The role of intraspinal sensory neurons in the control of quadrupedal locomotion

From swimming to walking and flying, animals have evolved specific locomotor strategies to thrive in different habitats. All types of locomotion depend on integration of motor commands and sensory information to generate precise movements. Cerebrospinal fluid-contacting neurons (CSF-cN) constitute a vertebrate sensory system that monitors CSF composition and flow. In fish, CSF-cN modulate swimming activity in response to changes in pH and bending of the spinal cord, yet their role in higher vertebrates remains unknown. We used mouse genetics to study their function in quadrupedal locomotion and found that CSF-cN are directly integrated into spinal motor circuits by forming connections with motor neurons and premotor interneurons. Elimination of CSF-cN selectively perturbs the accuracy of foot placement required for skilled movements at the balance beam and horizontal ladder. These results identify an important role for mouse CSF-cN in adaptive motor control and indicate that this sensory system evolved a novel function from lower vertebrates to accommodate the biomechanical requirements of terrestrial locomotion.

Cross-bridge mechanics estimated from skeletal muscles’ work-loop responses to impacts in legged locomotion

Legged locomotion has evolved as the most common form of terrestrial locomotion. When the leg makes contact with a solid surface, muscles absorb some of the shock-wave accelerations (impacts) that propagate through the body. We built a custom-made frame to which we fixated a rat (Rattus norvegicus, Wistar) muscle (m. gastrocnemius medialis and lateralis: GAS) for emulating an impact. We found that the fibre material of the muscle dissipates between 3.5 and $$23\,\upmu \hbox {J}$$ 23 μ J ranging from fresh, fully active to passive muscle material, respectively. Accordingly, the corresponding dissipated energy in a half-sarcomere ranges between 10.4 and $$68\,z\hbox {J}$$ 68 z J , respectively. At maximum activity, a single cross-bridge would, thus, dissipate 0.6% of the mechanical work available per ATP split per impact, and up to 16% energy in common, submaximal, activities. We also found the cross-bridge stiffness as low as $$2.2\,\hbox {pN}\,\hbox {nm}^{-1}$$ 2.2 pN nm - 1 , which can be explained by the Coulomb-actuating cross-bridge part dominating the sarcomere stiffness. Results of the study provide a deeper understanding of contractile dynamics during early ground contact in bouncy gait.

Relative skull size as one of the factors limiting skull shape variation in passerines

Despite a considerable interest of researchers to the issue of variation in skull shapes of birds and factors influencing it, some drivers associated with the design features of an entire bird body, which are important for both successful terrestrial locomotion and flight, are overlooked. One of such factors, in our opinion, is relative skull size (skull length in relation to body mass), which can affect the position of the body's center of gravity. We tested effects of relative skull size, allometry (i.e. absolute skull size), and diet on variation in skull shape. The study was conducted on 50 songbird species with a wide range of body mass (8.3g to 570g) and dietary preferences (granivores, insectivores/granivores, insectivores, omnivores). Skull shape was analyzed using 2D geometric morphometrics. We revealed that similar patterns of skull shape occur among passerines with different body sizes and diets. The relative skull size predicted skull shape to a similar extent and with a similar pattern as the absolute size. In our opinion, the effect of the relative skull size on skull shape variation is likely due to biomechanical constraints related to flight.

Modeling Sprawling Locomotion of the Stem Amniote Orobates: An Examination of Hindlimb Muscle Strains and Validation Using Extant Caiman

The stem amniote Orobates pabsti has been reconstructed to be capable of relatively erect, balanced, and mechanically power-saving terrestrial locomotion. This suggested that the evolution of such advanced locomotor capabilities preceded the origin of crown-group amniotes. We here further investigate plausible body postures and locomotion of Orobates by taking soft tissues into account. Freely available animation software BLENDER is used to first reconstruct the lines of action of hindlimb adductors and retractors for Orobates and then estimate the muscle strain of these muscles. We experimentally varied different body heights in modeled hindlimb stride cycles of Orobates to find the posture that maximizes optimal strains over the course of a stride cycle. To validate our method, we used Caiman crocodilus. We replicated the identical workflow used for the analysis of Orobates and compared the locomotor posture predicted for Caiman based on muscle strain analysis with this species’ actual postural data known from a previously published X-ray motion analysis. Since this validation experiment demonstrated a close match between the modeled posture that maximizes optimal adductor and retractor muscle strain and the in vivo posture employed by Caiman, using the same method for Orobates was justified. Generally, the use of muscle strain analysis for the reconstruction of posture in quadrupedal vertebrate fossils thus appears a promising approach. Nevertheless, results for Orobates remained inconclusive as several postures resulted in similar muscle strains and none of the postures could be entirely excluded. These results are not in conflict with the previously inferred moderately erect locomotor posture of Orobates and suggest considerable variability of posture during locomotion.

Body and Tail Coordination in the Bluespot Salamander (Ambystoma laterale) During Limb Regeneration

Animals are incredibly good at adapting to changes in their environment, a trait envied by most roboticists. Many animals use different gaits to seamlessly transition between land and water and move through non-uniform terrains. In addition to adjusting to changes in their environment, animals can adjust their locomotion to deal with missing or regenerating limbs. Salamanders are an amphibious group of animals that can regenerate limbs, tails, and even parts of the spinal cord in some species. After the loss of a limb, the salamander successfully adjusts to constantly changing morphology as it regenerates the missing part. This quality is of particular interest to roboticists looking to design devices that can adapt to missing or malfunctioning components. While walking, an intact salamander uses its limbs, body, and tail to propel itself along the ground. Its body and tail are coordinated in a distinctive wave-like pattern. Understanding how their bending kinematics change as they regrow lost limbs would provide important information to roboticists designing amphibious machines meant to navigate through unpredictable and diverse terrain. We amputated both hindlimbs of blue-spotted salamanders (Ambystoma laterale) and measured their body and tail kinematics as the limbs regenerated. We quantified the change in the body wave over time and compared them to an amphibious fish species, Polypterus senegalus. We found that salamanders in the early stages of regeneration shift their kinematics, mostly around their pectoral girdle, where there is a local increase in undulation frequency. Amputated salamanders also show a reduced range of preferred walking speeds and an increase in the number of bending waves along the body. This work could assist roboticists working on terrestrial locomotion and water to land transitions.

Terrestrial force production by the limbs of a semi-aquatic salamander provides insight into the evolution of terrestrial locomotor mechanics

Amphibious fishes and salamanders are valuable functional analogs for vertebrates that spanned the water-to-land transition. However, investigations of walking mechanics have focused on terrestrial salamanders and, thus, may better reflect the capabilities of stem tetrapods that were already terrestrial. The earliest tetrapods were aquatic, so salamanders that are not primarily terrestrial may yield more appropriate data for modelling the incipient stages of terrestrial locomotion. In the present study, locomotor biomechanics were quantified from semi-aquatic Pleurodeles waltl, a salamander that spends most of its adult life in water, and then compared to a primarily terrestrial salamander (Ambystoma tigrinum) and semi-aquatic fish (Periophthalmus barbarus) to evaluate whether walking mechanics show greater similarity between species with ecological versus phylogenetic similarities. Ground reaction forces (GRFs) from individual limbs or fins indicated that the pectoral appendages of each taxon had distinct patterns of force production, but hind limb forces were comparable between the salamanders. The rate of force development ('yank') was sometimes slower in P. waltl but generally comparable between the three species. Finally, medial inclination of the GRF in P. waltl was intermediate between semi-aquatic fish and terrestrial salamanders, potentially elevating bone stresses among more aquatic taxa as they move on land. These data provide a framework for modelling stem tetrapods using an earlier stage of quadrupedal locomotion that was powered primarily by the hind limbs (i.e., "rear-wheel drive"), and reveal mechanisms for appendages to generate propulsion in three locomotor strategies that are presumed to have occurred across the water-to-land transition in vertebrate evolution.

A mobile Mathieu oscillator model for vibrational locomotion of a bristlebot

Terrestrial locomotion that is produced by creating and exploiting frictional anisotropy is common amongst animals such as snakes, gastropods, limbless lizards. In this paper we present a model of a bristle bot that locomotes by generating frictional anisotropy due to the oscillatory motion of an internal mass and show that this is equivalent to a stick-slip Mathieu oscillator. Such vibrational robots have been available as toys and theoretical curiosities and have seen some applications such as the well known kilobot and in pipe line inspection, but much remains unknown about this type of terrestrial locomotion. In this paper, motivated by a toy model of a bristle bot made from a toothbrush, we derive a theoretical model for its dynamics and show that its dynamics can be classified into four modes of motion : purely stick (no locomotion), slip, stick-slip and hopping. In the stick mode, the dynamics of the system are those of a nonlinear Mathieu oscillator and large amplitude resonance oscillations lead to the slip mode of motion. The mode of motion depends on the amplitude and frequency of the periodic forcing. We compute a phase diagram that captures this behavior, that is reminiscent of the tongues of instability seen in a Mathieu oscillator. The broader result that emerges in this paper is that mobile limbless continuum or soft robots can exploit high frequency parametric oscillations to generate fast and efficient terrestrial motion.

Vibration-Based Locomotion of an Amphibious Robot

In this research, an innovative robot is presented that can move both on land and water thanks to a vibration-based locomotion mechanism. The robot consists of a U-shaped beam made of spring steel, two low-density feet that allow it to stand on the water surface without sinking, and a micro-DC motor with eccentric mass, which excites vibrations. The robot exhibits stable terrestrial and aquatic locomotion based on the synchronization between body vibrations and the centrifugal force due to the eccentric mass. On the one hand, in aquatic locomotion, the robot advances thanks to floating oscillations and the asymmetric shape of the floating feet. On the other hand, the terrestrial locomotion, which has already been demonstrated for a similar robot, exploits the modes of vibration of the elastic beam. In this study, the effect of different excitation frequencies on the locomotion speed in water is examined by means of experimental tests and a numerical model. A good agreement between experimental and numerical results is found. The maximum locomotion speed takes place when the floating modes of vibration are excited.

The scaling of ground reaction forces and duty factor in monitor lizards: implications for locomotion in sprawling tetrapods

Geometric scaling predicts a major challenge for legged, terrestrial locomotion. Locomotor support requirements scale identically with body mass ( α M 1 ), while force-generation capacity should scale α M 2/3 as it depends on muscle cross-sectional area. Mammals compensate with more upright limb postures at larger sizes, but it remains unknown how sprawling tetrapods deal with this challenge. Varanid lizards are an ideal group to address this question because they cover an enormous body size range while maintaining a similar bent-limb posture and body proportions. This study reports the scaling of ground reaction forces and duty factor for varanid lizards ranging from 7 g to 37 kg. Impulses (force×time) ( α M 0.99−1.34 ) and peak forces ( α M 0.73−1.00 ) scaled higher than expected. Duty factor scaled α M 0.04 and was higher for the hindlimb than the forelimb. The proportion of vertical impulse to total impulse increased with body size, and impulses decreased while peak forces increased with speed.