Hwang EJ, Smith MA, and Shadmehr R (2006) Adaptation and
generalization in acceleration dependent force fields, Experimental Brain
Research 169:496-506.
Abstract Any passive rigid inertial object
that we hold in our hand, e.g., a tennis racquet, imposes a field of forces on
the arm that depends on limb position, velocity, and acceleration. A
fundamental characteristic of this field is that the forces due to acceleration
and velocity are linearly separable in the intrinsic coordinates of the limb.
In order to learn such dynamics with a collection of basis elements, a control
system would generalize correctly and therefore perform optimally if the basis
elements that were sensitive to limb velocity were not sensitive to
acceleration, and vice versa. However, in the mammalian nervous system
proprioceptive sensors like muscle spindles encode a nonlinear combination of
all components of limb state, with sensitivity to velocity dominating
sensitivity to acceleration. Therefore, limb state in the space of proprioception
is not linearly separable despite the fact that this separation is a desirable
property of control systems that form models of inertial objects. In building
internal models of limb dynamics, does the brain use a representation that is
optimal for control of inertial objects, or a representation that is closely
tied to how peripheral sensors measure limb state? Here we show that in humans,
patterns of generalization of reaching movements in acceleration dependent
fields are strongly inconsistent with basis elements that are optimized for
control of inertial objects. Unlike a robot controller that models the dynamics
of the natural world and represents velocity and acceleration independently,
internal models of dynamics that people learn appear to be rooted in the
properties of proprioception, nonlinearly responding to the pattern of muscle
activation and representing velocity more strongly than acceleration.
[fulltext-pdf]