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The Journal of Neuroscience, August 3, 2005, 25(31):7238-7253; doi:10.1523/JNEUROSCI.1327-05.2005
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Behavioral/Systems/Cognitive
Coordination of Locomotion with Voluntary Movements in Humans
Yuri P. Ivanenko,1
Germana Cappellini,1
Nadia Dominici,1,2
Richard E. Poppele,4 and
Francesco Lacquaniti1,2,3
1Department of Neuromotor Physiology, Scientific Institute Foundation Santa Lucia, 00179 Rome, Italy, 2Department of Neuroscience and 3Centre of Space Bio-medicine, University of Rome Tor Vergata, 00173 Rome, Italy, and 4Department of Neuroscience, University of Minnesota, Minneapolis, Minnesota 55455
Muscle activity occurring during human locomotion can be accounted for by five basic temporal activation patterns in a variety of locomotion conditions. Here, we examined how these activation patterns interact with muscle activity required for a voluntary movement. Subjects produced a voluntary movement during locomotion, and we examined the resulting kinematics, kinetics, and EMG activity in 16-31 ipsilateral limb and trunk muscles during the tasks. There were four voluntary tasks added to overground walking ( 5 km/h) in which subjects kicked a ball, stepped over an obstacle, or reached down and grasped an object on the floor (weight support on either the right or the left foot). Statistical analyses of EMG waveforms showed that the five basic locomotion patterns were invariantly present in each task, although they could be differently weighted across muscles, suggesting a characteristic locomotion timing of muscle activations. We also observed a separate activation that was timed to the voluntary task. The coordination of locomotion with the voluntary task was accomplished by combining activation timings that were associated separately with the voluntary task and locomotion. Activation associated with the voluntary tasks was either synchronous with the timing for locomotion or had additional activations not represented in the basic locomotion timing. We propose that this superposition of an invariant locomotion timing pattern with a voluntary activation timing may be consistent with the proposal suggesting that compound movements are produced through a superposition of motor programs.
Key words: EMG activity; motor primitives; central pattern generators; spinal cord; human locomotion; walking
Received Jan 26, 2005;
revised June 16, 2005;
accepted June 17, 2005.
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