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  1. This study introduces a novel neuromechanical model of rat hindlimbs with biarticular muscles producing walking movements without ground contact. The design of the control network is informed by the findings from our previous investigations into two-layer central pattern generators (CPGs). Specifically, we examined one plausible synthetic nervous system (SNS) designed to actuate 3 biarticular muscles, including the Biceps femoris posterior (BFP) and Rectus femoris (RF), both of which provide torque about the hip and knee joints. We conducted multiple perturbation tests on the simulation model to investigate the contribution of these two biarticular muscles in stabilizing perturbed hindlimb walking movements. We tested the BFP and RF muscles under three conditions: active, only passive tension, and fully disabled. Our results show that when these two biarticular muscles were active, they not only reduced the impact of external torques, but also facilitated rapid coordination of motion phases. As a result, the hindlimb model with biarticular muscles demonstrated faster recovery compared to our previous monoarticular muscle model. 
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  2. This work presents an in-depth numerical investigation into a hypothesized two-layer central pattern generator (CPG) that controls mammalian walking and how different parameter choices might affect the stepping of a simulated neuromechanical model. Particular attention is paid to the functional role of features that have not received a great deal of attention in previous work: the weak cross-excitatory connectivity within the rhythm generator and the synapse strength between the two layers. Sensitivity evaluations of deafferented CPG models and the combined neuromechanical model are performed. Locomotion frequency is increased in two different ways for both models to investigate whether the model’s stability can be predicted by trends in the CPG’s phase response curves (PRCs). Our results show that the weak cross-excitatory connection can make the CPG more sensitive to perturbations and that increasing the synaptic strength between the two layers results in a trade-off between forced phase locking and the amount of phase delay that can exist between the two layers. Additionally, although the models exhibit these differences in behavior when disconnected from the biomechanical model, these differences seem to disappear with the full neuromechanical model and result in similar behavior despite a variety of parameter combinations. This indicates that the neural variables do not have to be fixed precisely for stable walking; the biomechanical entrainment and sensory feedback may cancel out the strengths of excitatory connectivity in the neural circuit and play a critical role in shaping locomotor behavior. Our results support the importance of including biomechanical models in the development of computational neuroscience models that control mammalian locomotion. 
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