A biomechanical perspective on spinal mechanisms of coordinated muscular action: an architecture principle.

A principle of organization of spinal circuitry which emerges from the studies reviewed here is that the structure of the distributed network of pathways in the spinal cord contains a detailed representation of the corresponding three dimensional architecture of the musculoskeletal system. The pertinent architectural features for a given muscle include (1) the number and identity of spanned joints, and (2) the line of action at a joint with respect to the torque directions of other muscles and to the gravity vector. In accordance with established ideas, muscles with common primary actions (synergists) at the ankle are linked by excitatory, length-dependent pathways. Those muscles which have opposite actions are linked by reciprocal inhibition, although muscles which are not principally involved in postural control are not connected in this way. Among antigravity and stabilizing muscles, force-dependent, inhibitory pathways link (1) muscles crossing different joints, and (2) members of different synergistic groups which exert torques in different directions. Therefore, each muscle has a unique set of actions in terms of joints spanned and line of action, and each muscle receives a unique combination of reflex inputs. The cross-joint coordination resulting from actions of force-dependent pathways becomes stronger at higher forces with a consequent reduction in degrees of freedom of the musculoskeletal system. Length-dependent pathways link muscles which share some, but not all, mechanical actions at a joint and may have different patterns of activation during locomotion. Length-dependent pathways appear to coordinate muscle responses to postural disturbances and enhance joint stiffness.