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During quiet standing the human "inverted pendulum" sways irregularly. In previous work where subjects balanced a real inverted pendulum, we investigated what contribution the intrinsic mechanical ankle stiffness makes to achieve stability. Using the results of a plausible model, we suggested that intrinsic ankle stiffness is inadequate for providing(More)
It has been widely assumed for nearly a century, that postural muscles operate in a spring-like manner and that muscle length signals joint angle (the mechano-reflex mechanism). Here we employ automated analysis of ultrasound images to resolve calf muscle (soleus and gastrocnemius) length changes as small as 10 mum in standing subjects. Previously, we have(More)
In humans, during standing the calf muscles soleus and gastrocnemius actively prevent forward toppling about the ankles. It has been generally assumed that these postural muscles behave like springs with dynamic stiffness reflecting their mechanical properties, reflex gain including higher derivatives, and central control. Here, for the first time, we have(More)
In cat medial gastrocnemius (MG), fibres supplied by individual motoneurones (muscle units) distribute extensively along the muscle longitudinal axis. In the human MG, the size of motor unit territory is unknown. It is uncertain if the absolute size of muscle unit territory or the size relative to the whole muscle is most comparable with the cat. By(More)
In standing, there are small sways of the body. Our interest is to use an artificial task to illuminate the mechanisms underlying the sways and to account for changes in their size. Using the ankle musculature, subjects balanced a large inverted pendulum. The equilibrium of the pendulum is unstable and quasi-regular sway was observed like that in quiet(More)
The paradigm of continuous control using internal models has advanced understanding of human motor control. However, this paradigm ignores some aspects of human control, including intermittent feedback, serial ballistic control, triggered responses and refractory periods. It is shown that event-driven intermittent control provides a framework to explain the(More)
While standing naturally and when manually or pedally balancing an equivalent inverted pendulum, the load sways slowly (characteristic unidirectional duration approximately 1 s) and the controller, calf muscles or hand, makes more frequent adjustments (characteristic unidirectional duration 400 ms). Here we test the hypothesis that these durations reflect(More)
Human balance is commonly described using linear-time-invariant (LTI) models. The feedback time delay determines the position of balance in the motor-control hierarchy. The extent of LTI control illuminates the automaticity of the control process. Using non-parametric analysis, we measured the feedback delay, extent of LTI control and visuo-motor transfer(More)
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(More)
During human standing, tonic ankle extensor torque is required to support the centre of mass (CoM) forward of the ankles, and dynamic torque modulation is required to maintain unstable balance. Passive mechanisms contribute to both but the extent is controversial. Some groups have revealed a substantial intrinsic stiffness (65-90%) normalized to load(More)