Afferent input, efference copy, signal noise, and biases in perception of joint angle during active versus passive elbow movements.

Psychophysical studies have reported an overestimation of limb position in the direction of movement during the early part of active movements. The main hypothesis tested in this study is that the overestimation results from a process of forward prediction of limb state driven by an efference copy of the outgoing motor command. This hypothesis predicts that position overestimation should decrease or disappear during passive movements, for which there should be no efference copy. Seven subjects were asked to remember and to report the perceived angle of their elbow joint at different times during active and passive movements. They showed a highly velocity-dependent overestimation of the elbow joint angle near the beginning of the movement in both active and passive trials. Toward the end of the movement, subjects showed a relatively velocity-independent underestimation of their elbow angle in all trials. Contrary to the prediction of the efference copy hypothesis, the amplitude and the velocity-dependent slope of the elbow angle overestimation were both greater during the early part of passive movements than active movements. This indicates that psychophysical evidence of early overestimation of arm position on its own is not a sufficient proof of forward prediction based on an efference copy, at least under the conditions of this study. Decreased errors during active movements suggest that an efference copy can improve the accuracy of state estimation during active movements. Error patterns seem to parallel the likely level of sensorimotor noise, suggesting a probabilistic mechanism for position estimation.