Hwang EJ and Shadmehr R (2005) Internal models of limb dynamics
and the encoding of limb state, Journal of Neural Engineering
2:S266-S278.
Abstract Studies of reaching suggest that
humans adapt to novel arm dynamics by building internal models that transform
planned sensory states of the limb, e.g., desired limb position and its
derivatives, into motor commands, e.g., joint torques. Earlier work modeled
this computation via a population of basis elements and used system
identification techniques to estimate the tuning properties of the bases from
the patterns of generalization. Here we hypothesized that the neural
representation of planned sensory states in the internal model might resemble
the signals from the peripheral sensors. These sensors normally encode the
limb's actual sensory state in which movement errors occurred. We developed a
set of equations based on properties of muscle spindles that estimated spindle
discharge as a function of the limb's state during reaching and drawing of
circles. We then implemented a simulation of a two-link arm that learned to
move in various force fields using these spindle-like bases. The system
produced a pattern of adaptation and generalization that accounted for a wide
range of previously reported behavioral results. In particular, the bases
showed gain-field interactions between encoding of limb position and velocity,
very similar to the gain fields inferred from behavioral studies. The poor
sensitivity of the bases to limb acceleration predicted behavioral results that
were confirmed by experiment. We suggest that the internal model of limb
dynamics is computed by the brain with neurons that encode the state of the
limb in a manner similar to that expected of muscle spindle afferents.
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