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First published online September 5, 2008
Journal of Experimental Biology 211, 2989-3000 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.014357
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Running on uneven ground: leg adjustment to vertical steps and self-stability

Sten Grimmer1,*, Michael Ernst1, Michael Günther1,2 and Reinhard Blickhan1

1 Friedrich-Schiller-Universität, Institut für Sportwissenschaft, Lehrstuhl für Bewegungswissenschaft, Seidelstraße 20, D-07749 Jena, Germany
2 Eberhard-Karls-Universität, Institut für Sportwissenschaft, Arbeitsbereich III, Wilhelmstraße 124, D-72074 Tübingen, Germany

* Author for correspondence (e-mail: sten.grimmer{at}uni-jena.de)

Accepted 11 June 2008

Human running is characterized by comparably simple whole-body dynamics. These dynamics can be modelled with a point mass bouncing on a spring leg. Theoretical studies using such spring–mass models predict that running can be self-stable. In simulations, this self-stability allows for running on uneven ground without paying attention to the ground irregularities. Whether humans actually use this property of the mechanical system in such an irregular environment is, however, unclear. One way to approach this question is to study how the leg stiffness in stance and the leg orientation in flight are changed in response to ground perturbations. Here, for 11 human subjects we studied two consecutive contacts during running on uneven ground with a force plate of adjustable height (step of +5, +10 and +15 cm). We found that runners adjust their leg stiffness to the height of a vertical step. The adjustment is characterized by a 9% increase in leg stiffness in preparation for the perturbation and by a systematic decrease in proportion to the step height. At the highest vertical step (+15 cm), leg stiffness was reduced by about 26%. We also observed that the angle of attack decreased from 68 deg. to 62 deg. with increasing ground height. These leg adjustments are in accordance with the predictions of a stable spring–mass system. Furthermore, we could describe the identified leg forces and leg compressions with a simple spring–mass simulation for a given body mass, leg stiffness, angle of attack and initial conditions. We compared the experimental findings with the self-stabilizing properties of the spring–mass model, and discuss how humans use a combination of strategies that include purely mechanical self-stabilization and active neuromuscular control. Finally, beyond self-stability, we suggest that control may apply to smooth centre of mass kinematics.

Key words: biomechanics, human locomotion, spring–mass model, leg stiffness, self-stability


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