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The Journal of Neurophysiology Vol. 83 No. 3 March 2000, pp. 1283-1299
Copyright ©2000 by the American Physiological Society
Allgemeine Zoologie und Neurobiologie, Ruhr-Universität, 44780 Bochum, Germany
Stuphorn, Veit,
Erhard Bauswein, and
Klaus-Peter Hoffmann.
Neurons in the Primate Superior Colliculus Coding for Arm
Movements in Gaze-Related Coordinates. J. Neurophysiol. 83: 1283-1299, 2000. In the intermediate and deep layers
of the superior colliculus (SC), a well-established oculomotor
structure, a substantial population of cells is involved in the control
of arm movements. To examine the reference frame of these neurons, we
recorded in two rhesus monkeys (Macaca mulatta) the
discharges of 331 neurons in the SC and the underlying mesencephalic
reticular formation (MRF) while monkeys reached to the same target
location during different gaze orientations. For 65 reach-related cells
with sufficient data and for simultaneously recorded electromyograms
(EMGs) of 11 arm muscles, we calculated an ANOVA (factors: target
position, gaze angle) and a gaze-dependency (GD) index. EMGs and the
activity of many (60%) of the reach-related neurons were not
influenced by the target representation on the retina or eye position.
We refer to these as "gaze-independent" reach neurons. For 40%,
however, the GD fell outside the range of the muscle modulation, and
the ANOVA showed a significant influence of gaze. These
"gaze-related" reach neurons discharge only when the monkey reaches
for targets having specific coordinates in relation to the gaze axis,
i.e., for targets in a gaze-related "reach movement field" (RMF).
Neuronal activity was not modulated by the specific path of the arm
movement, the muscle pattern that is necessary for its realization or
the arm that was used for the reach. In each SC we found gaze-related neurons with RMFs both in the contralateral and in the ipsilateral hemifield. The topographical organization of the gaze-related reach
neurons in the SC could not be matched with the well-known visual and
oculomotor maps. Gaze-related neurons were more modulated in their
strength of activity with different directions of arm movements than
were gaze-independent reach neurons. Gaze-related reach neurons were
recorded at a median depth of 2.03 mm below SC surface in the
intermediate layers, where they overlap with saccade-related burst
neurons (median depth: 1.55 mm). Most of the gaze-independent reach
cells were found in a median depth of 4.01 mm below the SC surface in
the deep layers and in the underlying MRF. The gaze-related reach
neurons operating in a gaze-centered coordinate system could signal
either the desired target position with respect to gaze direction or
the motor error between gaze axis and reach target. The
gaze-independent reach neurons, possibly operating in a shoulder- or
arm-centered reference frame, might carry signals closer to motor
output. Together these two types of reach neurons add evidence to our
hypothesis that the SC is involved in the sensorimotor transformation
for eye-hand coordination in primates.
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