Phase shift between joint rotation and actuation reflects dominant forces and predicts muscle activation patterns.

During behavior, the work done by actuators on the body can be resisted by the body's inertia, elastic forces, gravity, or viscosity. The dominant forces that resist actuation have major consequences on the control of that behavior. In the literature, features and actuation of locomotion, for example, have been successfully predicted by nondimensional numbers (e.g. Froude number and Reynolds number) that generally express the ratio between two of these forces (gravitational, inertial, elastic, and viscous). However, animals of different sizes or motions at different speeds may not share the same dominant forces within a behavior, making ratios of just two of these forces less useful. Thus, for a broad comparison of behavior across many orders of magnitude of limb length and cycle period, a dimensionless number that includes gravitational, inertial, elastic, and viscous forces is needed. This study proposes a nondimensional number that relates these four forces: the phase shift (ϕ) between the displacement of the limb and the actuator force that moves it. Using allometric scaling laws, ϕ for terrestrial walking is expressed as a function of the limb length and the cycle period at which the limb steps. Scale-dependent values of ϕ are used to explain and predict the electromyographic (EMG) patterns employed by different animals as they walk.

Investigating the role of low level reinforcement reflex loops in insect locomotion.

Insects are highly capable walkers, but many questions remain regarding how the insect nervous system controls locomotion. One particular question is how information is communicated between the 'lower level' ventral nerve cord (VNC) and the 'higher level' head ganglia to facilitate control. In this work, we seek to explore this question by investigating how systems traditionally described as 'positive feedback' may initiate and maintain stepping in the VNC with limited information exchanged between lower and higher level centers. We focus on the 'reflex reversal' of the stick insect femur-tibia joint between a resistance reflex (RR) and an active reaction in response to joint flexion, as well as the activation of populations of descending dorsal median unpaired (desDUM) neurons from limb strain as our primary reflex loops. We present the development of a neuromechanical model of the stick insect (Carausius morosus) femur-tibia (FTi) and coxa-trochanter joint control networks 'in-the-loop' with a physical robotic limb. The control network generates motor commands for the robotic limb, whose motion and forces generate sensory feedback for the network. We based our network architecture on the anatomy of the non-spiking interneuron joint control network that controls the FTi joint, extrapolated network connectivity based on known muscle responses, and previously developed mechanisms to produce 'sideways stepping'. Previous studies hypothesized that RR is enacted by selective inhibition of sensory afferents from the femoral chordotonal organ, but no study has tested this hypothesis with a model of an intact limb. We found that inhibiting the network's flexion position and velocity afferents generated a reflex reversal in the robot limb's FTi joint. We also explored the intact network's ability to sustain steady locomotion on our test limb. Our results suggested that the reflex reversal and limb strain reinforcement mechanisms are both necessary but individually insufficient to produce and maintain rhythmic stepping in the limb, which can be initiated or halted by brief, transient descending signals. Removing portions of this feedback loop or creating a large enough disruption can halt stepping independent of the higher-level centers. We conclude by discussing why the nervous system might control motor output in this manner, as well as how to apply these findings to generalized nervous system understanding and improved robotic control.

Neurodynamic modeling of the fruit fly Drosophila melanogaster.

This manuscript describes neuromechanical modeling of the fruit fly Drosophila melanogaster in the form of a hexapod robot, Drosophibot, and an accompanying dynamic simulation. Drosophibot is a testbed for real-time dynamical neural controllers modeled after the anatomy and function of the insect nervous system. As such, Drosophibot has been designed to capture features of the animal's biomechanics in order to better test the neural controllers. These features include: dynamically scaling the robot to match the fruit fly by designing its joint elasticity and movement speed; a biomimetic actuator control scheme that converts neural activity into motion in the same way as observed in insects; biomimetic sensing, including proprioception from all leg joints and strain sensing from all leg segments; and passively compliant tarsi that mimic the animal's passive compliance to the walking substrate. We incorporated these features into a dynamical simulation of Drosophibot, and demonstrate that its actuators and sensors perform in an animal-like way. We used this simulation to test a neural walking controller based on anatomical and behavioral data from insects. Finally, we describe Drosophibot's hardware and show that the animal-like features of the simulation transfer to the physical robot.

A dynamical model exploring sensory integration in the insect central complex substructures.

It is imperative that an animal has the ability to contextually integrate received sensory information to formulate appropriate behavioral responses. Determining a body heading based on a multitude of ego-motion cues and visual landmarks is an example of such a task that requires this context dependent integration. The work presented here simulates a sensory integrator in the insect brain called the central complex (CX). Based on the architecture of the CX, we assembled a dynamical neural simulation of two structures called the protocerebral bridge (PB) and the ellipsoid body (EB). Using non-spiking neuronal dynamics, our simulation was able to recreate in vivo neuronal behavior such as correlating body rotation direction and speed to activity bumps within the EB as well as updating the believed heading with quick secondary system updates. With this model, we performed sensitivity analysis of certain neuronal parameters as a possible means to control multi-system gains during sensory integration. We found that modulation of synapses in the memory network and EB inhibition are two possible mechanisms in which a sensory system could affect the memory stability and gain of another input, respectively. This model serves as an exploration in network design for integrating simultaneous idiothetic and allothetic cues in the task of body tracking and determining contextually dependent behavioral outputs.