The elastic stretch-shortening cycle of the Achilles tendon during walking can reduce the active work demands on the plantarflexor muscles in series. However, this does not explain why or when this ankle work, whether by muscle or tendon, needs to be performed during gait. We therefore employ a simple bipedal walking model to investigate how ankle work and series elasticity impact economical locomotion. Our model shows that ankle elasticity can use passive dynamics to aid push-off late in single support, redirecting the body's center-of-mass (COM) motion upward. An appropriately timed, elastic push-off helps to reduce dissipative collision losses at contralateral heelstrike, and therefore the positive work needed to offset those losses and power steady walking. Thus, the model demonstrates how elastic ankle work can reduce the total energetic demands of walking, including work required from more proximal knee and hip muscles. We found that the key requirement for using ankle elasticity to achieve economical gait is the proper ratio of ankle stiffness to foot length. Optimal combination of these parameters ensures proper timing of elastic energy release prior to contralateral heelstrike, and sufficient energy storage to redirect the COM velocity. In fact, there exist parameter combinations that theoretically yield collision-free walking, thus requiring zero active work, albeit with relatively high ankle torques. Ankle elasticity also allows the hip to power economical walking by contributing indirectly to push-off. Whether walking is powered by the ankle or hip, ankle elasticity may aid walking economy by reducing collision losses.

The human ankle produces a large burst of 'push-off' mechanical power late in the stance phase of walking, reduction of which leads to considerably poorer energy economy. It is, however, uncertain whether the energetic penalty results from poorer efficiency when the other leg joints substitute for the ankle's push-off work, or from a higher overall demand for work due to some fundamental feature of push-off. Here, we show that greater metabolic energy expenditure is indeed explained by a greater demand for work. This is predicted by a simple model of walking on pendulum-like legs, because proper push-off reduces collision losses from the leading leg. We tested this by experimentally restricting ankle push-off bilaterally in healthy adults (N=8) walking on a treadmill at 1.4 m s −1 , using ankle-foot orthoses with steel cables limiting motion. These produced up to ∼50% reduction in ankle pushoff power and work, resulting in up to ∼50% greater net metabolic power expenditure to walk at the same speed. For each 1 J reduction in ankle work, we observed 0.6 J more dissipative collision work by the other leg, 1.3 J more positive work from the leg joints overall, and 3.94 J more metabolic energy expended. Loss of ankle push-off required more positive work elsewhere to maintain walking speed; this additional work was performed by the knee, apparently at reasonably high efficiency. Ankle push-off may contribute to walking economy by reducing dissipative collision losses and thus overall work demand.