Abstract

The ability of vertebrates to generate rhythm within their spinal neural networks is essential for walking, running, and other rhythmic behaviors. The central pattern generator (CPG) network responsible for these behaviors is well-characterized with experimental and theoretical studies, and it can be formulated as a nonlinear dynamical system. The underlying mechanism responsible for locomotor behavior can be expressed as the process of leaky integration with resetting states generating appropriate phases for changing body velocity. The low-dimensional input to the CPG model generates the bilateral pattern of swing and stance modulation for each limb and is consistent with the desired limb speed as the input command. To test the minimal configuration of required parameters for this model, we reduced the system of equations representing CPG for a single limb and provided the analytical solution with two complementary methods. The analytical and empirical cycle durations were similar (R2 = 0.99) for the full range of walking speeds. The structure of solution is consistent with the use of limb speed as the input domain for the CPG network. Moreover, the reciprocal interaction between two leaky integration processes representing a CPG for two limbs was sufficient to capture fundamental experimental dynamics associated with the control of heading direction. This analysis provides further support for the embedded velocity or limb speed representation within spinal neural pathways involved in rhythm generation.

Highlights

  • The mechanism of spinal rhythmogenesis is an integral part of the mammalian locomotor system that fuses descending and sensory feedback signals with body dynamics (Dickinson et al, 2000)

  • Using an analytical central pattern generator (CPG) model, we have demonstrated previously that the asymmetric gait can be represented with the strengths of connections between intrinsic elements of a relatively simple bilateral CPG (Sobinov & Yakovenko, 2018)

  • The relationship between step cycle duration and the input ‘‘drive’’ to the analytical model was investigated in two complimentary solutions that rely on different assumptions: (i) the assumption of constant integration rate in a single limb model of CPG, and (ii) the expansion of function with the common Taylor series method

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Summary

Introduction

The mechanism of spinal rhythmogenesis is an integral part of the mammalian locomotor system that fuses descending and sensory feedback signals with body dynamics (Dickinson et al, 2000). The elusive mechanism of locomotor pattern generation remains poorly understood in the context of its regulation and integration within descending feedforward and sensory feedback pathways. One of the main obstacles is the definition of CPG’s essential function. We know that this neural element can compute control commands for the redundant musculoskeletal system (Gritsenko et al, 2016) that, in turn, shapes the activity of hierarchical neural mechanisms (Lillicrap & Scott, 2013) distributed along the neuraxis (Grillner, 1985). The spinal motor circuits are known to accommodate rewiring in healthy operation (Vahdat et al, 2015) and injured states (Stevenson et al, 2015; Liu et al, 2017)

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