicular to the cortical surface, indicating a columnar
organization of saccade direction. Saccade direction progressively
shifted across SEF; however, these orderly shifts were more indicative
of a hypercolumnar organization rather than a single global topography.
No systematic organization for saccade amplitude was evident. We
conclude that saccades are represented in SEF by congruent visual
receptive fields, presaccadic movement fields, and efferent mappings.
Thus SEF specifies saccade vectors as bursts of
activity by local groups of neurons with appropriate projections to
downstream oculomotor structures. In this respect, SEF is organized
like the superior colliculus and the frontal eye field even though SEF
lacks an overall global saccade topography. We contend that all
specialized oculomotor functions of SEF must operate within the context
of this fundamental organization.
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