Controlling multijoint motor behavior

Abstract

Much can be learned about the central nervous system from a study of motor coordination, but its true richness and complexity become evident only in a multiarticular system. Despite the intrinsic complexity of multiarticular actions, they offer an unparalleled opportunity to learn about the central nervous system in a quantitative and experimentally testable way. For example, the observation that unconstrained, unperturbed arm movements are coordinated in terms of hand motion shows that motor control is organized in a hierarchy of increasing levels of abstraction. These arm motions are organized as though a disembodied hand could be moved in space; the details of how this is to be achieved must then be supplied by a different level in the hierarchy. The essence of human behavior is its adaptability. Just as the true complexity of coordination is evident only in multiarticular actions, the sophistication and subtlety of adaptive behavior are evident only in dynamic, interactive tasks. A study of movement alone is not sufficient to understand this behavior. The dynamic response of the limbs becomes the overriding concern and must be controlled by the central nervous system. The dynamic response of a limb is usually associated with its posture, rather than its movement, but in a functional task such as the use of a tool, the postural dynamics are an integral part of the action. This perspective on motor behavior leads to some useful insights. Coordination is not a problem for movement alone; in a multiarticular system, even posture requires coordination and control. Muscles do not merely act reciprocally to generate forces about the joints; the net mechanical impedance of the limb may be controlled by synergistic activation of all muscles, including antagonists. Controlling dynamic behavior is a far more demanding task than controlling motion. Consequently, features of the neuromusculoskeletal system that appear to be redundant or unnecessary for movement control can play a functional role in controlling dynamic behavior. Polyarticular muscles contribute to the mechanical impedance in a unique way. Skeletal redundancies have a profound influence on all aspects of dynamic behavior, including the apparent inertia of the limbs. Redundancies are commonly perceived as a complicating factor in the control of motion, a problem that must be solved by the central nervous system. Rather than presenting a problem requiring solution, they may present a solution to a problem. Posture is not merely the outcome of a motor act; it is one of the important preparatory stages in the production of motor behavior.