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The Journal of Neurophysiology Vol. 84 No. 5 November 2000, pp. 2302-2316
Copyright ©2000 by the American Physiological Society
1Medical Research Council Group for Action and Perception, Centre for Vision Research and Departments of Psychology and Biology, York University, Toronto, Ontario M3J 1P3, Canada; and 2Department of Medical Physics and Biophysics, University of Nijmegen, NL 6525 EZ Nijmegen, The Netherlands
Medendorp, W. P.,
J. D. Crawford,
D.Y.P. Henriques,
J.A.M. Van Gisbergen, and
C.C.A.M. Gielen.
Kinematic Strategies for Upper Arm-Forearm Coordination in
Three Dimensions. J. Neurophysiol. 84: 2302-2316, 2000. This study addressed the question of how the
three-dimensional (3-D) control strategy for the upper arm depends on
what the forearm is doing. Subjects were instructed to point a
laser
attached in line with the upper arm
toward various visual
targets, such that two-dimensional (2-D) pointing directions of the
upper arm were held constant across different tasks. For each such
task, subjects maintained one of several static upper arm-forearm
configurations, i.e., each with a set elbow angle and forearm
orientation. Upper arm, forearm, and eye orientations were measured
with the use of 3-D search coils. The results confirmed that Donders'
law (a behavioral restriction of 3-D orientation vectors to a 2-D
"surface") does not hold across all pointing tasks,
i.e., for a given pointing target, upper arm torsion varied widely.
However, for any one static elbow configuration, torsional variance was
considerably reduced and was independent of previous arm position,
resulting in a thin, Donders-like surface of orientation vectors. More
importantly, the shape of this surface (which describes
upper arm torsion as a function of its 2-D pointing direction) depended
on both elbow angle and forearm orientation. For pointing with the arm
fully extended or with the elbow flexed in the horizontal plane, a
Listing's-law-like strategy was observed, minimizing shoulder
rotations to and from center at the cost of position-dependent tilts in
the forearm. In contrast, when the arm was bent in the vertical plane,
the surface of best fit showed a Fick-like twist that
increased continuously as a function of static elbow flexion, thereby
reducing position-dependent tilts of the forearm with respect to
gravity. In each case, the torsional variance from these surfaces
remained constant, suggesting that Donders' law was obeyed equally
well for each task condition. Further experiments established that
these kinematic rules were independent of gaze direction and eye
orientation, suggesting that Donders' law of the arm does not
coordinate with Listing's law for the eye. These results revive the
idea that Donders' law is an important governing principle for the
control of arm movements but also suggest that its various forms may
only be limited manifestations of a more general set of
context-dependent kinematic rules. We propose that these rules are
implemented by neural velocity commands arising as a function of
initial arm orientation and desired pointing direction, calculated such
that the torsional orientation of the upper arm is implicitly
coordinated with desired forearm posture.
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