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The Journal of Neurophysiology Vol. 81 No. 4 April 1999, pp. 1458-1468
Copyright ©1999 by the American Physiological Society
1Kawato Dynamic Brain Project,
Osu, Rieko and
Hiroaki Gomi.
Multijoint muscle regulation mechanisms examined by measured
human arm stiffness and EMG signals. Stiffness properties of the musculo-skeletal system can be controlled by regulating muscle activation and neural feedback gain. To understand the regulation of
multijoint stiffness, we examined the relationship between human arm
joint stiffness and muscle activation during static force control in
the horizontal plane by means of surface electromyographic (EMG)
studies. Subjects were asked to produce a specified force in a
specified direction without cocontraction or they were asked to keep
different cocontractions while producing or not producing an external
force. The stiffness components of shoulder, elbow, and their
cross-term and the EMG of six related muscles were measured during the
tasks. Assuming that the EMG reflects the corresponding muscle
stiffness, the joint stiffness was predicted from the EMG by using a
two-link six-muscle arm model and a constrained least-square-error regression method. Using the parameters estimated in this regression, single-joint stiffness (diagonal terms of the joint-stiffness matrix)
was decomposed successfully into biarticular and monoarticular muscle
components. Although biarticular muscles act on both shoulder and
elbow, they were found to covary strongly with elbow monoarticular muscles. The preferred force directions of biarticular muscles were
biased to the directions of elbow monoarticular muscles. Namely, the
elbow joint is regulated by the simultaneous activation of
monoarticular and biarticular muscles, whereas the shoulder joint is
regulated dominantly by monoarticular muscles. These results suggest
that biarticular muscles are innervated mainly to control the elbow
joint during static force-regulation tasks. In addition, muscle
regulation mechanisms for static force control tasks were found to be
quite different from those during movements previously reported. The
elbow single-joint stiffness was always higher than cross-joint
stiffness (off-diagonal terms of the matrix) in static tasks while
elbow single-joint stiffness is reported to be sometimes as small as
cross-joint stiffness during movement. That is, during movements, the
elbow monoarticular muscles were occasionally not activated when
biarticular muscles were activated. In static tasks, however,
monoarticular muscle components in single-joint stiffness were
increased considerably whenever biarticular muscle components in
single- and cross-joint stiffness increased. These observations suggest
that biarticular muscles are not simply coupled with the innervation of
elbow monoarticular muscles but also are regulated independently
according to the required task. During static force-regulation tasks,
covariation between biarticular and elbow monoarticular muscles may be
required to increase stability and/or controllability or to distribute
effort among the appropriate muscles.
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