Kinematic control of walking.
The planar law of inter-segmental co-ordination we described may emerge from the coupling of neural oscillators between each other and with limb mechanical oscillators. Muscle contraction intervenes at variable times to re-excite the intrinsic oscillations of the system when energy is lost. The hypothesis that a law of coordinative control results from a minimal active tuning of the passive inertial and viscoelastic coupling among limb segments is congruent with the idea that movement has evolved according to minimum energy criteria (1, 8). It is known that multi-segment motion of mammals locomotion is controlled by a network of coupled oscillators (CPGs, see 18, 33, 37). Flexible combination of unit oscillators gives rise to different forms of locomotion. Inter-oscillator coupling can be modified by changing the synaptic strength (or polarity) of the relative spinal connections. As a result, unit oscillators can be coupled in phase, out of phase, or with a variable phase, giving rise to different behaviors, such as speed increments or reversal of gait direction (from forward to backward). Supra-spinal centers may drive or modulate functional sets of coordinating interneurons to generate different walking modes (or gaits). Although it is often assumed that CPGs control patterns of muscle activity, an equally plausible hypothesis is that they control patterns of limb segment motion instead (22). According to this kinematic view, each unit oscillator would directly control a limb segment, alternately generating forward and backward oscillations of the segment. Inter-segmental coordination would be achieved by coupling unit oscillators with a variable phase. Inter-segmental kinematic phase plays the role of global control variable previously postulated for the network of central oscillators. In fact, inter-segmental phase shifts systematically with increasing speed both in man (4) and cat (38). Because this phase-shift is correlated with the net mechanical power output over a gait cycle (3, 4), phase control could be used for limiting the overall energy expenditure with increasing speed (22). Adaptation to different walking conditions, such as changes in body posture, body weight unloading and backward walk, also involves inter-segmental phase tuning, as does the maturation of limb kinematics in toddlers.
Lacquaniti
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Human Physiology Section, University of Rome Tor Vergata, Scientific Institute Santa Lucia, Via Ardeatina 306, 00179 Rome, Italy. lacquaniti@caspur.it
Ivanenko
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YP
Zago
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D000818:Animals; D001683:Biological Clocks / Q000502:physiology*; D001696:Biomechanical Phenomena; D005121:Extremities / Q000502:physiology*; D005684:Gait / Q000502:physiology*; D006801:Humans; D008124:Locomotion / Q000502:physiology*; D009068:Movement / Q000502:physiology*; D009119:Muscle Contraction / Q000502:physiology; D011187:Posture / Q000502:physiology