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The hypothesis of modular control, which stands on the existence of muscle synergies as building blocks of muscle coordination, has been investigated in a great variety of motor tasks and species. Yet, its role during learning processes is still largely unexplored. To what extent is such modular control flexible, in terms of spatial structure and temporal activation, to externally or internally induced adaptations, is a debated issue. To address this question, we designed a biofeedback experiment to induce changes in the timing of muscle activations during leg cycling movements. The protocol consisted in delaying the peak of activation of one target muscle and using its electromyography (EMG) envelope as visual biofeedback. For each of the 10 healthy participants, the protocol was repeated for three different target muscles: Tibialis Anterioris (TA), Gastrocnemius Medialis (GM), and Vastus Lateralis (VL). To explore the effects of the conditioning protocol, we analyzed changes in the activity of eight lower limb muscles by applying different models of modular motor control [i.e., fixed spatial components (FSC) and fixed temporal components (FTC)]. Our results confirm the hypothesis that visual EMG biofeedback is able to induce changes in muscle coordination. Subjects were able to shift the peak of activation of the target muscle, with a delay of (49 ± 27°) across subjects and conditions. This time shift generated a reorganization of all the other muscles in terms of timing and amplitude. By using different models of modular motor control, we demonstrated that neither spatially invariant nor temporally invariant muscle synergies alone were able to account for these changes in muscle coordination after learning, while temporally invariant muscle synergies with adjustments in timing could capture most of muscle activity adaptations observed after the conditioning protocol. These results suggest that short-term learning in rhythmic tasks is built upon synergistic temporal commands that are robust to changes in the task demands.

Reorganization of Muscle Coordination Underlying Motor Learning in Cycling Tasks