Soft tissue deformations explain most of the mechanical work variations of human walking

Humans perform mechanical work during walking, some by leg joints actuated by muscles, and some by passive, dissipative soft tissues. Dissipative losses must be restored by active muscle work, potentially in amounts sufficient to cost substantial metabolic energy. The most dissipative, and therefore costly, walking conditions might be predictable from the pendulum-like dynamics of the legs. If this behavior is systematic, it may also predict the work distribution between active joints and passive soft tissues. We therefore tested whether the overall negative work of walking, and the fraction owing to soft tissue dissipation, are both predictable by a simple dynamic walking model across a wide range of conditions. The model predicts whole-body negative work from the leading leg's impact with the ground (termed the collision), to increase with the squared product of walking speed and step length. We experimentally tested this in humans (N=9) walking in 26 different combinations of speed (0.7-2.0 m s-1) and step length (0.5-1.1 m), with recorded motions and ground reaction forces. Whole-body negative collision work increased as predicted (R2=0.73), with a consistent fraction of approximately 63% (R2=0.88) owing to soft tissues. Soft tissue dissipation consistently accounted for approximately 56% of the variation in total whole-body negative work, across a wide range of speed and step length combinations. During typical walking, active work to restore dissipative losses could account for 31% of the net metabolic cost. Soft tissue dissipation, not included in most biomechanical studies, explains most of the variation in negative work of walking, and could account for a substantial fraction of the metabolic cost.