Walking is like slithering: A unifying, data-driven view of locomotion.

Legged movement is ubiquitous in nature and of increasing interest for robotics. Most legged animals routinely encounter foot slipping, yet detailed modeling of multiple contacts with slipping exceeds current simulation capacity. Here we present a principle that unifies multilegged walking (including that involving slipping) with slithering and Stokesian (low Reynolds number) swimming. We generated data-driven principally kinematic models of locomotion for walking in low-slip animals (Argentine ant, 4.7% slip ratio of slipping to total motion) and for high-slip robotic systems (BigANT hexapod, slip ratio 12 to 22%; Multipod robots ranging from 6 to 12 legs, slip ratio 40 to 100%). We found that principally kinematic models could explain much of the variability in body velocity and turning rate using body shape and could predict walking behaviors outside the training data. Most remarkably, walking was principally kinematic irrespective of leg number, foot slipping, and turning rate. We find that grounded walking, with or without slipping, is governed by principally kinematic equations of motion, functionally similar to frictional swimming and slithering. Geometric mechanics thus leads to a unified model for swimming, slithering, and walking. Such commonality may shed light on the evolutionary origins of animal locomotion control and offer new approaches for robotic locomotion and motion planning.

Phase response curves and the role of coordinates.

The "infinitesimal phase response curve" (PRC) is a common tool used to analyze phase resetting in the natural sciences in general and neuroscience in particular. We make the observation that the PRC with respect to a coordinate v actually depends on the choice of other coordinates. As a consequence, a complete delay embedding reconstruction of the dynamics using v which would allow phase to be computed still does not allow the v PRC to be computed. We give a coordinate-free definition of the PRC making this observation obvious. This leads to an experimental protocol: first collect an appropriate ensemble of measurements by intermittently controlling neuron voltage. Then, for any suitable current carrier dynamic postulated, we show how the ensemble can be used to compute the voltage PRC with that current carrier. The approach extends to many oscillators measured and controlled through a subset of their coordinates.

Longitudinal quasi-static stability predicts changes in dog gait on rough terrain.

Legged animals utilize gait selection to move effectively and must recover from environmental perturbations. We show that on rough terrain, domestic dogs, Canis lupus familiaris, spend more time in longitudinal quasi-statically stable patterns of movement. Here, longitudinal refers to the rostro-caudal axis. We used an existing model in the literature to quantify the longitudinal quasi-static stability of gaits neighbouring the walk, and found that trot-like gaits are more stable. We thus hypothesized that when perturbed, the rate of return to a stable gait would depend on the direction of perturbation, such that perturbations towards less quasi-statically stable patterns of movement would be more rapid than those towards more stable patterns of movement. The net result of this would be greater time spent in longitudinally quasi-statically stable patterns of movement. Limb movement patterns in which diagonal limbs were more synchronized (those more like a trot) have higher longitudinal quasi-static stability. We therefore predicted that as dogs explored possible limb configurations on rough terrain at walking speeds, the walk would shift towards trot. We gathered experimental data quantifying dog gait when perturbed by rough terrain and confirmed this prediction using GPS and inertial sensors (n=6, P<0.05). By formulating gaits as trajectories on the n-torus we are able to make tractable the analysis of gait similarity. These methods can be applied in a comparative study of gait control which will inform the ultimate role of the constraints and costs impacting locomotion, and have applications in diagnostic procedures for gait abnormalities, and in the development of agile legged robots.

Morphology and the gradient of a symmetric potential predict gait transitions of dogs.

Gaits and gait transitions play a central role in the movement of animals. Symmetry is thought to govern the structure of the nervous system, and constrain the limb motions of quadrupeds. We quantify the symmetry of dog gaits with respect to combinations of bilateral, fore-aft, and spatio-temporal symmetry groups. We tested the ability of symmetries to model motion capture data of dogs walking, trotting and transitioning between those gaits. Fully symmetric models performed comparably to asymmetric with only a [Formula: see text] increase in the residual sum of squares and only one-quarter of the parameters. This required adding a spatio-temporal shift representing a lag between fore and hind limbs. Without this shift, the symmetric model residual sum of squares was [Formula: see text] larger. This shift is related to (linear regression, [Formula: see text], [Formula: see text]) dog morphology. That this symmetry is respected throughout the gaits and transitions indicates that it generalizes outside a single gait. We propose that relative phasing of limb motions can be described by an interaction potential with a symmetric structure. This approach can be extended to the study of interaction of neurodynamic and kinematic variables, providing a system-level model that couples neuronal central pattern generator networks and mechanical models.

Thermal adaptation and phosphorus shape thermal performance in an assemblage of rainforest ants.

We studied the Thermal Performance Curves (TPCs) of 87 species of rainforest ants and found support for both the Thermal Adaptation and Phosphorus-Tolerance hypotheses. TPCs relate a fitness proxy (here, worker speed) to environmental temperature. Thermal Adaptation posits that thermal generalists (ants with flatter, broader TPCs) are favored in the hotter, more variable tropical canopy compared to the cooler, less variable litter below. As predicted, species nesting in the forest canopy 1) had running speeds less sensitive to temperature; 2) ran over a greater range of temperatures; and 3) ran at lower maximum speeds. Tradeoffs between tolerance and maximum performance are often invoked for constraining the evolution of thermal generalists. There was no evidence that ant species traded off thermal tolerance for maximum speed, however. Phosphorus-Tolerance is a second mechanism for generating ectotherms able to tolerate thermal extremes. It posits that ants active at high temperatures invest in P-rich machinery to buffer their metabolism against thermal extremes. Phosphorus content in ant tissue varied three-fold, and as predicted, temperature sensitivity was lower and thermal range was higher in P-rich species. Combined, we show how the vertical distribution of hot and variable vs. cooler and stable microclimates in a single forest contribute to a diversity of TPCs and suggest that a widely varying P stoichiometry among these ants may drive some of these differences.

Instantaneous kinematic phase reflects neuromechanical response to lateral perturbations of running cockroaches.

Instantaneous kinematic phase calculation allows the development of reduced-order oscillator models useful in generating hypotheses of neuromechanical control. When perturbed, changes in instantaneous kinematic phase and frequency of rhythmic movements can provide details of movement and evidence for neural feedback to a system-level neural oscillator with a time resolution not possible with traditional approaches. We elicited an escape response in cockroaches (Blaberus discoidalis) that ran onto a movable cart accelerated laterally with respect to the animals' motion causing a perturbation. The specific impulse imposed on animals (0.50 [Formula: see text] 0.04 m s[Formula: see text]; mean, SD) was nearly twice their forward speed (0.25 [Formula: see text] 0.06 m s[Formula: see text]. Instantaneous residual phase computed from kinematic phase remained constant for 110 ms after the onset of perturbation, but then decreased representing a decrease in stride frequency. Results from direct muscle action potential recordings supported kinematic phase results in showing that recovery begins with self-stabilizing mechanical feedback followed by neural feedback to an abstracted neural oscillator or central pattern generator. Trials fell into two classes of forward velocity changes, while exhibiting statistically indistinguishable frequency changes. Animals pulled away from the side with front and hind legs of the tripod in stance recovered heading within 300 ms, whereas animals that only had a middle leg of the tripod resisting the pull did not recover within this period. Animals with eight or more legs might be more robust to lateral perturbations than hexapods.

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.

Active tails enhance arboreal acrobatics in geckos.

Geckos are nature's elite climbers. Their remarkable climbing feats have been attributed to specialized feet with hairy toes that uncurl and peel in milliseconds. Here, we report that the secret to the gecko's arboreal acrobatics includes an active tail. We examine the tail's role during rapid climbing, aerial descent, and gliding. We show that a gecko's tail functions as an emergency fifth leg to prevent falling during rapid climbing. A response initiated by slipping causes the tail tip to push against the vertical surface, thereby preventing pitch-back of the head and upper body. When pitch-back cannot be prevented, geckos avoid falling by placing their tail in a posture similar to a bicycle's kickstand. Should a gecko fall with its back to the ground, a swing of its tail induces the most rapid, zero-angular momentum air-righting response yet measured. Once righted to a sprawled gliding posture, circular tail movements control yaw and pitch as the gecko descends. Our results suggest that large, active tails can function as effective control appendages. These results have provided biological inspiration for the design of an active tail on a climbing robot, and we anticipate their use in small, unmanned gliding vehicles and multisegment spacecraft.

Challenges in dynamic mode decomposition.

Dynamic mode decomposition (DMD) is a powerful tool for extracting spatial and temporal patterns from multi-dimensional time series, and it has been used successfully in a wide range of fields, including fluid mechanics, robotics and neuroscience. Two of the main challenges remaining in DMD research are noise sensitivity and issues related to Krylov space closure when modelling nonlinear systems. Here, we investigate the combination of noise and nonlinearity in a controlled setting, by studying a class of systems with linear latent dynamics which are observed via multinomial observables. Our numerical models include system and measurement noise. We explore the influences of dataset metrics, the spectrum of the latent dynamics, the normality of the system matrix and the geometry of the dynamics. Our results show that even for these very mildly nonlinear conditions, DMD methods often fail to recover the spectrum and can have poor predictive ability. Our work is motivated by our experience modelling multilegged robot data, where we have encountered great difficulty in reconstructing time series for oscillatory systems with intermediate transients, which decay only slightly faster than a period.

Insects running on elastic surfaces.

In nature, cockroaches run rapidly over complex terrain such as leaf litter. These substrates are rarely rigid, and are frequently very compliant. Whether and how compliant surfaces change the dynamics of rapid insect locomotion has not been investigated to date largely due to experimental limitations. We tested the hypothesis that a running insect can maintain average forward speed over an extremely soft elastic surface (10 N m(-1)) equal to 2/3 of its virtual leg stiffness (15 N m(-1)). Cockroaches Blaberus discoidalis were able to maintain forward speed (mean +/- s.e.m., 37.2+/-0.6 cm s(-1) rigid surface versus 38.0+/-0.7 cm s(-1) elastic surface; repeated-measures ANOVA, P=0.45). Step frequency was unchanged (24.5+/-0.6 steps s(-1) rigid surface versus 24.7+/-0.4 steps s(-1) elastic surface; P=0.54). To uncover the mechanism, we measured the animal's centre of mass (COM) dynamics using a novel accelerometer backpack, attached very near the COM. Vertical acceleration of the COM on the elastic surface had a smaller peak-to-peak amplitude (11.50+/-0.33 m s(-2), rigid versus 7.7+/-0.14 m s(-2), elastic; P=0.04). The observed change in COM acceleration over an elastic surface required no change in effective stiffness when duty factor and ground stiffness were taken into account. Lowering of the COM towards the elastic surface caused the swing legs to land earlier, increasing the period of double support. A feedforward control model was consistent with the experimental results and provided one plausible, simple explanation of the mechanism.

Multi-legged steering and slipping with low DoF hexapod robots.

Thanks to their sprawled posture and multi-legged support, stability is not as hard to achieve for hexapedal robots as it is for bipeds and quadrupeds. A key engineering challenge with hexapods has been to produce insect-like agility and maneuverability, of which steering is an essential part. However, the mechanisms of multi-legged steering are not always clear, especially for robots with underactuated legs. Here we propose a formal definition of steering, and show why steering is difficult for robots with 6 or more underactuated legs. We show that for many of these robots, steering is impossible without slipping, and present experimental results which demonstrate the importance of allowing for slipping to occur intentionally when optimizing steering ability. Our results suggest that a non-holonomic multi-legged slipping model might be more appropriate than dynamic models for representing such robots, and that conventional non-slip contact models might miss significant parts of the performance envelope.

Estimating the phase of synchronized oscillators.

The state of a collection of phase-locked oscillators is determined by a single phase variable or cyclic coordinate. This paper presents a computational method, Phaser, for estimating the phase of phase-locked oscillators from limited amounts of multivariate data in the presence of noise and measurement errors. Measurements are assumed to be a collection of multidimensional time series. Each series consists of several cycles of the same or similar systems. The oscillators within each system are not assumed to be identical. Using measurements of the noise covariance for the multivariate input, data from the individual oscillators in the system are combined to reduce the variance of phase estimates for the whole system. The efficacy of the algorithm is demonstrated on experimental and model data from biomechanics of cockroach running and on simulated oscillators with varying levels of noise.

Constructing predictive models of human running.

Running is an essential mode of human locomotion, during which ballistic aerial phases alternate with phases when a single foot contacts the ground. The spring-loaded inverted pendulum (SLIP) provides a starting point for modelling running, and generates ground reaction forces that resemble those of the centre of mass (CoM) of a human runner. Here, we show that while SLIP reproduces within-step kinematics of the CoM in three dimensions, it fails to reproduce stability and predict future motions. We construct SLIP control models using data-driven Floquet analysis, and show how these models may be used to obtain predictive models of human running with six additional states comprising the position and velocity of the swing-leg ankle. Our methods are general, and may be applied to any rhythmic physical system. We provide an approach for identifying an event-driven linear controller that approximates an observed stabilization strategy, and for producing a reduced-state model which closely recovers the observed dynamics.

Finding the dimension of slow dynamics in a rhythmic system.

Dynamical systems with asymptotically stable periodic orbits are generic models for rhythmic processes in dissipative physical systems. This paper presents a method for reconstructing the dynamics near a periodic orbit from multivariate time-series data. It is used to test theories about the control of legged locomotion, a context in which time series are short when compared with previous work in nonlinear time-series analysis. The method presented here identifies appropriate dimensions of reduced order models for the deterministic portion of the dynamics. The paper also addresses challenges inherent in identifying dynamical models with data from different individuals.