The unique structure of the human foot is seen as a crucial adaptation for bipedalism. Its arched shape makes it possible to stiffen the foot to withstand high loads when pushing off, without compromising the range of motion. Experimental studies demonstrated that manipulating foot stiffness had considerable effects on gait. In clinical practise, altered foot structure is associated with pathological gait. Yet our understanding of how foot structure influences gait mechanics is still poor. Here we used predictive simulations to explore causal relations between foot properties and whole-body gait. Our dynamic three-segment foot model with longitudinal arch improved gait predictions compared to one- and two-segment foot models and can explain measured ankle-foot kinematics and energetics. We identified three properties of the ankle-foot complex that are crucial for healthy walking: (1) compliant Achilles tendon, (2) stiff heel pad, (3) the ability to stiffen the foot. The latter requires sufficient arch height and contributions of plantar fascia, intrinsic and extrinsic foot muscles. Insufficient foot stiffness results in walking patterns with reduced push-off power. During terminal stance plantar fascia and intrinsic foot muscles transfer energy from the metatarsophalangeal to midtarsal joint, which further increases push-off power.
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