Geometric phase predicts locomotion performance in undulating living systems across scales.

Self-propelling organisms locomote via generation of patterns of self-deformation. Despite the diversity of body plans, internal actuation schemes and environments in limbless vertebrates and invertebrates, such organisms often use similar traveling waves of axial body bending for movement. Delineating how self-deformation parameters lead to locomotor performance (e.g. speed, energy, turning capabilities) remains challenging. We show that a geometric framework, replacing laborious calculation with a diagrammatic scheme, is well-suited to discovery and comparison of effective patterns of wave dynamics in diverse living systems. We focus on a regime of undulatory locomotion, that of highly damped environments, which is applicable not only to small organisms in viscous fluids, but also larger animals in frictional fluids (sand) and on frictional ground. We find that the traveling wave dynamics used by mm-scale nematode worms and cm-scale desert dwelling snakes and lizards can be described by time series of weights associated with two principal modes. The approximately circular closed path trajectories of mode weights in a self-deformation space enclose near-maximal surface integral (geometric phase) for organisms spanning two decades in body length. We hypothesize that such trajectories are targets of control (which we refer to as "serpenoid templates"). Further, the geometric approach reveals how seemingly complex behaviors such as turning in worms and sidewinding snakes can be described as modulations of templates. Thus, the use of differential geometry in the locomotion of living systems generates a common description of locomotion across taxa and provides hypotheses for neuromechanical control schemes at lower levels of organization.

Web Vibrations in Intraspecific Contests of Female Black Widow Spiders, Latrodectus hesperus.

Female black widow spiders, Latrodectus hesperus Chamberlin and Ivie (Araneae: Theridiidae), are solitary predators of arthropods with no tolerance for intruders on the webs. In California, L. hesperus are found in urban and agricultural settings and can be a phytosanitary pest in fresh produce. Spatial separation of L. hesperus webs could be determined by seasonal population densities, with territorial competition expected under high densities in the environment. However, little is known about female-female communication behaviors in this species. In 1-hr laboratory observations, displays of female-female rivalry included production of vibrational signals in a majority (20 of 30) of trials. The number of signals produced by both females was highest during the initial 10 min of trials, with signaling rate (time interval between signals) peaking during the 40-50 min observation period. The overall ratio of signals produced by the resident female and the introduced female was about 5:1, with the number of signals produced by the resident female higher than the number of signals produced by the introduced female. Analysis of rivalry signals showed a peak in magnitude (about 0.4 m/s) ranging from 6 to 23 Hz and smaller peaks at about 29, 38, and 47 Hz. Collectively, these results demonstrate that female L. hesperus exhibit territorial rivalry and that female-female rivalry is mediated by emission of vibrational signals through the web. Understanding the mechanisms of intraspecific competition in L. hesperus is required for elucidating interspecific interactions in the environment and may lead to development of novel methods to prevent spiders from colonizing crops.

Curvilinear Kirigami Skins Let Soft Bending Actuators Slither Faster.

The locomotion of soft snake robots is dependent on frictional interactions with the environment. Frictional anisotropy is a morphological characteristic of snakeskin that allows snakes to engage selectively with surfaces and generate propulsive forces. The prototypical slithering gait of most snakes is lateral undulation, which requires a significant lateral resistance that is lacking in artificial skins of existing soft snake robots. We designed a set of kirigami lattices with curvilinearly-arranged cuts to take advantage of in-plane rotations of the 3D structures when wrapped around a soft bending actuator. By changing the initial orientation of the scales, the kirigami skin produces high lateral friction upon engagement with surface asperities, with lateral to cranial anisotropic friction ratios above 4. The proposed design increased the overall velocity of the soft snake robot more than fivefold compared to robots without skin.

Validation of a Novel Stereo Vibrometry Technique for Spiderweb Signal Analysis.

From courtship rituals, to prey identification, to displays of rivalry, a spider's web vibrates with a symphony of information. Examining the modality of information being transmitted and how spiders interact with this information could lead to new understanding how spiders perceive the world around them through their webs, and new biological and engineering techniques that leverage this understanding. Spiders interact with their webs through a variety of body motions, including abdominal tremors, bounces, and limb jerks along threads of the web. These signals often create a large enough visual signature that the web vibrations can be analyzed using video vibrometry on high-speed video of the communication exchange. Using video vibrometry to examine these signals has numerous benefits over the conventional method of laser vibrometry, such as the ability to analyze three-dimensional vibrations and the ability to take measurements from anywhere in the web, including directly from the body of the spider itself. In this study, we developed a method of three-dimensional vibration analysis that combines video vibrometry with stereo vision, and verified this method against laser vibrometry on a black widow spiderweb that was experiencing rivalry signals from two female spiders.

Data-driven geometric system identification for shape-underactuated dissipative systems.

Modeling system dynamics becomes challenging when the properties of individual system components cannot be directly measured, and often requires identification of properties from observed motion. In this paper, we show that systems whose movement is highly dissipative have features which provide an opportunity to more easily identify models and more quickly optimize motions than would be possible with general techniques. Geometric mechanics provides means for reduction of the dynamics by environmental homogeneity, while the dissipative nature minimizes the role of second order (inertial) features in the dynamics. Here we extend the tools of geometric system identification to 'shape-underactuated dissipative systems (SUDS)'-systems whose motions are more dissipative than inertial, but whose actuation is restricted to a subset of the body shape coordinates. Many animal motions are SUDS, including micro-swimmers such as nematodes and flagellated bacteria, and granular locomotors such as snakes and lizards. Many soft robots are also SUDS, particularly robots that incorporate highly damped series elastic actuators to reduce the rigidity of their interactions with their environments during locomotion and manipulation. We motivate the use of SUDS models, and validate their ability to predict motion of a variety of simulated viscous swimming platforms. For a large class of SUDS, we show how the shape velocity actuation inputs can be directly converted into torque inputs, suggesting that systems with soft pneumatic or dielectric elastomer actuators can be modeled with the tools presented. Based on fundamental assumptions in the physics, we show how our model complexity scales linearly with the number of passive shape coordinates. This scaling offers a large reduction on the number of trials needed to identify the system model from experimental data, and may reduce overfitting. The sample efficiency of our method suggests its use in modeling, control, and optimization in robotics, and as a tool for the study of organismal motion in friction dominated regimes.

Optimal gaits for drag-dominated swimmers with passive elastic joints.

In this paper we identify optimal swimming strategies for drag-dominated swimmers with a passive elastic joint. We use resistive force theory to obtain the dynamics of the system. We then use frequency-domain analysis to relate the motion of the passive joint to the motion of the actuated joint. We couple this analysis with elements of the geometric framework introduced in our previous work aimed at identifying useful gaits for systems in drag-dominated environments to identify speed-maximizing and efficiency-maximizing gaits for drag-dominated swimmers with a passive elastic joint.

Surprising simplicities and syntheses in limbless self-propulsion in sand.

Animals moving on and in fluids and solids move their bodies in diverse ways to generate propulsion and lift forces. In fluids, animals can wiggle, stroke, paddle or slap, whereas on hard frictional terrain, animals largely engage their appendages with the substrate to avoid slip. Granular substrates, such as desert sand, can display complex responses to animal interactions. This complexity has led to locomotor strategies that make use of fluid-like or solid-like features of this substrate, or combinations of the two. Here, we use examples from our work to demonstrate the diverse array of methods used and insights gained in the study of both surface and subsurface limbless locomotion in these habitats. Counterintuitively, these seemingly complex granular environments offer certain experimental, theoretical, robotic and computational advantages for studying terrestrial movement, with the potential for providing broad insights into morphology and locomotor control in fluids and solids, including neuromechanical control templates and morphological and behavioral evolution. In particular, granular media provide an excellent testbed for a locomotion framework called geometric mechanics, which was introduced by particle physicists and control engineers in the last century, and which allows quantitative analysis of alternative locomotor patterns and morphology to test for control templates, optimality and evolutionary alternatives. Thus, we posit that insights gained from movement in granular environments can be translated into principles that have broader applications across taxa, habitats and movement patterns, including those at microscopic scales.

A review on locomotion robophysics: the study of movement at the intersection of robotics, soft matter and dynamical systems.

Discovery of fundamental principles which govern and limit effective locomotion (self-propulsion) is of intellectual interest and practical importance. Human technology has created robotic moving systems that excel in movement on and within environments of societal interest: paved roads, open air and water. However, such devices cannot yet robustly and efficiently navigate (as animals do) the enormous diversity of natural environments which might be of future interest for autonomous robots; examples include vertical surfaces like trees and cliffs, heterogeneous ground like desert rubble and brush, turbulent flows found near seashores, and deformable/flowable substrates like sand, mud and soil. In this review we argue for the creation of a physics of moving systems-a 'locomotion robophysics'-which we define as the pursuit of principles of self-generated motion. Robophysics can provide an important intellectual complement to the discipline of robotics, largely the domain of researchers from engineering and computer science. The essential idea is that we must complement the study of complex robots in complex situations with systematic study of simplified robotic devices in controlled laboratory settings and in simplified theoretical models. We must thus use the methods of physics to examine both locomotor successes and failures using parameter space exploration, systematic control, and techniques from dynamical systems. Using examples from our and others' research, we will discuss how such robophysical studies have begun to aid engineers in the creation of devices that have begun to achieve life-like locomotor abilities on and within complex environments, have inspired interesting physics questions in low dimensional dynamical systems, geometric mechanics and soft matter physics, and have been useful to develop models for biological locomotion in complex terrain. The rapidly decreasing cost of constructing robot models with easy access to significant computational power bodes well for scientists and engineers to engage in a discipline which can readily integrate experiment, theory and computation.

BIOMECHANICS. Why the seahorse tail is square.

Whereas the predominant shapes of most animal tails are cylindrical, seahorse tails are square prisms. Seahorses use their tails as flexible grasping appendages, in spite of a rigid bony armor that fully encases their bodies. We explore the mechanics of two three-dimensional-printed models that mimic either the natural (square prism) or hypothetical (cylindrical) architecture of a seahorse tail to uncover whether or not the square geometry provides any functional advantages. Our results show that the square prism is more resilient when crushed and provides a mechanism for preserving articulatory organization upon extensive bending and twisting, as compared with its cylindrical counterpart. Thus, the square architecture is better than the circular one in the context of two integrated functions: grasping ability and crushing resistance.

Sidewinding with minimal slip: snake and robot ascent of sandy slopes.

Limbless organisms such as snakes can navigate nearly all terrain. In particular, desert-dwelling sidewinder rattlesnakes (Crotalus cerastes) operate effectively on inclined granular media (such as sand dunes) that induce failure in field-tested limbless robots through slipping and pitching. Our laboratory experiments reveal that as granular incline angle increases, sidewinder rattlesnakes increase the length of their body in contact with the sand. Implementing this strategy in a physical robot model of the snake enables the device to ascend sandy slopes close to the angle of maximum slope stability. Plate drag experiments demonstrate that granular yield stresses decrease with increasing incline angle. Together, these three approaches demonstrate how sidewinding with contact-length control mitigates failure on granular media.

Geometric visualization of self-propulsion in a complex medium.

Combining geometric mechanics theory, laboratory robotic experiment, and numerical simulation, we study the locomotion in granular media of the simplest noninertial swimmer, the Purcell three-link swimmer. Using granular resistive force laws as inputs, the theory relates translation and rotation of the body to shape changes (movements of the links). This allows analysis, visualization, and prediction of effective movements that are verified by experiment. The geometric approach also facilitates comparison between swimming in granular media and in viscous fluids.