R. Shadmehr et Fa. Mussaivaldi, ADAPTIVE REPRESENTATION OF DYNAMICS DURING LEARNING OF A MOTOR TASK, The Journal of neuroscience, 14(5), 1994, pp. 3208-3224
We investigated how the CNS learns to control movements in different d
ynamical conditions, and how this learned behavior is represented. In
particular, we considered the task of making reaching movements in the
presence of externally imposed forces from a mechanical environment.
This environment was a force field produced by a robot manipulandum, a
nd the subjects made reaching movements while holding the end-effector
of this manipulandum. Since the force field significantly changed the
dynamics of the task, subjects' initial movements in the force field
were grossly distorted compared to their movements in free space. Howe
ver, with practice, hand trajectories in the force field converged to
a path very similar to that observed in free space. This indicated tha
t for reaching movements, there was a kinematic plan independent of dy
namical conditions. The recovery of performance within the changed mec
hanical environment is motor adaptation. In order to investigate the m
echanism underlying this adaptation, we considered the response to the
sudden removal of the field after a training phase. The resulting tra
jectories, named aftereffects, were approximately mirror images of tho
se that were observed when the subjects were initially exposed to the
field. This suggested that the motor controller was gradually composin
g a model of the force field, a model that the nervous system used to
predict and compensate for the forces imposed by the environment. In o
rder to explore the structure of the model, we investigated whether ad
aptation to a force field, as presented in a small region, led to afte
reffects in other regions of the workspace. We found that indeed there
were aftereffects in workspace regions where no exposure to the field
had taken place; that is, there was transfer beyond the boundary of t
he training data. This observation rules out the hypothesis that the s
ubject's model of the force field was constructed as a narrow associat
ion between visited states and experienced forces; that is, adaptation
was not via composition of a look-up table. In contrast, subjects mod
eled the force field by a combination of computational elements whose
output was broadly tuned across the motor state space. These elements
formed a model that extrapolated to outside the training region in a c
oordinate system similar to that of the joints and muscles rather than
end-point forces. This geometric property suggests that the elements
of the adaptive process represent dynamics of a motor task in terms of
the intrinsic coordinate system of the sensors and actuators.