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J Physiol Volume 577, Number 1, 417-432, November 15, 2006 DOI: 10.1113/jphysiol.2006.118786
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INTEGRATIVE

The frequency of human, manual adjustments in balancing an inverted pendulum is constrained by intrinsic physiological factors

Ian D. Loram1, Peter J. Gawthrop2 and Martin Lakie3

1 Institute for Biophysical and Clinical Research into Human Movement, Manchester Metropolitan University, Alsager ST7 2HL, UK
2 Department of Mechanical Engineering, Centre for Systems and Control, University of Glasgow, Glasgow G12 8QQ, UK
3 Applied Physiology Research Group, School of Sport and Exercise Sciences, University of Birmingham, Birmingham B15 2TT, UK

While standing naturally and when manually or pedally balancing an equivalent inverted pendulum, the load sways slowly (characteristic unidirectional duration ~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 load properties rather than some intrinsic property of the human neuromuscular system. Using a specialized set-up mechanically analogous to real standing, subjects manually balanced inverted pendulums with different moments of inertia through a compliant spring representing the Achilles tendon. The spring bias was controlled by a sensitive joystick via a servo motor and accurate visual feedback was provided on an oscilloscope. As moment of inertia decreased, inverted pendulum sway size increased and it became difficult to sustain successful balance. The mean duration of unidirectional balance adjustments did not change. Moreover, the mean duration of unidirectional inverted pendulum sway reduced only slightly, remaining around 1 s. The simplest explanation is that balance was maintained by a process of manual adjustments intrinsically limited to a mean frequency of two to three unidirectional adjustments per second corresponding to intermittent control observed in manual tracking experiments. Consequently the inverted pendulum sway duration, mechanically related to the bias duration, reflects an intrinsic constraint of the neuromuscular control system. Given the similar durations of sway and muscle adjustments observed in real standing, we postulate that the characteristic duration of unidirectional standing sway reflects intrinsic intermittent control rather than the inertial properties of the body.

(Received 7 August 2006; accepted after revision 12 September 2006; first published online 14 September 2006)
Corresponding author I. Loram: Institute for Biophysical and Clinical Research into Human Movement, Manchester Metropolitan University, Alsager ST7 2HL, UK.  Email: i.loram{at}mmu.ac.uk




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