The neuromechanical tuning hypothesis

Simulations performed with neuromechanical models are providing insight into the neural control of locomotion that would be hard if not impossible to obtain in any other way. We first discuss the known properties of the neural mechanisms controlling locomotion, with a focus on mammalian systems. The rhythm-generating properties of central pattern generators (CPGs) are discussed in light of results indicating that cycle characteristics may be preset by tonic drive to spinal interneuronal networks. We then describe neuromechanical simulations that have revealed some basic rules of interaction between CPGs, sensory-mediated switching mechanisms and the biomechanics of locomotor movements. We posit that the spinal CPG timer and the sensory-mediated switch operate in parallel, the former being driven primarily by descending inputs and the latter by the kinematics. The CPG timer produces extensor and flexor phase durations, which covary along specific lines in a plot of phase- versus cycle-duration. We coined the term "phase-duration characteristics" to describe such plots. Descending input from higher centers adjusts the operating points on the phase-duration characteristics according to anticipated biomechanical requirements. In well-predicted movements, CPG-generated phase durations closely match those required by the kinematics, minimizing the corrections in phase duration required of the sensory switching mechanism. We propose the term "neuromechanical tuning" to describe this process of matching the CPG to the kinematics.