RESUMEN
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.
Asunto(s)
Lagartos , Locomoción , Animales , Locomoción/fisiología , Lagartos/fisiología , Serpientes/fisiología , Fenómenos Biomecánicos , Modelos BiológicosRESUMEN
Locomotion is typically studied either in continuous media where bodies and legs experience forces generated by the flowing medium or on solid substrates dominated by friction. In the former, centralized whole-body coordination is believed to facilitate appropriate slipping through the medium for propulsion. In the latter, slip is often assumed minimal and thus avoided via decentralized control schemes. We find in laboratory experiments that terrestrial locomotion of a meter-scale multisegmented/legged robophysical model resembles undulatory fluid swimming. Experiments varying waves of leg stepping and body bending reveal how these parameters result in effective terrestrial locomotion despite seemingly ineffective isotropic frictional contacts. Dissipation dominates over inertial effects in this macroscopic-scaled regime, resulting in essentially geometric locomotion on land akin to microscopic-scale swimming in fluids. Theoretical analysis demonstrates that the high-dimensional multisegmented/legged dynamics can be simplified to a centralized low-dimensional model, which reveals an effective resistive force theory with an acquired viscous drag anisotropy. We extend our low-dimensional, geometric analysis to illustrate how body undulation can aid performance in non-flat obstacle-rich terrains and also use the scheme to quantitatively model how body undulation affects performance of biological centipede locomotion (the desert centipede Scolopendra polymorpha) moving at relatively high speeds (â¼0.5 body lengths/sec). Our results could facilitate control of multilegged robots in complex terradynamic scenarios.
RESUMEN
Although typically possessing four limbs and short bodies, lizards have evolved diverse morphologies, including elongate trunks with tiny limbs. Such forms are hypothesized to aid locomotion in cluttered/fossorial environments but propulsion mechanisms (e.g., the use of body and/or limbs to interact with substrates) and potential body/limb coordination remain unstudied. Here, we use biological experiments, a geometric theory of locomotion, and robophysical models to investigate body-limb coordination in diverse lizards. Locomotor field studies in short-limbed, elongate lizards (Brachymeles and Lerista) and laboratory studies of fully limbed lizards (Uma scoparia and Sceloporus olivaceus) and a snake (Chionactis occipitalis) reveal that body-wave dynamics can be described by a combination of standing and traveling waves; the ratio of the amplitudes of these components is inversely related to the degree of limb reduction and body elongation. The geometric theory (which replaces laborious calculation with diagrams) helps explain our observations, predicting that the advantage of traveling-wave body undulations (compared with a standing wave) emerges when the dominant thrust-generation mechanism arises from the body rather than the limbs and reveals that such soil-dwelling lizards propel via "terrestrial swimming" like sand-swimming lizards and snakes. We test our hypothesis by inducing the use of traveling waves in stereotyped lizards via modulating the ground-penetration resistance. Study of a limbed/undulatory robophysical model demonstrates that a traveling wave is beneficial when propulsion is generated by body-environment interaction. Our models could be valuable in understanding functional constraints on the evolutionary processes of elongation and limb reduction as well as advancing robot designs.
Asunto(s)
Lagartos , Natación , Animales , Evolución Biológica , Extremidades/anatomía & histología , Extremidades/fisiología , Lagartos/anatomía & histología , Lagartos/fisiología , Serpientes/anatomía & histología , Serpientes/fisiologíaRESUMEN
Centipedes coordinate body and limb flexion to generate propulsion. On flat, solid surfaces, the limb-stepping patterns can be characterized according to the direction in which limb-aggregates propagate, opposite to (retrograde) or with the direction of motion (direct). It is unknown how limb and body dynamics are modified in terrain with terradynamic complexity more representative of these animal's natural heterogeneous environments. Here, we investigated how centipedes that use retrograde and direct limb-stepping patterns, Scolopendra polymorpha and Scolopocryptops sexspinosus, respectively, coordinate their body and limbs to navigate laboratory environments which present footstep challenges and terrain rugosity. We recorded the kinematics and measured the locomotive performance of these animals traversing two rugose terrains with randomly distributed step heights and compared the kinematics with those on a flat frictional surface. Scolopendra polymorpha exhibited similar body and limb dynamics across all terrains and a decrease in speed with increased terrain rugosity. Unexpectedly, when placed in a rugose terrain, S. sexspinosus changed the direction of the limb-stepping pattern from direct to retrograde. Further, for both species, traversal of these rugose terrains was facilitated by hypothesized passive mechanics: upon horizontal collision of a limb with a block, the limb bent and later continued the stepping pattern. Although centipedes have many degrees of freedom, our results suggest these animals negotiate limb-substrate interactions and navigate complex terrains leveraging the innate flexibility of their limbs to simplify control.
Asunto(s)
Quilópodos , Extremidades , Animales , Fenómenos Biomecánicos , Ambiente , Locomoción , MarchaRESUMEN
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.
Asunto(s)
Lagartos/fisiología , Locomoción , Arena , Serpientes/fisiología , Animales , Fenómenos Biomecánicos , Extremidades/anatomía & histologíaRESUMEN
Colloidal particles with asymmetric surface chemistry (Janus particles) have unique bifunctional properties. The size of these particles is an important determinant for their applications in diverse fields from drug delivery to chemical catalysis. The size of Janus particles, with a core surface coated with carboxylate and a partially encapsulating silica shell, depends upon several factors, including the core size and the concentration of carboxylate coating. The role of the carboxylate coating on the Janus particle size is well-understood; however, the role of the core size is not well defined. The role of the carboxylated polystyrene (cPS) core size on the cPS-silica Janus particle morphology (its size and shape) was examined by testing two different silica sizes and five different cPS core sizes. Results from electron microscopy (EM) and dynamic light scattering (DLS) analysis indicate that the composite cPS-silica particle acquires two distinct shapes: (i) when the size of the cPS core is much smaller than the non-cPS silica (b-SiO2) sphere, partially encapsulated Janus particles are formed, and (ii) when the cPS core is larger than or equal to the b-SiO2 sphere, a raspberry-like structure rather than a Janus particle is formed. The cPS-silica Janus particles of â¼100-500 nm size were obtained when the size of the cPS core was much smaller than the non-cPS silica (b-SiO2) sphere. These scalable nanoscale Janus particles will have wide application in a multifunctional delivery platform and catalysis.
Asunto(s)
Sistemas de Liberación de Medicamentos , Poliestirenos/química , Dióxido de Silicio/química , Coloides , Tamaño de la PartículaRESUMEN
Whereas the transport of matter by wheeled vehicles or legged robots can be guaranteed in engineered landscapes such as roads or rails, locomotion prediction in complex environments such as collapsed buildings or crop fields remains challenging. Inspired by the principles of information transmission, which allow signals to be reliably transmitted over "noisy" channels, we developed a "matter-transport" framework that demonstrates that noninertial locomotion can be provably generated over noisy rugose landscapes (heterogeneities on the scale of locomotor dimensions). Experiments confirm that sufficient spatial redundancy in the form of serially connected legged robots leads to reliable transport on such terrain without requiring sensing and control. Further analogies from communication theory coupled with advances in gaits (coding) and sensor-based feedback control (error detection and correction) can lead to agile locomotion in complex terradynamic regimes.
RESUMEN
Limbless locomotors, from microscopic worms to macroscopic snakes, traverse complex, heterogeneous natural environments typically using undulatory body wave propagation. Theoretical and robophysical models typically emphasize body kinematics and active neural/electronic control. However, we contend that because such approaches often neglect the role of passive, mechanically controlled processes (those involving "mechanical intelligence"), they fail to reproduce the performance of even the simplest organisms. To uncover principles of how mechanical intelligence aids limbless locomotion in heterogeneous terradynamic regimes, here we conduct a comparative study of locomotion in a model of heterogeneous terrain (lattices of rigid posts). We used a model biological system, the highly studied nematode worm Caenorhabditis elegans, and a robophysical device whose bilateral actuator morphology models that of limbless organisms across scales. The robot's kinematics quantitatively reproduced the performance of the nematodes with purely open-loop control; mechanical intelligence simplified control of obstacle navigation and exploitation by reducing the need for active sensing and feedback. An active behavior observed in C. elegans, undulatory wave reversal upon head collisions, robustified locomotion via exploitation of the systems' mechanical intelligence. Our study provides insights into how neurally simple limbless organisms like nematodes can leverage mechanical intelligence via appropriately tuned bilateral actuation to locomote in complex environments. These principles likely apply to neurally more sophisticated organisms and also provide a design and control paradigm for limbless robots for applications like search and rescue and planetary exploration.
Asunto(s)
Caenorhabditis elegans , Robótica , Animales , Locomoción , Serpientes , Fenómenos BiomecánicosRESUMEN
Serially connected robots are promising candidates for performing tasks in confined spaces such as search and rescue in large-scale disasters. Such robots are typically limbless, and we hypothesize that the addition of limbs could improve mobility. However, a challenge in designing and controlling such devices lies in the coordination of high-dimensional redundant modules in a way that improves mobility. Here we develop a general framework to discover templates to control serially connected multi-legged robots. Specifically, we combine two approaches to build a general shape control scheme which can provide baseline patterns of self-deformation ('gaits') for effective locomotion in diverse robot morphologies. First, we take inspiration from a dimensionality reduction and a biological gait classification scheme to generate cyclic patterns of body deformation and foot lifting/lowering, which facilitate the generation of arbitrary substrate contact patterns. Second, we extend geometric mechanics, which was originally introduced to study swimming at low Reynolds numbers, to frictional environments, allowing the identification of optimal body-leg coordination in this common terradynamic regime. Our scheme allows the development of effective gaits on flat terrain with diverse numbers of limbs (4, 6, 16, and even 0 limbs) and backbone actuation. By properly coordinating the body undulation and leg placement, our framework combines the advantages of both limbless robots (modularity and narrow profile) and legged robots (mobility). Our framework can provide general control schemes for the rapid deployment of general multi-legged robots, paving the way toward machines that can traverse complex environments. In addition, we show that our framework can also offer insights into body-leg coordination in living systems, such as salamanders and centipedes, from a biomechanical perspective.