Variational analysis of sensory feedback mechanisms in powerstroke–recovery systems

Although the raison d'etre of the brain is the survival of the body, there are relatively few theoretical studies of closed-loop rhythmic motor control systems. In this paper we provide a unified framework, based on variational analysis, for investigating the dual goals of performance and robustness in powerstroke-recovery systems. To demonstrate our variational method, we augment two previously published closed-loop motor control models by equipping each model with a performance measure based on the rate of progress of the system relative to a spatially extended external substrate-such as a long strip of seaweed for a feeding task, or progress relative to the ground for a locomotor task. The sensitivity measure quantifies the ability of the system to maintain performance in response to external perturbations, such as an applied load. Motivated by a search for optimal design principles for feedback control achieving the complementary requirements of efficiency and robustness, we discuss the performance-sensitivity patterns of the systems featuring different sensory feedback architectures. In a paradigmatic half-center oscillator-motor system, we observe that the excitation-inhibition property of feedback mechanisms determines the sensitivity pattern while the activation-inactivation property determines the performance pattern. Moreover, we show that the nonlinearity of the sigmoid activation of feedback signals allows the existence of optimal combinations of performance and sensitivity. In a detailed hindlimb locomotor system, we find that a force-dependent feedback can simultaneously optimize both performance and robustness, while length-dependent feedback variations result in significant performance-versus-sensitivity tradeoffs. Thus, this work provides an analytical framework for studying feedback control of oscillations in nonlinear dynamical systems, leading to several insights that have the potential to inform the design of control or rehabilitation systems.