Control variables and proprioceptive feedback in fast single-joint movement.

Sensorimotor mechanisms were studied on the basis of kinematic and electromyographic data as well as the static torque developed by the muscles as a function of joint angle. The latter relationship is known as the torque/angle characteristic. Fast single-joint movement may result from a shift in this characteristic and a change in its slope. Such movements were studied at the wrist in 9 normal and 1 deafferented subject. After training to flex the wrist to a target, subjects repeated the same movements but in random test trials movements were opposed by the load generated by linear position feedback to a torque motor. At the end of the loaded trials, the load was suddenly removed. In the second experiment, subjects made wrist movements to the target that were opposed by the load and, on random test trials, the movements were not loaded. In these test trials, the wrist arrived in a static position outside the target zone. In both experiments, subjects were instructed not to correct errors. The final torque/angle characteristics specified in the movements were reconstructed on the basis of the static wrist positions and torques before and after unloading. Normal subjects made movements by shifting the position of the torque/angle characteristic and by increasing its slope. If subjects indeed maintained the same pattern of control variables (descending commands), the same final position of the characteristic would be reproduced from trial to trial regardless of load perturbations. This assumption of equifinality was tested by comparing the final position of the wrist in nonloaded movements with that after removal of the load in loaded movements. Equifinality was observed in normal subjects. Movements in the deafferented subject were also associated with a shift of the torque/angle characteristic and a change in its slope. However, she was unable to consistently reproduce its final position. In spite of muscle coactivation, her maximal stiffness was lower than in normal subjects. In the absence of vision, the subject made movements with the load by increasing the slope of the characteristic instead of by shifting its position far enough. Load perturbation affected her final wrist position (inequifinality), which may reflect the presence of a significant hysteresis of the characteristic as a result of the absence of stretch reflexes. The deficits following deafferentation presumably result from the destruction of biomechanical and sensorimotor mechanisms including the ability of control variables to specify the positional frame of reference for afferent and descending systems.