The significance of closed kinematic chains to biological movement and dynamic stability

Closed kinematic chains (CKCs) are widely used in mechanical engineering because they provide a simple and efficient mechanism with multiple applications, but they are much less appreciated in living tissues. Biomechanical research has been dominated by the use of lever models and their kinematic analysis, which has largely ignored the geometric organization of these ubiquitous and evolutionary-conserved systems, yet CKCs contribute substantially to our understanding of biological motion. Closed-chain kinematics couple multiple parts into continuous mechanical loops that allow the structure itself to regulate complex movements, and are described in a wide variety of different organisms, including humans. In a biological context, CKCs are modular units nested within others at multiple size scales as part of an integrated movement system that extends throughout the organism and can act in synergy with the nervous system, where present. They provide an energy-efficient mechanism that enables multiple mechanical functions to be optimized during embryological development an increases evolutionary diversity.