Adaptive Locomotion Research Articles

Anisotropic magnetized tubular microrobots for bioinspired adaptive locomotion

Mimicry of motile microorganisms offers new concepts and strategies to design and manipulate microrobots. As a representative instance, Escherichia coli switches its motility between swimming and tumbling to adapt to surroundings. Inspired by this, we present a tubular microrobot behaving rolling and tumbling motion modes analogous to the motion of Escherichia coli, as a result of the anisotropic magnetized status with radially magnetized status at rolling and longitudinally magnetized one at tumbling. The motion behavior of microrobot as rolling or tumbling is determined by the frequency of rotating magnetic field. The threshold frequency for switching is closely related to the amplitude of magnetic field, viscosity of solution and size of microrobot. The net locomotion near substrate comes from the friction and pressure, and the motion switching is due to the magnetized statuses and discrepancy of drag force. The switch from rolling to tumbling endows the microrobot with adaptive locomotion to cross obstacles. Furthermore, microfluidic manipulation through rotational microrobot is achieved based on the local microvortex, which is generated from the hydrodynamic interaction between microrobot and substrate. This manipulation is applied in cargo delivery and release by altering surrounding microfluid via switchable motion modes. This work opens up new possibilities in improving moving ability and extending functions of microrobots through bioinspired motion behaviors.

Morphological correlates of distal fibular morphology with locomotion in great apes, humans, and Australopithecus afarensis.

ObjectivesRecent studies highlighted the importance of the fibula to further our understanding of locomotor adaptations in fossil hominins. In this study, we present a three‐dimensional geometric morphometric (3D‐GM) investigation of the distal fibula in extant hominids and Australopithecus afarensis with the aim of pointing out morphological correlations to arboreal behavior.MethodsThree‐dimensional surface meshes of the distal fibula were obtained using computer tomography for 40 extant hominid specimens and laser scanner for five A. afarensis specimens. Distal fibula morphology was quantified positioning 11 fixed landmarks, 40 curve semilandmarks, and 20 surface landmarks on each specimen. A generalized Procrustes analysis (GPA) was carried out on all landmark coordinates followed by Procrustes ANOVA. Principal component analysis (PCA) was performed on the GPA‐aligned shape coordinates. Kruskal‐Wallis tests and Mann–Whitney test were performed on scores along PCs.ResultsGreat apes are characterized by a shorter subcutaneous triangular surface (STS), more downward facing fibulotalar articular facets, more anteriorly facing lateral malleolus and wider/deeper malleolar fossa than humans. Within great apes, orangutans are characterized by more medially facing fibulotalar articular facets. Australopithecus afarensis shows a unique distal fibular morphology with several traits that are generally associated more to arboreality and less to bipedalism such as a short STS, a more anteriorly facing, laterally pointing malleolus and deeper and larger malleolar fossa.ConclusionsThe distal fibula morphology is indicative of locomotor patterns within extant hominids. The 3D‐GM method presented here can be successfully used to further our understanding of arboreal adaptations in fossil hominins.

Genomic adaptations for arboreal locomotion in Asian flying treefrogs

SignificanceTo adapt to arboreal lifestyles, treefrogs have evolved a suite of complex traits that support vertical movement and gliding, thus presenting a unique case for studying the genetic basis for traits causally linked to vertical niche expansion. Here, based on two de novo-assembled Asian treefrog genomes, we determined that genes involved in limb development and keratin cytoskeleton likely played a role in the evolution of their climbing systems. Behavioral and morphological evaluation and time-ordered gene coexpression network analysis revealed the developmental patterns and regulatory pathways of the webbed feet used for gliding in Rhacophorus kio.

Adjustable whole-body dynamics for adaptive locomotion: the influence of upper body movements and its interactions with the lower body parts on the stable locomotion of a simple bipedal robot

AbstractThis paper investigates the influence of adding an upper body to a bipedal robot on its stable walking behavior. The robot’s parts are mutually interconnected through an actuator network system. Therefore, the movement pattern of the upper body depends on the type of interactions created with other limbs. Throughout the experiments, various interactions among the different body parts were tested. The results showed that a robot with a motionless upper body exhibited unstable walking behavior. However, once the same upper body was involved and interacted properly, with other body parts, its movement significantly helped to stabilize the behavior of the robot.

Use of the Azure Kinect to measure foot clearance during obstacle crossing: A validation study.

Obstacle crossing is typical adaptive locomotion known to be related to the risk of falls. Previous conventional studies have used elaborate and costly optical motion capture systems, which not only represent a considerable expense but also require participants to visit a laboratory. To overcome these shortcomings, we aimed to develop a practical and inexpensive solution for measuring obstacle-crossing behavior by using the Microsoft Azure Kinect, one of the most promising markerless motion capture systems. We validated the Azure Kinect as a tool to measure foot clearance and compared its performance to that of an optical motion capture system (Qualisys). We also determined the effect of the Kinect sensor placement on measurement performance. Sixteen healthy young men crossed obstacles of different heights (50, 150, and 250 mm). Kinect sensors were placed in front of and beside the obstacle as well as diagonally between those positions. As indices of measurement quality, we counted the number of measurement failures and calculated the systematic and random errors between the foot clearance measured by the Kinect and Qualisys. We also calculated the Pearson correlation coefficients between the Kinect and Qualisys measurements. The number of measurement failures and the systematic and random error were minimized when the Kinect was placed diagonally in front of the obstacle on the same side as the trail limb. The high correlation coefficient (r > 0.890) observed between the Kinect and Qualisys measurements suggest that the Azure Kinect has excellent potential for measuring foot clearance during obstacle-crossing tasks.

Context-Specificity of Locomotor Learning Is Developed during Childhood.

Humans can perform complex movements with speed and agility in the face of constantly changing task demands. To accomplish this, motor plans are adapted to account for errors in our movements because of changes in our body (e.g., growth or injury) or in the environment (e.g., walking on sand vs ice). It has been suggested that adaptation that occurs in response to changes in the state of our body will generalize across different movement contexts and environments, whereas adaptation that occurs with alterations in the external environment will be context-specific. Here, we asked whether the ability to form generalizable versus context-specific motor memories develops during childhood. We performed a cross-sectional study of context-specific locomotor adaptation in 35 children (3–18 years old) and 7 adults (19–31 years old). Subjects first adapted their gait and learned a new walking pattern on a split-belt treadmill, which has two belts that move each leg at a different speed. Then, subjects walked overground to assess the generalization of the adapted walking pattern across different environments. Our results show that the generalization of treadmill after-effects to overground walking decreases as subjects’ age increases, indicating that age and experience are critical factors regulating the specificity of motor learning. Our results suggest that although basic locomotor patterns are established by two years of age, brain networks required for context-specific locomotor learning are still being developed throughout youth.

Articulating Resilience: Adaptive Locomotion of Wheeled Tensegrity Robot

Resilience plays an important role in improving robustness for robots in harsh environments such as planetary exploration and unstructured terrains. As a naturally compliant structure, tensegrity presents advantageous flexibility for enhancing resilience in robotic applications according to existing research. However, tensegrity robots to date are normally based on monolithic morphologies and are slow in locomotion. In this paper, we demonstrate how we adopt such flexibility to improve the robustness of wheeled robots by articulating modules with tensegrity mechanisms. The test results reveal the robot is resistant and resilient to external hazards in a fully passive manner owing to the continuous elasticity in the structure network. It possesses a good number of DoFs and can adapt to various terrains easily either with actuation or not. The robot is also capable of crawling locomotion aside from wheeled locomotion to traverse uneven surfaces and provide self-recovery from rollovers. It demonstrates good robustness and mobility at the same time compared with existing tensegrity robots and shows the competitiveness with conventional rigid robots in harsh scenarios. The proposed robot presents the capability of tensegrity robots with resilience, robustness, agility, and mobility without compromise. In a broader perspective, it widens the potential of tensegrity robots in practical applications.

Real-time adaptive leg-stiffness for roll compensation via magnetorheological control in a legged robot

Over the recent few decades, the evolving research-field of legged robotics has seen various mechanical and control-based developments. Inspired by biological species, a significant adaptation in modern mechanical leg designs has been the implementation of adjustable stiffness, shifting from what were previously simple linkages to more-complex variable stiffness actuators. Physiological studies previously demonstrated leg-stiffness modulation was not only a common trait in multiple biological locomotors, but also played a key role in disturbance recovery for humans. Guided by this, recent robotics research has shown that this can also be applied to legged robots to achieve similar locomotion adaptations, albeit often limited by the tuning time of leg stiffness in such circumstances. This study proposes real-time adaptive stiffness robot legs which are governed by fast-response magnetorheological fluid dampers, enabling stiffness adjustment upon a single step. Through experimental characterisation and model validation, these legs are shown to achieve a maximum stiffness shift of 114%. Enabled by real-time control during locomotion, improved performance and roll-angle stability is experimentally demonstrated for a bipedal robot test platform. Such improvement to locomotion is found through typical legged locomotion scenarios, with the platform encountering: obstacles, valleys, and coronal gradients in a comprehensive series of experiments.

Electromagnetic Feet With Soft Toes for Adaptive, Versatile, and Stable Locomotion of an Inchworm-Inspired Pipe Crawling Robot.

Feet play an important role in the adaptive, versatile, and stable locomotion of legged creatures. Accordingly, several robotic research studies have used biological feet as the inspiration for the design of robot feet in traversing complex terrains. However, so far, no robot feet can allow legged robots to adaptively, versatilely, and robustly crawl on various curved metal pipes, including flat surfaces for pipe inspection. To address this issue, we propose here a novel hybrid rigid-soft robot-foot design inspired by the leg morphology of an inchworm. The foot consists of a rigid section with an electromagnet and a soft toe covering for enhanced adhesion to a metal pipe. Finite element analysis , performed under different loading conditions, reveals that due to its compliance, the soft toe can undergo recoverable deformation with adaptability to various curved metal pipes and plain metal surfaces. We have successfully implemented electromagnetic feet with soft toes (EROFT) on an inchworm-inspired pipe crawling robot for adaptive, versatile, and stable locomotion. Foot-to-surface adaptability is provided by the inherent elasticity of the soft toe, making the robot a versatile and stable metal pipe crawler. Experiments show that the robot crawling success rate reaches 100% on large diameter metal pipes. The proposed hybrid rigid-soft feet (i.e., electromagnetic feet with soft toes) can solve the problem of continuous surface adaptation for the robot in a stable and efficient manner, irrespective of the surface curvature, without the need to manually change the robot feet for specific surfaces. To this end, the foot development enables the robot to meet a set of deployment requirements on large oil and gas pipelines for potential use in inspecting various faults and leakages.

Effects of sensory manipulations on locomotor adaptation to split-belt treadmill walking in healthy younger and older adults

Locomotor adaptation relies on processes of both the peripheral and central nervous systems that may be compromised with advanced age (e.g., proprioception, sensorimotor integration). Age-related changes to these processes may result in reduced rates of locomotor adaptation under normal conditions and should cause older adults to be disproportionately more affected by sensory manipulations during adaptation compared to younger adults. 17 younger and 10 older adults completed five separate 5-minute split-belt walking trials: three under normal sensory conditions, one with 30% bodyweight support (meant to reduce proprioceptive input), and one with goggles that constrained the visual field (meant to reduce visual input). We fit step length symmetry data from each participant in each trial with a single exponential function and used the time constant to quantify locomotor adaption rate. Group by trial ANOVAs were used to test the effects of age, condition, and their interaction on adaptation rates. Contrary to our hypothesis, we found no evidence that sensory manipulations disproportionately affected older compared to younger adults, at least in our relatively small sample. In fact, in both groups, adaptation rates remained unaffected across all trials, including both normal and sensory manipulated trials. Our results provide evidence that both younger and older adults were able to adequately reweight sources of sensory information based on environmental constraints, indicative of well-functioning neural processes of motor adaptation.

Voltage imaging identifies spinal circuits that modulate locomotor adaptation in zebrafish

Motor systems must continuously adapt their output to maintain a desired trajectory. While the spinal circuits underlying rhythmic locomotion are well described, little is known about how the network modulates its output strength. A major challenge has been the difficulty of recording from spinal neurons during behavior. Here, we use voltage imaging to map the membrane potential of large populations of glutamatergic neurons throughout the spinal cord of the larval zebrafish during fictive swimming in a virtual environment. We characterized a previously undescribed subpopulation of tonic-spiking ventral V3 neurons whose spike rate correlated with swimming strength and bout length. Optogenetic activation of V3 neurons led to stronger swimming and longer bouts but did not affect tail beat frequency. Genetic ablation of V3 neurons led to reduced locomotor adaptation. The power of voltage imaging allowed us to identify V3 neurons as a critical driver of locomotor adaptation in zebrafish.

Crab-inspired compliant leg design method for adaptive locomotion of a multi-legged robot

Chinese mitten crab has unique limb structures composed of a hard exoskeleton and flexible muscles. They enable the crab to locomote adaptively and safely on various terrains. In this work, we investigated the limb structures, motion principle, and gaits of the crab using a high-speed camera and a press machine. Then, a novel compliant robot leg design method is proposed, inspired by the crab limb. The leg comprises six hard scleromeres and a flexible thin-wall spring steel sheet (FSSS) mimicking the exoskeleton and muscle. The scleromeres connected one by one with rotational joints are designed with slots. The front end of the FSSS is fixed on the scleromere close to the ground. The rear end crosses the slots and is mounted at the shaft of a linear actuator installed at the rear scleromere. The leg bends and stretches when the actuator pushes and pulls the FSSS, respectively. The kinematic modeling, rigid-flexible coupling dynamic simulations, and leg prototype tests are conducted, which verify the leg design approach. Thirdly, we put forward a multi-legged robot with eight compliant legs and design its gait using the gaits of the crab. Finally, the robot’s performance is evaluated, including the capabilities of walking on different terrains at adjustable speeds and body heights, traversing low channels, walking on slopes, and carrying loads. The results prove that the single-motor-actuated compliant legs and their dynamic coupling with the rigid robot body frame can enable them to have the ground clearance ability and realize the adaptive walking of the robot. The leg design methodology can be used to design multi-legged robots with the merits of compact, light, low mechanical complexity, high safety, and easy to control, for many applications, such as environmental monitoring, search and rescue.

Coordinating Clearance and Postural Reorientation When Avoiding Physical and Virtual Pedestrians.

Community ambulation requires efficient locomotor adaptations to avoid collisions with other pedestrians. Virtual reality (VR) offers the possibility to experimentally manipulate the environment, allowing researchers to safely assess locomotor responses during pedestrian interactions. In this paper, locomotor adjustments in response to interferers approaching from different directions were examined in both a physical and a virtual environment. Further analysis compared the extent to which locomotor adjustments were influenced by different circumvention strategies. To achieve these goals, twelve healthy young participants were assessed while walking towards a target and avoiding pedestrians approaching from the left, middle, or right, in both VR and the real world. Results showed that in VR, participants walked with slower velocities, maintained larger minimum distances, and reached larger trajectory deviations. Additionally, trajectory deviations and postural reorientations were executed earlier in VR. There were additional differences according to whether participants had to give way to a head-on approaching interferer or choose to pass in front or behind a diagonally approaching interferer. Compared to the other circumvention strategies, passing in front was associated with faster walking speeds as well as smaller and later trajectory deviations and postural reorientations. Lastly, while a cephalocaudal sequence of segment reorientation was observed in both environments, no difference between the onset of head and thorax reorientation was observed in VR. In conclusion, obstacle clearance and postural reorientation are modulated by the environment and circumvention strategies. These modulations should be considered when designing experiments or clinical interventions.

Split-Belt Training but Not Cerebellar Anodal tDCS Improves Stability Control and Reduces Risk of Fall in Patients with Multiple Sclerosis.

The objective of this study was to examine the therapeutic potential of multiple sessions of training on a split-belt treadmill (SBT) combined with cerebellar anodal transcranial direct current stimulation (tDCS) on gait and balance in People with Multiple Sclerosis (PwMS). Twenty-two PwMS received six sessions of anodal (PwMSreal, n = 12) or sham (PwMSsham, n = 10) tDCS to the cerebellum prior to performing the locomotor adaptation task on the SBT. To evaluate the effect of the intervention, functional gait assessment (FGA) scores and distance walked in 2 min (2MWT) were measured at the baseline (T0), day 6 (T5), and at the 4-week follow up (T6). Locomotor performance and changes of motor outcomes were similar in PwMSreal and PwMSsham independently from tDCS mode applied to the cerebellum (anodal vs. sham, on FGA, p = 0.23; and 2MWT, p = 0.49). When the data were pooled across the groups to investigate the effects of multiple sessions of SBT training alone, significant improvement of gait and balance was found on T5 and T6, respectively, relative to baseline (FGA, p < 0.001 for both time points). The FGA change at T6 was significantly higher than at T5 (p = 0.01) underlining a long-lasting improvement. An improvement of the distance walked during the 2MWT was also observed on T5 and T6 relative to T0 (p = 0.002). Multiple sessions of SBT training resulted in a lasting improvement of gait stability and endurance, thus potentially reducing the risk of fall as measured by FGA and 2MWT. Application of cerebellar tDCS during SBT walking had no additional effect on locomotor outcomes.

Gait Adaptation to a Phase-Specific Nociceptive Electrical Stimulation Applied at the Ankle: A Model to Study Musculoskeletal-Like Pain.

Introduction: Lower limb pain, whether induced experimentally or as a result of a musculoskeletal injury, can impair motor control, leading to gait adaptations such as increased muscle stiffness or modified load distribution around joints. These adaptations may initially reduce pain but can also lead to longer-term maladaptive plasticity and to the development of chronic pain. In humans, many current experimental musculoskeletal-like pain models are invasive, and most don’t accurately reproduce the movement-related characteristics of musculoskeletal pain. The main objective of this study was to measure pain adaptation strategies during gait of a musculoskeletal-like experimental pain protocol induced by phase-specific, non-invasive electrical stimulation.Methods: Sixteen healthy participants walked on a treadmill at 4 km/h for three consecutive periods (BASELINE, PAIN, and POST-PAIN). Painful electrical stimulations were delivered at heel strike for the duration of heel contact (HC) using electrodes placed around the right lateral malleolus to mimic ankle sprains. Gait adaptations were quantified bilaterally using instrumented pressure-sensitive insoles. One-way ANOVAs and group time course analyses were performed to characterize the impact of electrical stimulation on heel and forefoot contact pressure and contact duration.Results: During the first few painful strides, peak HC pressure decreased on the painful side (8.6 ± 1.0%, p < 0.0001) and increased on the non-stimulated side (11.9 ± 0.9%, p < 0.0001) while HC duration was significantly reduced bilaterally (painful: 12.1 ± 0.9%, p < 0.0001; non-stimulated: 4.8 ± 0.8%, p < 0.0001). No clinically meaningful modifications were observed for the forefoot. One minute after the onset of painful stimulation, perceived pain levels stabilized and peak HC pressure remained significantly decreased on the painful side, while the other gait adaptations returned to pre-stimulation values.Discussion: These results demonstrate that a non-invasive, phase-specific pain can produce a stable painful gait pattern. Therefore, this protocol will be useful to study musculoskeletal pain locomotor adaptation strategies under controlled conditions.

Younger and Late Middle-Aged Adults Exhibit Different Patterns of Cognitive-Motor Interference During Locomotor Adaptation, With No Disruption of Savings.

It has been proposed that motor adaptation and subsequent savings (or faster relearning) of an adapted movement pattern are mediated by cognitive processes. Here, we evaluated the pattern of cognitive-motor interference that emerges when young and late middle-aged adults perform an executive working memory task during locomotor adaptation. We also asked if this interferes with savings of a newly learned walking pattern, as has been suggested by a study of reaching adaptation. We studied split-belt treadmill adaptation and savings in young (21 ± 2 y/o) and late middle-aged (56 ± 6 y/o) adults with or without a secondary 2-back task during adaptation. We found that young adults showed similar performance on the 2-back task during baseline and adaptation, suggesting no effect of the dual-task on cognitive performance; however, dual-tasking interfered with adaptation over the first few steps. Conversely, dual-tasking caused a decrement in cognitive performance in late middle-aged adults with no effect on adaptation. To determine if this effect was specific to adaptation, we also evaluated dual-task interference in late middle-aged adults that dual-tasked while walking in a complex environment that did not induce motor adaptation. This group exhibited less cognitive-motor interference than late middle-aged adults who dual-tasked during adaptation. Savings was unaffected by dual-tasking in both young and late middle-aged adults, which may indicate different underlying mechanisms for savings of reaching and walking. Collectively, our findings reveal an age-dependent effect of cognitive-motor interference during dual-task locomotor adaptation and no effect of dual-tasking on savings, regardless of age. Young adults maintain cognitive performance and show a mild decrement in locomotor adaptation, while late middle-aged adults adapt locomotion at the expense of cognitive performance.

Locomotion Control With Frequency and Motor Pattern Adaptations.

Existing adaptive locomotion control mechanisms for legged robots are usually aimed at one specific type of adaptation and rarely combined with others. Adaptive mechanisms thus stay at a conceptual level without their coupling effect with other mechanisms being investigated. However, we hypothesize that the combination of adaptation mechanisms can be exploited for enhanced and more efficient locomotion control as in biological systems. Therefore, in this work, we present a central pattern generator (CPG) based locomotion controller integrating both a frequency and motor pattern adaptation mechanisms. We use the state-of-the-art Dual Integral Learner for frequency adaptation, which can automatically and quickly adapt the CPG frequency, enabling the entire motor pattern or output signal of the CPG to be followed at a proper high frequency with low tracking error. Consequently, the legged robot can move with high energy efficiency and perform the generated locomotion with high precision. The versatile state-of-the-art CPG-RBF network is used as a motor pattern adaptation mechanism. Using this network, the motor patterns or joint trajectories can be adapted to fit the robot's morphology and perform sensorimotor integration enabling online motor pattern adaptation based on sensory feedback. The results show that the two adaptation mechanisms can be combined for adaptive locomotion control of a hexapod robot in a complex environment. Using the CPG-RBF network for motor pattern adaptation, the hexapod learned basic straight forward walking, steering, and step climbing. In general, the frequency and motor pattern mechanisms complement each other well and their combination can be seen as an essential step toward further studies on adaptive locomotion control.

How big can a walking fish be? A theoretical inference based on observations on four land-dwelling fish genera of South Vietnam.

Comparative study of terrestrial locomotion of 4 fish genera including Anabas, Channa, Clarias, and Monopterus, was performed in experimental setting with the substrate surface of wet clay. No special adaptations for terrestrial locomotion were found. Every fish uses for propulsion on land what it already has. Eel-shaped Monopterus crawls by body undulations in a serpentine or sidewinding technique, the latter of which was not previously observed beyond snakes. The other 3 fish genera walk by body oscillations using stiff appendages as propulsors. When they are located anteriorly, as the serrate operculum in Anabas and the preaxial spine of the pectoral fin in Clarias, the propulsion is termed prolocomotor, when posteriorly, as the spiny anal fin in Channa-metalocomotor. Channa is the heaviest fish walking out of water in our days, quite comparable in size with first Devonian tetrapods Acanthostega and Tulerpeton. A theoretical calculation is suggested for the upper size limit of a fish capable of terrestrial walking without special locomotor adaptations. It should be roughly 20 cm in the vertical dimension of the trunk, which is just a little above the known size of Devonian tetrapodomorph fishes Panderichthys and Elpistostege. The metalocomotor walking technique of Channa is suggested as the closest extant model for terrestrial locomotion at the fish-tetrapod transition. The major difference is that the metalocomotor propulsor in Channa is represented by the anal fin, while in tetrapodomorphs by the pelvic fins. The sprawled pelvic fins were advantageous in respect of reduced requirement for side-to-side tail swinging.

Circuits for State-Dependent Modulation of Locomotion.

Brain-wide neural circuits enable bi- and quadrupeds to express adaptive locomotor behaviors in a context- and state-dependent manner, e.g., in response to threats or rewards. These behaviors include dynamic transitions between initiation, maintenance and termination of locomotion. Advances within the last decade have revealed an intricate coordination of these individual locomotion phases by complex interaction of multiple brain circuits. This review provides an overview of the neural basis of state-dependent modulation of locomotion initiation, maintenance and termination, with a focus on insights from circuit-centered studies in rodents. The reviewed evidence indicates that a brain-wide network involving excitatory circuit elements connecting cortex, midbrain and medullary areas appears to be the common substrate for the initiation of locomotion across different higher-order states. Specific network elements within motor cortex and the mesencephalic locomotor region drive the initial postural adjustment and the initiation of locomotion. Microcircuits of the basal ganglia, by implementing action-selection computations, trigger goal-directed locomotion. The initiation of locomotion is regulated by neuromodulatory circuits residing in the basal forebrain, the hypothalamus, and medullary regions such as locus coeruleus. The maintenance of locomotion requires the interaction of an even larger neuronal network involving motor, sensory and associative cortical elements, as well as defined circuits within the superior colliculus, the cerebellum, the periaqueductal gray, the mesencephalic locomotor region and the medullary reticular formation. Finally, locomotor arrest as an important component of defensive emotional states, such as acute anxiety, is mediated via a network of survival circuits involving hypothalamus, amygdala, periaqueductal gray and medullary premotor centers. By moving beyond the organizational principle of functional brain regions, this review promotes a circuit-centered perspective of locomotor regulation by higher-order states, and emphasizes the importance of individual network elements such as cell types and projection pathways. The realization that dysfunction within smaller, identifiable circuit elements can affect the larger network function supports more mechanistic and targeted therapeutic intervention in the treatment of motor network disorders.

Friction modulation in limbless, three-dimensional gaits and heterogeneous terrains

Motivated by a possible convergence of terrestrial limbless locomotion strategies ultimately determined by interfacial effects, we show how both 3D gait alterations and locomotory adaptations to heterogeneous terrains can be understood through the lens of local friction modulation. Via an effective-friction modeling approach, compounded by 3D simulations, the emergence and disappearance of a range of locomotory behaviors observed in nature is systematically explained in relation to inhabited environments. Our approach also simplifies the treatment of terrain heterogeneity, whereby even solid obstacles may be seen as high friction regions, which we confirm against experiments of snakes ‘diffracting’ while traversing rows of posts, similar to optical waves. We further this optic analogy by illustrating snake refraction, reflection and lens focusing. We use these insights to engineer surface friction patterns and demonstrate passive snake navigation in complex topographies. Overall, our study outlines a unified view that connects active and passive 3D mechanics with heterogeneous interfacial effects to explain a broad set of biological observations, and potentially inspire engineering design.