Turtling the salamander: tail movements mitigate need for kinematic limb changes during walking in tiger salamanders (Ambystoma tigrinum) with restricted lateral movement

Lateral undulation and trunk flexibility offer performance benefits to maneuverability, stability, and stride length (via speed and distance traveled). These benefits make them key characteristics of the locomotion of tetrapods with sprawling posture, with the exception of turtles. Despite their bony carapace preventing lateral undulations, turtles are able to improve their locomotor performance by increasing stride length via greater limb protraction. The goal of this study was to quantify the effect of reduced lateral flexibility in a generalized sprawling tetrapod, the tiger salamander (Ambystoma tigrinum). We had two potential predictions: (1) either salamanders completely compensate by changing their limb kinematics, or (2) their performance (i.e., speed) will suffer due to the reduced lateral flexibility. This reduction was performed by artificially limiting trunk flexibility by attaching a 2-piece shell around the body between the pectoral and pelvic girdles. Adult tiger salamanders (n = 3, SVL = 9 cm-14.5 cm) walked on a 1 m trackway under three different conditions: unrestricted, flexible shell (Tygon tubing), and rigid shell (PVC tubing). Trials were filmed in a single, dorsal view, and kinematics of entire midline and specific body regions (head, trunk, tail), as well as the fore and hindlimbs, were calculated. Tygon individuals had significantly higher curvature than both PVC and unrestricted individuals for the body, but this trend was primarily driven by changes in tail movements. PVC individuals had significantly lower curvature in the trunk region compared to unrestricted individuals or Tygon; however, there was no difference between unrestricted and Tygon individuals suggesting the shells performed as expected. PVC and Tygon individuals had significantly higher curvature in the tails compared to unrestricted individuals. There were no significant differences for any limb kinematic variables among treatments including average, minimum, maximum angles. Thus, salamanders respond to decreased lateral movement in their trunk by increasing movements in their tail, without changes in limb kinematics. These results suggest that tail undulations may be a more critical component to sprawling-postured tetrapod locomotion than previously recognized.

4. How amphibians move

‘How amphibians move’ examines how amphibians move. The three kinds of living amphibians share the same basic biology and life history. However, the anatomy of the skeleton and muscles is very different amongst them. This reflects the different ways that the locomotion of the three respective ancestors evolved. The urodeles retained the most primitive way, with a long body and tail. Salamanders and newts use lateral undulation when swimming, but they also coupled it with limbs for walking on land. The anurans became far more modified by shortening the body, losing the tail altogether, and elongating the back legs. Meanwhile, the caecilians evolved a limbless burrowing mode.

Slithering CSF: Cerebrospinal Fluid Dynamics in the Stationary and Moving Viper Boa, Candoia aspera

In the viper boa (Candoia aspera), the cerebrospinal fluid (CSF) shows two stable overlapping patterns of pulsations: low-frequency (0.08 Hz) pulses with a mean amplitude of 4.1 mmHg that correspond to the ventilatory cycle, and higher-frequency (0.66 Hz) pulses with a mean amplitude of 1.2 mmHg that correspond to the cardiac cycle. Manual oscillations of anesthetized C. aspera induced propagating sinusoidal body waves. These waves resulted in a different pattern of CSF pulsations with frequencies corresponding to the displacement frequency of the body and with amplitudes greater than those of the cardiac or ventilatory cycles. After recovery from anesthesia, the snakes moved independently using lateral undulation and concertina locomotion. The episodes of lateral undulation produced similar influences on the CSF pressure as were observed during the manual oscillations, though the induced CSF pulsations were of lower amplitude during lateral undulation. No impact on the CSF was found while C. aspera was performing concertina locomotion. The relationship between the propagation of the body and the CSF pulsations suggests that the body movements produce an impulse on the spinal CSF.

Generation of propulsive force via vertical undulations in snakes

Lateral undulation is the most widespread mode of terrestrial vertebrate limbless locomotion, in which posteriorly propagating horizontal waves press against environmental asperities (e.g. grass, rocks) and generate propulsive reaction forces. We hypothesized that snakes can generate propulsion using a similar mechanism of posteriorly propagating vertical waves pressing against suitably oriented environmental asperities. Using an array of horizontally oriented cylinders, one of which was equipped with force sensors, and a motion capture system, we found snakes generated substantial propulsive force and propulsive impulse with minimal contribution from lateral undulation. Additional tests showed that snakes could propel themselves via vertical undulations from a single suitable contact point, and this mechanism was replicated in a robotic model. Vertical undulations can provide snakes a valuable locomotor tool for taking advantage of vertical asperities in a variety of habitats, potentially in combination with lateral undulation, to fully exploit the 3D structure of the habitat.

Functional consequences of convergently evolved microscopic skin features on snake locomotion

The small structures that decorate biological surfaces can significantly affect behavior, yet the diversity of animal–environment interactions essential for survival makes ascribing functions to structures challenging. Microscopic skin textures may be particularly important for snakes and other limbless locomotors, where substrate interactions are mediated solely through body contact. While previous studies have characterized ventral surface features of some snake species, the functional consequences of these textures are not fully understood. Here, we perform a comparative study, combining atomic force microscopy measurements with mathematical modeling to generate predictions that link microscopic textures to locomotor performance. We discover an evolutionary convergence in the ventral skin structures of a few sidewinding specialist vipers that inhabit sandy deserts—an isotropic texture that is distinct from the head-to-tail-oriented, micrometer-sized spikes observed on a phylogenetically broad sampling of nonsidewinding vipers and other snakes from diverse habitats and wide geographic range. A mathematical model that relates structural directionality to frictional anisotropy reveals that isotropy enhances movement during sidewinding, whereas anisotropy improves movement during slithering via lateral undulation of the body. Our results highlight how an integrated approach can provide quantitative predictions for structure–function relationships and insights into behavioral and evolutionary adaptations in biological systems.

Lateral Undulation Aids Biological and Robotic Earthworm Anchoring and Locomotion

Earthworms (Lumbricus terrestris) are characterized by soft, highly flexible and extensible bodies, and are capable of locomoting in most terrestrial environments. Previous studies of earthworm movement have focused on the use of retrograde peristaltic gaits in which controlled contraction of longitudinal and circular muscles results in waves of shortening/thickening and thinning/lengthening of the hydrostatic skeleton. These waves can propel the animal across ground as well as into soil. However, worms can also benefit from axial body bends during locomotion. Such lateral undulation dynamics can aid locomotor function via hooking/anchoring (to provide propulsion), modify travel orientation (to avoid obstacles and generate turns) and even generate snake-like undulatory locomotion in environments where peristaltic locomotion results in poor performance. To the best of our knowledge, the important aspects of locomotion associated with the lateral undulation of an earthworm body are yet to be systematically investigated. In this study, we observed that within confined environments, the worm uses lateral undulation to anchor its body to the walls of their burrows and tip (nose) bending to search the environment. This relatively simple locomotion strategy drastically improved the performance of our soft bodied robophysical model of the earthworm both in a confined (in an acrylic tube) and above-ground heterogeneous environment (rigid pegs), where the peristaltic gait often fails. In summary, lateral undulation facilitates the mobility of earthworm locomotion in diverse environments and can play an important role in the creation of low cost soft robotic devices capable of traversing a variety of environments.

Coordination of lateral body bending and leg movements for sprawled posture quadrupedal locomotion

Many animals generate propulsive forces by coordinating legs, which contact and push against the surroundings, with bending of the body, which can only indirectly influence these forces. Such body–leg coordination is not commonly employed in quadrupedal robotic systems. To elucidate the role of back bending during quadrupedal locomotion, we study a model system: the salamander, a sprawled-posture quadruped that uses lateral bending of the elongate back in conjunction with stepping of the limbs during locomotion. We develop a geometric approach that yields a low-dimensional representation of the body and limb contributions to the locomotor performance quantified by stride displacement. For systems where the damping forces dominate inertial forces, our approach offers insight into appropriate coordination patterns, and improves the computational efficiency of optimization techniques. In particular, we demonstrate effect of the lateral undulation coordinated with leg movement in the forward, rotational, and lateral directions of the robot motion. We validate the theoretical results using numerical simulations, and then successfully test these approaches using robophysical experiments on granular media, a model deformable, frictional substrate. Although our focus lies primarily on robotics, we also demonstrate that our tools can accurately predict optimal body bending of a living salamander Salamandra salamandra.

A fast non-dominated sorting multi-objective symbiotic organism search algorithm for energy efficient locomotion of snake robot

This paper deals with energy efficient locomotion of a wheel-less snake robot. This is very crucial for potential applications of untethered snake robots. The optimum gait parameters for the energy efficient locomotion of the snake robot are obtained with two different multi-objective algorithms based on symbiotic organism search algorithm by considering both minimizing the average power consumption and maximizing the forward velocity of the robot. This paper also investigates the energy efficient locomotion of the snake robot under different environment conditions. The obtained results demonstrate that both proposed methods achieve satisfying stable results regarding power consumption reduction with optimal forward velocity for lateral undulation motion. However, it is seen that fast non-dominated sorting multi-objective symbiotic organism search algorithm provides advantage on obtaining a uniformly distributed solution set with a good diversity only in a single run. This paper is important in terms of presenting useful results for developing efficient motion and environmental adaptability of the snake robot.