24Walking movements are orchestrated by the activation of a large number of muscles. The 25 control of numerous muscles during walking is believed to be simplified by flexible activation 26 of groups of muscles called muscle synergies. Although significant corticomuscular 27 connectivity during walking has been reported, the level at which the cortex controls locomotor 28 muscle activity (i.e., muscle synergy or individual muscle level) remains unclear. Here, we 29 examined cortical involvement in muscle control during walking by brain decoding of the 30 activation of muscle synergies and individual muscles from electroencephalographic (EEG) 31 signals using linear decoder models. First, we demonstrated that activation of locomotor muscle 32 synergies was decoded from slow cortical waves with significant accuracy. In addition, we 33 found that decoding accuracy for muscle synergy activation was greater than that for individual 34 muscle activation and that decoding of individual muscle activation was based on muscle 35 synergy-related cortical information. Taken together, these results provide indirect evidence that 36 the cerebral cortex hierarchically controls multiple muscles through a few muscle synergies 37 during walking. Our findings extend the current understanding of the role of the cortex in 38 muscular control during walking and could accelerate the development of effective 39 brain-machine interfaces for people with locomotor disabilities. 40 41 42 43 44 45 46 47 48 49 3 50
Introduction 51Human locomotor movement is organized by the coordinated activation of a large 52 number of muscles. It has been suggested that complex muscle activity is generated from a 53 small number of groups of muscle activations called muscle synergies [1][2][3][4][5]. Locomotor muscle 54 synergies are thought to be structured in the spinal circuitry [6, 7]. Based on previous studies 55 examining synergy activation among different subject groups, it has been suggested that the 56 cortex activates locomotor muscle synergies [1, 2, 7]. These studies reported that locomotor 57 muscle synergy in healthy adults exhibited activation that was sharply timed around gait events 58 [1], whereas locomotor muscle synergy in neonates [2] and complete spinal cord injury (SCI) 59 patients [7] exhibited smooth prolonged activation. The differences in the patterns in neonates 60 and SCI patients could be caused by immature and injured corticospinal pathways, respectively.
61Based on these findings, it is thought that cortical descending commands modulate basic 62 locomotor muscle synergy activation generated by subcortical structures, particularly in the 63 spinal cord. However, there is currently no direct evidence of cortico-muscle synergy 64 relationships supported by simultaneous recordings of cortical activity and muscle synergy 65 activation during walking.
66Unlike quadruped animals [8, 9], human bipedal walking is characterized by 67 significant cortical activity even during undemanding steady-state walking [10][11][12][13][14][15][16]...