Control theory arose from a need to control synthetic systems. From regulating steam engines to tuning radios to devices capable of autonomous movement, it provided a formal mathematical basis for understanding the role of feedback in the stability (or change) of dynamical systems. It provides a framework for understanding any system with regulation via feedback, including biological ones such as regulatory gene networks, cellular metabolic systems, sensorimotor dynamics of moving animals, and even ecological or evolutionary dynamics of organisms and populations. Here, we focus on four case studies of the sensorimotor dynamics of animals, each of which involves the application of principles from control theory to probe stability and feedback in an organism’s response to perturbations. We use examples from aquatic (two behaviors performed by electric fish), terrestrial (following of walls by cockroaches), and aerial environments (flight control by moths) to highlight how one can use control theory to understand the way feedback mechanisms interact with the physical dynamics of animals to determine their stability and response to sensory inputs and perturbations. Each case study is cast as a control problem with sensory input, neural processing, and motor dynamics, the output of which feeds back to the sensory inputs. Collectively, the interaction of these systems in a closed loop determines the behavior of the entire system.

In this paper, we analyze the self-stability properties of planar running with a dissipative spring-mass model driven by torque actuation at the hip. We first show that a two-dimensional, approximate analytic return map for uncontrolled locomotion with this system under a fixed touchdown leg angle policy and an open-loop ramp torque profile exhibits only marginal self-stability that does not always persist for the exact system. We then propose a per-stride feedback strategy for the hip torque that explicitly compensates for damping losses, reducing the return map to a single dimension and substantially improving the robust stability of fixed points. Subsequent presentation of simulation evidence establishes that the predictions of this approximate model are consistent with the behavior of the exact plant model. We illustrate the relevance and utility of our model both through the qualitative correspondence of its predictions to biological data as well as its use in the design of a task-level running controller.

This note proposes a new methodology for subspace-based state-space identification for linear time-periodic (LTP) systems. Since LTP systems can be lifted to equivalent linear time-invariant (LTI) systems, we first lift input-output data from the unknown LTP system as if it was collected from an equivalent LTI system. Then, we use frequency-domain subspace identification methods to find an LTI system estimate. Subsequently, we propose a novel method to obtain a time-periodic realization for the estimated lifted LTI system by exploiting the specific parametric structure of Fourier series coefficients of the frequency-domain lifting method. Our method can be used to both obtain state-space estimates for unknown LTP systems as well as to obtain Floquet transforms for known LTP systems.

Stability and performance during rhythmic motor behaviors such as locomotion are critical for survival across taxa: falling down would bode well for neither cheetah nor gazelle. Little is known about how haptic feedback, particularly during discrete events such as the heel-strike event during walking, enhances rhythmic behavior. To determine the effect of haptic cues on rhythmic motor performance, we investigated a virtual paddle juggling behavior, analogous to bouncing a table tennis ball on a paddle. Here, we show that a force impulse to the hand at the moment of ball-paddle collision categorically improves performance over visual feedback alone, not by regulating the rate of convergence to steady state (e.g., via higher gain feedback or modifying the steady-state hand motion), but rather by reducing cycle-to-cycle variability. This suggests that the timing and state cues afforded by haptic feedback decrease the nervous system's uncertainty of the state of the ball to enable more accurate control but that the feedback gain itself is unaltered. This decrease in variability leads to a substantial increase in the mean first passage time, a measure of the long-term metastability of a stochastic dynamical system. Rhythmic tasks such as locomotion and juggling involve intermittent contact with the environment (i.e., hybrid transitions), and the timing of such transitions is generally easy to sense via haptic feedback. This timing information may improve metastability, equating to less frequent falls or other failures depending on the task.