Long-latency responses during reaching account for the mechanical interaction between the shoulder and elbow joints

Although considerable research indicates that reaching movements rely on knowledge of the arm's mechanical properties and environment to anticipate and counter predictable loads, far less research has examined whether this degree of sophistication is present for on-line corrections during reaching. Here we examine the R2/3 response to mechanical perturbations (45-100 ms, also called the long-latency reflex), which is highly flexible and includes the fastest possible contribution from primary motor cortex, a key neural substrate for self-initiated action. Torque perturbations were occasionally and unexpectedly applied to the subject's shoulder and/or elbow in the course of performing reaching movements. Critically, these perturbations would evoke different patterns of feedback corrections from a shoulder extensor muscle if it countered only the local shoulder displacement relative to unperturbed motion or accounted for the mechanical interactions between the shoulder and elbow joints and countered the underlying shoulder torque. Our results show that the earliest response (R1: 20-45 ms) reflected local shoulder displacement, whereas the R2/3 response (45-100 ms) reflected knowledge of multijoint dynamics. Moreover, the same pattern of feedback occurred whether the shoulder muscle helped initiate the movement (during its agonist phase) or helped terminate the movement (during its antagonist phase). These results contribute to the accumulating evidence that highly sophisticated feedback control underlies motor behavior and are consistent with a shared neural substrate, such as primary motor cortex, for feedforward and feedback control.