Motor patterns for human gait: backward versus forward locomotion.
Seven healthy subjects walked forward (FW) and backward (BW) at different freely chosen speeds, while their motion, ground reaction forces, and electromyographic (EMG) activity from lower limb muscles were recorded. We considered the time course of the elevation angles of the thigh, shank, and foot segments in the sagittal plane, the anatomic angles of the hip, knee, and ankle joints, the vertical and longitudinal ground reaction forces, and the rectified EMGs. The elevation angles were the most reproducible variables across trials in each walking direction. After normalizing the time course of each variable over the gait cycle duration, the waveforms of all elevation angles in BW gait were essentially time reversed relative to the corresponding waveforms in FW gait. Moreover, the changes of the thigh, shank, and foot elevation covaried along a plane during the whole gait cycle in both FW and BW directions. Cross-correlation analysis revealed that the phase coupling among these elevation angles is maintained with a simple reversal of the delay on the reversal of walking direction. The extent of FW-BW correspondence also was good for the hip angle, but it was smaller for the knee and ankle angles and for the ground reaction forces. The EMG patterns were drastically different in the two movement directions as was the organization of the muscular synergies measured by cross-correlation analysis. Moreover, at any given speed, the mean EMG activity over the gait cycle was generally higher in BW than in FW gait, suggesting a greater level of energy expenditure in the former task. We argue that conservation of kinematic templates across gait reversal at the expense of a complete reorganization of muscle synergies does not arise from biomechanical constraints but may reflect a behavioral goal achieved by the central networks involved in the control of locomotion.
Human Physiology Section, Scientific Institute Santa Lucia, National Research Council, University of Tor Vergata, 00179 Rome, Italy.
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Grasso
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Bianchi
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Lacquaniti
D000328:Adult; D004576:Electromyography; D005260:Female; D005684:Gait / Q000502:physiology*; D006801:Humans; D007866:Leg; D008297:Male; D009068:Movement / Q000502:physiology*; D018482:Muscle, Skeletal / Q000502:physiology; D015203:Reproducibility of Results; D013997:Time Factors