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The Journal of Neuroscience, October 10, 2007, 27(41):11149-11161; doi:10.1523/JNEUROSCI.2644-07.2007

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Behavioral/Systems/Cognitive
Modular Control of Limb Movements during Human Locomotion

Yuri P. Ivanenko,¹ Germana Cappellini,¹ Nadia Dominici,¹ Richard E. Poppele,² and Francesco Lacquaniti^1,3,4

¹Department of Neuromotor Physiology, Santa Lucia Foundation, 00179 Rome, Italy, ²Department of Neuroscience, University of Minnesota, Minneapolis, Minnesota 55455, and ³Department of Neuroscience and ⁴Centre of Space Bio-Medicine, University of Rome Tor Vergata, 00173 Rome, Italy

Correspondence should be addressed to Dr. Yuri P. Ivanenko, Department of Neuromotor Physiology, Scientific Institute Foundation Santa Lucia, 306 via Ardeatina, 00179 Rome, Italy. Email: y.ivanenko{at}hsantalucia.it

The idea that the CNS may control complex interactions by modular decomposition has received considerable attention. We explored this idea for human locomotion by examining limb kinematics. The coordination of limb segments during human locomotion has been shown to follow a planar law for walking at different speeds, directions, and levels of body unloading. We compared the coordination for different gaits. Eight subjects were asked to walk and run on a treadmill at different speeds or to walk, run, and hop over ground at a preferred speed. To explore various constraints on limb movements, we also recorded stepping over an obstacle, walking with the knees flexed, and air-stepping with body weight support. We found little difference among covariance planes that depended on speed, but there were differences that depended on gait. In each case, we could fit the planar trajectories with a weighted sum of the limb length and orientation trajectories. This suggested that limb length and orientation might provide independent predictors of limb coordination. We tested this further by having the subjects step, run, and hop in place, thereby varying only limb length and maintaining limb orientation fixed, and also by marching with knees locked to maintain limb length constant while varying orientation. The results were consistent with a modular control of limb kinematics where limb movements result from a superposition of separate length- and orientation-related angular covariance. The hypothesis finds support in the animal findings that limb proprioception may also be encoded in terms of these global limb parameters.

Key words: kinematics; leg; locomotion; humans; motor primitives; proprioception

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Received June 11, 2007; revised Aug. 6, 2007; accepted Aug. 16, 2007.

Correspondence should be addressed to Dr. Yuri P. Ivanenko, Department of Neuromotor Physiology, Scientific Institute Foundation Santa Lucia, 306 via Ardeatina, 00179 Rome, Italy. Email: y.ivanenko{at}hsantalucia.it

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