Long-latency reflexes of the human arm reflect an internal model of limb dynamics

A key feature of successful motor control is the ability to counter unexpected perturbations. This process is complicated in multijoint systems, like the human arm, by the fact that loads applied at one joint will create motion at other joints [1-3]. Here, we test whether our most rapid corrections, i.e., reflexes, address this complexity through an internal model of the limb's mechanical properties. By selectively applying torque perturbations to the subject's shoulder and/or elbow, we revealed a qualitative difference between the arm's short-latency/spinal reflexes and long-latency/cortical reflexes. Short-latency reflexes of shoulder muscles were linked exclusively to shoulder motion, whereas its long-latency reflexes were sensitive to both shoulder and elbow motion, i.e., matching the underlying shoulder torque. In fact, a long-latency reflex could be evoked without even stretching or lengthening the shoulder muscle but by displacing just the elbow joint. Further, the shoulder's long-latency reflexes were appropriately modified across the workspace to account for limb-geometry changes that affect the transformation between joint torque and joint motion. These results provide clear evidence that long-latency reflexes possess an internal model of limb dynamics, a degree of motor intelligence previously reserved for voluntary motor control [3-5]. The use of internal models for both voluntary and reflex control is consistent with substantial overlap in their neural substrates and current notions of intelligent feedback control [6-8].