Locomotion in Extinct Giant Kangaroos: Were Sthenurines Hop-Less Monsters?

Sthenurine kangaroos (Marsupialia, Diprotodontia, Macropodoidea) were an extinct subfamily within the family Macropodidae (kangaroos and rat-kangaroos). These “short-faced browsers” first appeared in the middle Miocene, and radiated in the Plio-Pleistocene into a diversity of mostly large-bodied forms, more robust than extant forms in their build. The largest (Procoptodon goliah) had an estimated body mass of 240 kg, almost three times the size of the largest living kangaroos, and there is speculation whether a kangaroo of this size would be biomechanically capable of hopping locomotion. Previously described aspects of sthenurine anatomy (specialized forelimbs, rigid lumbar spine) would limit their ability to perform the characteristic kangaroo pentapedal walking (using the tail as a fifth limb), an essential gait at slower speeds as slow hopping is energetically unfeasible. Analysis of limb bone measurements of sthenurines in comparison with extant macropodoids shows a number of anatomical differences, especially in the large species. The scaling of long bone robusticity indicates that sthenurines are following the “normal” allometric trend for macropodoids, while the large extant kangaroos are relatively gracile. Other morphological differences are indicative of adaptations for a novel type of locomotor behavior in sthenurines: they lacked many specialized features for rapid hopping, and they also had anatomy indicative of supporting their body with an upright trunk (e.g., dorsally tipped ischiae), and of supporting their weight on one leg at a time (e.g., larger hips and knees, stabilized ankle joint). We propose that sthenurines adopted a bipedal striding gait (a gait occasionally observed in extant tree-kangaroos): in the smaller and earlier forms, this gait may have been employed as an alternative to pentapedal locomotion at slower speeds, while in the larger Pleistocene forms this gait may have enabled them to evolve to body sizes where hopping was no longer a feasible form of more rapid locomotion.     

Morphological variation of the thoracolumbar vertebrae in Macropodidae and its functional relevance

The present study was designed to investigate how the form of the marsupial thoracolumbar vertebrae varies to cope with the particular demands of diverse loading and locomotor behaviors. The vertebral columns of 10 species of Macropodidae, with various body masses and modes of locomotion, together with two other arboreal marsupials, koala and cuscus, were selected. Seventy-four three-dimensional landmark coordinates were acquired on each of the 10 last presacral vertebrae of the 70 vertebral columns. The interspecific variations of the third lumbar vertebra (L3, which approximates the mean) and the transitional patterns of the thoracolumbar segments were examined using the combined approach of generalized Procrustes analysis (GPA) and principal components analysis (PCA). The results of analyses of an individual vertebra (L3) and of the transitional patterns indicate significant interspecific differences. In the L3 study the first PC shows allometric shape variation, while the second PC seems to relate to adaptation for terrestrial versus arboreal locomotion. When the L3 vertebrae of the common spotted cuscus and koala are included for comparison, the vertebra of the tree kangaroo occupies an intermediate position between the hopping kangaroo and these arboreal marsupials. The L3 vertebrae in the arboreal marsupials possess a distinct dorsoventrally expanded vertebral body, and perpendicularly orientated spinous and transverse processes. The results of the present study suggest that vertebral shape in the kangaroo and wallaroos provides a structural adaptation to hopping through a relatively enlarged loading area and powerful lever system. In contrast, the small-sized bettongs (or rat kangaroos) have a relatively flexible column and elongated levers for the action of back muscles that extend and laterally flex the spine. The complex pattern of vertebral shape transition in the last 10 presacral vertebrae was examined using PCAs that compare between species information about vertebral shape variation along the thoracolumbar column. The results reinforce and emphasize important aspects of the patterns of variation seen in the detailed analysis of the third lumbar vertebra. The results also imply that size, spinal loading pattern, and locomotor behavior exert an influence on shaping the vertebra. Further, the morphological adaptations are consistent within these marsupials and this opens up the possibility that this kind of analysis may be useful in making functional inferences from fossil material.     

Toward Fast and Efficient Mobility in Aquatic Environment: A Robot with Compliant Swimming Appendages Inspired by a Water Beetle

Water beetles are proficient drag-powered swimmers,with oar-like legs.Inspired by this mechamsm,here we propose a miniature robot,with mobility provided by a pair of legs with swimming appendages.The robot has optimized linkage structure to maximize the stroke angle,which is actuated by a single DC motor with a series of gears and a spring.A simplified swimming appendage model is proposed to calculate the deflection due to the applied drag force,and is compared with simulated data using COMSOL Multiphysics.Also,the swimming appendages are optimized by considering their locations on the legs using two fitness functions,and six different configurations are selected.We investigate the performance of the robot with various types of appendage using a high-speed camera,and motion capture cameras.The robot with the proposed configuration exhibits fast and efficient movement compared with other robots.In addition,the locomotion of the robot is analyzed by considering its dynamics,and compared with that of a water boatman (Corixidae).     

Bird terrestrial locomotion as revealed by 3D kinematics

Most birds use at least two modes of locomotion: flying and walking (terrestrial locomotion). Whereas the wings and tail are used for flying, the legs are mainly used for walking. The role of other body segments remains, however, poorly understood. In this study, we examine the kinematics of the head, the trunk, and the legs during terrestrial locomotion in the quail (Coturnix coturnix). Despite the trunk representing about 70% of the total body mass, its function in locomotion has received little scientific interest to date. This prompted us to focus on its role in terrestrial locomotion. We used high-speed video fluoroscopic recordings of quails walking at voluntary speeds on a trackway. Dorso-ventral and lateral views of the motion of the skeletal elements were recorded successively and reconstructed in three dimensions using a novel method based on the temporal synchronisation of both views. An analysis of the trajectories of the body parts and their coordination showed that the trunk plays an important role during walking. Moreover, two sub-systems participate in the gait kinematics: (i) the integrated 3D motion of the trunk and thighs allows for the adjustment of the path of the centre of mass; (ii) the motion of distal limbs transforms the alternating forward motion of the feet into a continuous forward motion at the knee and thus assures propulsion. Finally, head bobbing appears qualitatively synchronised to the movements of the trunk. An important role for the thigh muscles in generating the 3D motion of the trunk is suggested by an analysis of the pelvic anatomy.     

Stance leg control: variation of leg parameters supports stable hopping

The spring-loaded inverted pendulum describes the planar center-of-mass dynamics of legged locomotion. This model features linear springs with constant parameters as legs. In biological systems, however, spring-like properties of limbs can change over time. Therefore, in this study, it is asked how variation of spring parameters during ground contact would affect the dynamics of the spring-mass model. Neglecting damping initially, it is found that decreasing stiffness and increasing rest length of the leg during a stance phase are required for orbitally stable hopping. With damping, stable hopping is found for a larger region of rest-length rates and stiffness rates. Here, also increasing stiffness and decreasing rest length can result in stable hopping. Within the predicted range of leg parameter variations for stable hopping, there is no need for precise parameter tuning. Since hopping gaits form a subset of the running gaits (with vanishing horizontal velocity), these results may help to improve leg design in robots and prostheses.     

Development of a Biomimetic Quadruped Robot

This paper presents the design and prototype of a small quadruped robot whose walking motion is realized by two piezocomposite actuators. In the design, biomimetic ideas are employed to obtain the agility of motions and sustainability of a heavy load. The design of the robot legs is inspired by the leg configuration of insects, two joints （hip and knee） of the leg enable two basic motions, lifting and stepping. The robot frame is designed to have a slope relative to the horizontal plane, which makes the robot move forward. In addition, the bounding locomotion of quadruped animals is implemented in the robot. Experiments show that the robot can carry an additional load of about 100 g and run with a fairly high velocity. The quadruped prototype can be an important step towards the goal of building an autonomous mobile robot actuated by piezocomposite actuators.     

Velocity Control of a Bounding Quadruped via Energy Control and Vestibular Reflexes

In this paper a bio-inspired approach of velocity control for a quadruped robot running with a bounding gait on compliant legs is set up. The dynamic properties ofa sagittal plane model of the robot are investigated. By analyzing the stable fixed points based on Poincare map, we find that the energy change of the system is the main source for forward velocity adjustment. Based on the analysis of the dynamics model of the robot, a new simple linear running controller is proposed using the energy control idea, which requires minimal task level feedback and only controls both the leg torque and ending impact angle. On the other hand, the functions of mammalian vestibular reflexes are discussed, and a reflex map between forward velocity and the pitch movement is built through statistical regression analysis. Finally, a velocity controller based on energy control and vestibular reflexes is built, which has the same structure as the mammalian nervous mechanism for body posture control. The new con- troller allows the robot to run autonomously without any other auxiliary equipment and exhibits good speed adjustment capa- bility. A series simulations and experiments were set to show the good movement agility, and the feasibility and validity of the robot system.     

Collision-based mechanics of bipedal hopping

The muscle work required to sustain steady-speed locomotion depends largely upon the mechanical energy needed to redirect the centre of mass and the degree to which this energy can be stored and returned elastically. Previous studies have found that large bipedal hoppers can elastically store and return a large fraction of the energy required to hop, whereas small bipedal hoppers can only elastically store and return a relatively small fraction. Here, we consider the extent to which large and small bipedal hoppers (tammar wallabies, approx. 7 kg, and desert kangaroo rats, approx. 0.1 kg) reduce the mechanical energy needed to redirect the centre of mass by reducing collisions. We hypothesize that kangaroo rats will reduce collisions to a greater extent than wallabies since kangaroo rats cannot elastically store and return as high a fraction of the mechanical energy of hopping as wallabies. We find that kangaroo rats use a significantly smaller collision angle than wallabies by employing ground reaction force vectors that are more vertical and center of mass velocity vectors that are more horizontal and thereby reduce their mechanical cost of transport. A collision-based approach paired with tendon morphometry may reveal this effect more generally among bipedal runners and quadrupedal trotters.     

Sensory feedback in CNN-based central pattern generators

Central Pattern Generators (CPGs) are a suitable paradigm to solve the problem of locomotion control in walking robots. CPGs are able to generate feed-forward signals to achieve a proper coordination among the robot legs. In literature they are often modelled as networks of coupled nonlinear systems. However the topic of feedback in these systems is rarely addressed. On the other hand feedback is essential for locomotion. In this paper the CPG for a hexapod robot is implemented through Cellular Neural Networks (CNNs). Feedback is included in the CPG controller by exploiting the dynamic properties of the CPG motor-neurons, such as synchronization issue and local bifurcations. These universal paradigms provide the essential issues to include sensory feedback in CPG architectures based on coupled nonlinear systems. Experiments on a dynamic model of a hexapod robot are presented to validate the approach introduced.     

How does a hopping kangaroo breathe?

We developed a model to demonstrate how a hopping kangaroo breathes. Interestingly, a kangaroo uses less energy to breathe while hopping than while standing still. This occurs, in part, because rather than using muscle power to move air into and out of the lungs, air is pulled into (inspiration) and pushed out of (expiration) the lungs as the abdominal organs "flop" within the kangaroo's body. Specifically, as the kangaroo hops upward, the abdominal organs lag behind, and the insertion of the diaphragm is pulled toward its origin, flattening the dome and increasing the vertical dimension of the thoracic cavity (the thoracic cavity and lungs enlarge). Increasing the volume of the thoracic cavity reduces alveolar pressure below atmospheric pressure (barometric pressure), and air moves into the alveoli by bulk flow. In contrast, the impact of the organs against the diaphragm at each landing causes expiration. Specifically, upon landing, the abdominal organs flop into the diaphragm, causing it to return to its dome shape and decreasing the vertical dimension of the thoracic cavity. This compresses the alveolar gas volume and elevates alveolar pressure above barometric pressure, so air is expelled. To demonstrate this phenomenon, the plunger of a syringe model of the respiratory system was inserted through a compression spring. Holding the syringe and pressing the plunger firmly against a hard surface expels air from the lungs (the balloon within the syringe deflates) and compresses the spring. This models the kangaroo landing after a hop forward. Subsequently, the compression spring provides the energy for the "kangaroo" to "hop" forward upon the release of the syringe, and air enters the lungs (the balloon within the syringe inflates). The model accurately reflects how a hopping kangaroo breathes. A model was chosen to demonstrate this phenomenon because models engage and inspire students as well as significantly enhance student understanding.     

Biomimetic Design and Optimal Swing of a Hexapod Robot Leg

Biological inspiration has spawned a wealth of solutions to both mechanical design and control schemes in the efforts to develop agile legged machines. This paper presents a compliant leg mechanism for a small six-legged robot, HITCR-ll, based on abstracted anatomy from insect legs. Kinematic structure, relative proportion of leg segment lengths and actuation system were analyzed in consideration of anatomical structure as well as muscle system of insect legs and desired mobility. A spring based passive compliance mechanism inspired by musculoskeletal structures of biological systems was integrated into distal segment of the leg to soften foot impact on touchdown. In addition, an efficient locomotion planner capable of generating natural movements for the legs during swing phase was proposed. The problem of leg swing was formulated as an optimal control procedure that satisfies a series of locomotion task terms while minimizing a biologically-based objective function, which was solved by a Gauss Pseudospectral Method （GPM） based numerical technique. We applied this swing generation algorithm to both a simulation platform and a robot prototype. Results show that the proposed leg structure and swing planner are able to successfully perform effective swing movements on rugged terrains.     

A cockroach that jumps

We report on a newly discovered cockroach (Saltoblattella montistabularis) from South Africa, which jumps and therefore differs from all other extant cockroaches that have a scuttling locomotion. In its natural shrubland habitat, jumping and hopping accounted for 71 per cent of locomotory activity. Jumps are powered by rapid and synchronous extension of the hind legs that are twice the length of the other legs and make up 10 per cent of the body weight. In high-speed images of the best jumps the body was accelerated in 10 ms to a take-off velocity of 2.1 m s(-1) so that the cockroach experienced the equivalent of 23 times gravity while leaping a forward distance of 48 times its body length. Such jumps required 38 μJ of energy, a power output of 3.4 mW and exerted a ground reaction force through both hind legs of 4 mN. The large hind legs have grooved femora into which the tibiae engage fully in advance of a jump, and have resilin, an elastic protein, at the femoro-tibial joint. The extensor tibiae muscles contracted for 224 ms before the hind legs moved, indicating that energy must be stored and then released suddenly in a catapult action to propel a jump. Overall, the jumping mechanisms and anatomical features show remarkable convergence with those of grasshoppers with whom they share their habitat and which they rival in jumping performance.     

On the applicability of the decentralized control mechanism extracted from the true slime mold: a robotic case study with a serpentine robot

A systematic method for an autonomous decentralized control system is still lacking, despite its appealing concept. In order to alleviate this, we focused on the amoeboid locomotion of the true slime mold, and extracted a design scheme for the decentralized control mechanism that leads to adaptive behavior for the entire system, based on the so-called discrepancy function. In this paper, we intensively investigate the universality of this design scheme by applying it to a different type of locomotion based on a 'synthetic approach'. As a first step, we implement this design scheme to the control of a real physical two-dimensional serpentine robot that exhibits slithering locomotion. The experimental results show that the robot exhibits adaptive behavior and responds to the environmental changes; it is also robust against malfunctions of the body segments due to the local sensory feedback control that is based on the discrepancy function. We expect the results to shed new light on the methodology of autonomous decentralized control systems.     

Energy management that generates terrain following versus apex-preserving hopping in man and machine

While hopping, 12 subjects experienced a sudden step down of 5 or 10 cm. Results revealed that the hopping style was “terrain following”. It means that the subjects pursued to keep the distance between maximum hopping height (apex) and ground profile constant. The spring-loaded inverse pendulum (SLIP) model, however, which is currently considered as template for stable legged locomotion would predict apex-preserving hopping, by which the absolute maximal hopping height is kept constant regardless of changes of the ground level. To get more insight into the physics of hopping, we outlined two concepts of energy management: “constant energy supply”, by which in each bounce—regardless of perturbations—the same amount of mechanical energy is injected, and “lost energy supply”, by which the mechanical energy that is going to be dissipated in the current cycle is assessed and replenished. When tested by simulations and on a robot testbed capable of hopping, constant energy supply generated stable and robust terrain following hopping, whereas lost energy supply led to something like apex-preserving hopping, which, however, lacks stability as well as robustness. Comparing simulated and machine hopping with human hopping suggests that constant energy supply has a good chance to be used by humans to generate hopping.     

Quantitative estimation of pancreatic polypeptide (PP) cells of islet tissue of pancreas in Australian mammals

Pancreases from three male and three female echidnas ( Tachyglossus aculeatus ), possums ( Trichosurus vulpecula ), grey kangaroos ( Macropus fuliginosus ), spinifex hopping mice ( Notomys alexis ) and water rats ( Hydromys chrysogaster ) were studied using a point-intercept method and Weibel graticule (Weibel, Kistler & Scherle, 1966) on sections stained by an immunoperoxidase method for pancreatic polypeptide (PP). PP cells were assessed in the head, neck, body and tail regions. There was a greater percentage proportion of PP cells in the head than in the other regions of echidna, possum and water rat. PP cells were not identified in grey kangaroo and spinifex hopping mouse, although control sections were positive.
These results show some agreement with a previously proposed hypothesis (Bonner-Weir & Weir, 1979) concerning the distribution of cell types in the mammalian endocrine pancreas.     

Scaling of elastic energy storage in mammalian limb tendons: do small mammals really lose out?

It is widely believed that elastic energy storage is more important in the locomotion of larger mammals. This is based on: (a) comparison of kangaroos with the smaller kangaroo rat; and (b) calculations that predict that the capacity for elastic energy storage relative to body mass increases with size. Here we argue that: (i) data from kangaroos and kangaroo rats cannot be generalized to other mammals; (ii) the elastic energy storage capacity relative to body mass is not indicative of the importance of elastic energy to an animal; and (iii) the contribution of elastic energy to the mechanical work of locomotion will not increase as rapidly with size as the mass-specific energy storage capacity, because larger mammals must do relatively more mechanical work per stride. We predict how the ratio of elastic energy storage to mechanical work will change with size in quadrupedal mammals by combining empirical scaling relationships from the literature. The results suggest that the percentage contribution of elastic energy to the mechanical work of locomotion decreases with size, so that elastic energy is more important in the locomotion of smaller mammals. This now needs to be tested experimentally.     

Human hopping on damped surfaces: strategies for adjusting leg mechanics

Fast-moving legged animals bounce along the ground with spring-like legs and agilely traverse variable terrain. Previous research has shown that hopping and running humans maintain the same bouncing movement of the body's centre of mass on a range of elastic surfaces by adjusting their spring-like legs to exactly offset changes in surface stiffness. This study investigated human hopping on damped surfaces that dissipated up to 72% of the hopper's mechanical energy. On these surfaces, the legs did not act like pure springs. Leg muscles performed up to 24-fold more net work to replace the energy lost by the damped surface. However, considering the leg and surface together, the combination appeared to behave like a constant stiffness spring on all damped surfaces. By conserving the mechanics of the leg-surface combination regardless of surface damping, hoppers also conserved centre-of-mass motions. Thus, the normal bouncing movements of the centre of mass in hopping are not always a direct result of spring-like leg behaviour. Conserving the trajectory of the centre of mass by maintaining spring-like mechanics of the leg-surface combination may be an important control strategy for fast-legged locomotion on variable terrain.     

Parasitic Robot System for Waypoint Navigation of Turtle

In research on small mobile robots and biomimetic robots,locomotion ability remains a major issue despite many advances in technology.However,evolution has led to there being many real animals capable of excellent locomotion.This paper presents a "parasitic robot system" whereby locomotion abilities of an animal are applied to a robot task.We chose a turtle as our first host animal and designed a parasitic robot that can perform "operant conditioning".The parasitic robot,which is attached to the turtle,can induce object-tracking behavior of the turtle toward a Light Emitting Diode (LED) and positively reinforce the behavior through repeated stimulus-response interaction.After training sessions over five weeks,the robot could successfully control the direction of movement of the trained turtles in the waypoint navigation task.This hybrid animal-robot interaction system could provide an alternative solution to some of the limitations of conventional mobile robot systems in various fields,and could also act as a useful interaction system for the behavioral sciences.     

Kinematic Analysis and Experimental Verification on the Locomotion of Gecko

This paper presents a kinematic analysis of the locomotion of a gecko,and experimental verification of the kinematicmodel.Kinematic analysis is important for parameter design,dynamic analysis,and optimization in biomimetic robot research.The proposed kinematic analysis can simulate,without iteration,the locomotion of gecko satisfying the constraint conditionsthat maintain the position of the contacted feet on the surface.So the method has an advantage for analyzing the climbing motionof the quadruped mechanism in a real time application.The kinematic model of a gecko consists of four legs based on 7-degreesof freedom spherical-revolute-spherical joints and two revolute joints in the waist.The motion of the kinematic model issimulated based on measurement data of each joint.The motion of the kinematic model simulates the investigated real gecko’smotion by using the experimental results.The analysis solves the forward kinematics by considering the model as a combinationof closed and open serial mechanisms under the condition that maintains the contact positions of the attached feet on the ground.The motions of each joint are validated by comparing with the experimental results.In addition to the measured gait,three othergaits are simulated based on the kinematic model.The maximum strides of each gait are calculated by workspace analysis.Theresult can be used in biomimetic robot design and motion planning.