Adaptive load feedback robustly signals force dynamics in robotic model of Carausius morosus stepping

Animals utilize a number of neuronal systems to produce locomotion. One type of sensory organ that contributes in insects is the campaniform sensillum (CS) that measures the load on their legs. Groups of the receptors are found on high stress regions of the leg exoskeleton and they have significant effects in adapting walking behavior. Recording from these sensors in freely moving animals is limited by technical constraints. To better understand the load feedback signaled by CS to the nervous system, we have constructed a dynamically scaled robotic model of the Carausius morosus stick insect middle leg. The leg steps on a treadmill and supports weight during stance to simulate body weight. Strain gauges were mounted in the same positions and orientations as four key CS groups (Groups 3, 4, 6B, and 6A). Continuous data from the strain gauges were processed through a previously published dynamic computational model of CS discharge. Our experiments suggest that under different stepping conditions (e.g., changing “body” weight, phasic load stimuli, slipping foot), the CS sensory discharge robustly signals increases in force, such as at the beginning of stance, and decreases in force, such as at the end of stance or when the foot slips. Such signals would be crucial for an insect or robot to maintain intra- and inter-leg coordination while walking over extreme terrain

Neurobiology: Reconstructing the Neural Control of Leg Coordination

SummaryWalking is adaptable because the timing of movements of individual legs can be varied while maintaining leg coordination. Recent work in stick insects shows that leg coordination set by interactions of pattern generating circuits can be overridden by sensory feedback

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Biological systems can provide useful insights into principles of design and control of locomotion that can be applied to legged robots. In this paper we review our work on cockroaches using finite element analysis to model how loads are sensed and regulated in walking and climbing. A number of biological studies have shown that sensors that detect forces in the legs of insects are of particular importance in controlling walking and adapting locomotion to non-horizontal terrains. Our analysis strongly suggests that (i) the system can detect specific force vectors (body load versus propulsion) via sensors located in the leg in positions close to the body and (ii) the system uses this information in positive load feedback to regulate walking movements. These principles and design elements provide examples that can be applied in legged locomotion in walking machines. KEY WORDS—strain sensors, control, cockroaches, campaniform sensilla, finite element analysi

Encoding of force increases and decreases by tibial campaniform sensilla in the stick insect, Carausius morosus

Zill SN, Büschges A, Schmitz J. Encoding of force increases and decreases by tibial campaniform sensilla in the stick insect, Carausius morosus. Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology. 2011;197(8):851-867

Speed-dependent interplay between local pattern-generating activity and sensory signals during walking in Drosophila

In insects, the coordinated motor output required for walking is based on the interaction between local pattern-generating networks providing basic rhythmicity and leg sensory signals, which modulate this output on a cycle-to-cycle basis. How this interplay changes speed-dependently and thereby gives rise to the different coordination patterns observed at different speeds is not sufficiently understood. Here, we used amputation to reduce sensory signals in single legs and decouple them mechanically during walking in Drosophila. This allowed for the dissociation between locally generated motor output in the stump and coordinating influences from intact legs. Leg stumps were still rhythmically active during walking. Although the oscillatory frequency in intact legs was dependent on walking speed, stumps showed a high and relatively constant oscillation frequency at all walking speeds. At low walking speeds we found no strict cycle-to-cycle coupling between stumps and intact legs. In contrast, at high walking speeds stump oscillations were strongly coupled to the movement of intact legs on a one-to-one basis. Although during slow walking there was no preferred phase between stumps and intact legs, we nevertheless found a preferred time interval between touch-down or lift-off events in intact legs and levation or depression of stumps. Based on these findings, we hypothesize that, as in other insects, walking speed in Drosophila is predominantly controlled by indirect mechanisms and that direct modulation of basic pattern-generating circuits plays a subsidiary role. Furthermore, interleg coordination strength seems to be speed-dependent and greater coordination is evident at higher walking speeds

Force feedback reinforces muscle synergies in insect legs

The nervous system solves complex biomechanical problems by activating muscles in modular, synergist groups. We have studied how force feedback in substrate grip is integrated with effects of sense organs that monitor support and propulsion in insects. Campaniform sensilla are mechanoreceptors that encode forces as cuticular strains. We tested the hypothesis that integration of force feedback from receptors of different leg segments during grip occurs through activation of specific muscle synergies. We characterized the effects of campaniform sensilla of the feet (tarsi) and proximal segments (trochanter and femur) on activities of leg muscles in stick insects and cockroaches. In both species, mechanical stimulation of tarsal sensilla activated the leg muscle that generates substrate grip (retractor unguis), as well as proximal leg muscles that produce inward pull (tibial flexor) and support/propulsion (trochanteral depressor). Stimulation of campaniform sensilla on proximal leg segments activated the same synergistic group of muscles. In stick insects, the effects of proximal receptors on distal leg muscles changed and were greatly enhanced when animals made active searching movements. In insects, the task-specific reinforcement of muscle synergies can ensure that substrate adhesion is rapidly established after substrate contact to provide a stable point for force generation. (C) 2015 Elsevier Ltd. All rights reserved