Neural, mechanical, and geometric factors subserving arm posture in humans.

When the hand is displaced from an equilibrium posture by an external disturbance, a force is generated to restore the original position. We developed a new experimental method to measure and represent the field of elastic forces associated with posture of the hand in the horizontal plane. While subjects maintained a given posture, small displacements of the hand along different directions were delivered by torque motors. The hand was held in the displaced positions and, at that time, we measured the corresponding restoring forces before the onset of any voluntary reaction. The stiffness in the vicinity of the hand equilibrium position was estimated by analyzing the force and displacement vectors. We chose to represent the stiffness both numerically, as a matrix, and graphically, as an ellipse characterized by three parameters: magnitude (the area), shape (the ratio of axis) and orientation (direction of the major axis). The latter representation captures the main geometrical features of the elastic force field associated with posture. We also evaluated the conservative and nonconservative components of this elastic force field. We found that the former were much larger than the latter and concluded that the behavior of the neuromuscular system of the multiarticular arm is predominantly spring-like. Our data indicated that the shape and orientation of the stiffness were invariant over subjects and over time. We also investigated the ability of our subjects to produce voluntary and adaptive changes in the stiffness. Our findings indicated that, when a disturbance acting along a fixed and predictable direction was imposed, the magnitude of the stiffness was increased but only minor changes in shape and orientation occurred. Taken together, all of these experiments represent a step toward the understanding of the interactions between geometrical and neural factors involved in maintaining hand posture and its interactions with the environment.