Human balancing of an inverted pendulum with a compliant linkage: neural control by anticipatory intermittent bias.

These experiments were prompted by the recent discovery that the intrinsic stiffness of the ankle is inadequate to stabilise passively the body in standing. Our hope was that showing how a large inverted pendulum was manually balanced with low intrinsic stiffness would elucidate the active control of human standing. The results show that the pendulum can be satisfactorily stabilised when intrinsic stiffness is low. Analysis of sway size shows that intrinsic stiffness actually plays little part in stabilisation. The sway duration is also substantially independent of intrinsic stiffness. This suggests that the characteristic sway of the pendulum, rather than being dictated by stiffness and inertia, may result from the control pattern of hand movements. The key points revealed by these experiments are that with low intrinsic stiffness the hand provides pendulum stability by intermittently altering the bias of the spring and, on average, the hand moves in opposition to the load. The results lead to a new and testable hypothesis; namely that in standing, the calf muscle shortens as the body sways forward and lengthens as it sways backwards. These findings are difficult to reconcile with stretch reflex control of the pendulum and are of particular relevance to standing. They may also be relevant to postural maintenance in general whenever the CNS controls muscles which operate through compliant linkages. The results also suggest that in standing, rather than providing passive stability, the intrinsic stiffness acts as an energy efficient buffer which provides decoupling between muscle and body.