The human foot and heel–sole–toe walking strategy: a mechanism enabling an inverted pendular gait with low isometric muscle force?

Mechanically, the most economical gait for slow bipedal locomotion requires walking as an ‘inverted pendulum’, with: I, an impulsive, energy-dissipating leg compression at the beginning of stance; II, a stiff-limbed vault; and III, an impulsive, powering push-off at the end of stance. The characteristic ‘M’-shaped vertical ground reaction forces of walking in humans reflect this impulse–vault–impulse strategy. Humans achieve this gait by dissipating energy during the heel-to-sole transition in early stance, approximately stiff-limbed, flat-footed vaulting over midstance and ankle plantarflexion (powering the toes down) in late stance. Here, we show that the ‘M’-shaped walking ground reaction force profile does not require the plantigrade human foot or heel–sole–toe stance; it is maintained in tip–toe and high-heel walking as well as in ostriches. However, the unusual, stiff, human foot structure—with ground-contacting heel behind ankle and toes in front—enables both mechanically economical inverted pendular walking and physiologically economical muscle loading, by producing extreme changes in mechanical advantage between muscles and ground reaction forces. With a human foot, and heel–sole–toe strategy during stance, the shin muscles that dissipate energy, or calf muscles that power the push-off, need not be loaded at all—largely avoiding the ‘cost of muscle force’—during the passive vaulting phase.