Quadrupedal Gait Research Articles

Adaptations of large marsupials to survival in winter snow cover: locomotion and foraging

The small extent of seasonally snow-covered Australian mountains means that there has not been a great selective pressure on the mammalian fauna for adaptations to this environment. Only one large marsupial, the common wombat (Vombatus ursinus (Shaw, 1800)), is widespread above the winter snowline. In the past 20 years, with snow depth and duration declining, the swamp wallaby (Wallabia bicolor (Desmarest, 1804)) has become more common above the winter snowline. The red-necked wallaby (Macropus rufogriseus (Desmarest, 1817)) is common in alpine Tasmania where seasonal snow cover is neither as deep nor as long-lasting as on the mainland, but has only been recorded regularly above the winter snowline in the mainland Snowy Mountains since 2011. This study examines morphological aspects of locomotion of these three herbivorous marsupials in snow. The wombat is the best adapted to snow, with quadrupedal gait and an expanded home range allowing it to locate and feed on the same plant groups as it does at lower elevation. Wallabies are poorly adapted to locomotion in snow, but the browsing swamp wallaby is able to maintain its dietary habit by feeding on exposed shrubs in deep snow, whereas the red-necked wallaby, which depends more on grazing, appears constrained to areas where snow is shallow.

A hyper-robust sauropodomorph dinosaur ilium from the Upper Triassic–Lower Jurassic Elliot Formation of South Africa: Implications for the functional diversity of basal Sauropodomorpha

It has generally been held that the locomotory habits of sauropodomorph dinosaurs moved in a relatively linear evolutionary progression from bipedal through “semi-bipedal” to the fully quadrupedal gait of Sauropoda. However, there is now a growing appreciation of the range of locomotory strategies practiced amongst contemporaneous taxa of the latest Triassic and earliest Jurassic. Here we present on the anatomy of a hyper-robust basal sauropodomorph ilium from the Late Triassic–Early Jurassic Elliot Formation of South Africa. This element, in addition to highlighting the unexpected range of bauplan diversity throughout basal Sauropodomorpha, also has implications for our understanding of the relevance of “robusticity” to sauropodomorph evolution beyond generalized limb scaling relationships. Possibly representing a unique form of hindlimb stabilization during phases of bipedal locomotion, the autapomorphic morphology of this newly rediscovered ilium provides additional insight into the myriad ways in which basal Sauropodomorpha managed the inherited behavioural and biomechanical challenges of increasing body-size, hyper-herbivory, and a forelimb primarily adapted for use in a bipedal context.

Effects of Substrate Size and Orientation on Quadrupedal Gait Kinematics in Mouse Lemurs (Microcebus murinus).

As the smallest living primate, the mouse lemur is a suitable model for reconstructing the locomotor mechanisms by which primate ancestors might have responded to the challenges of an arboreal environment. In this study, we tested the effects of substrate diameter and orientation on quadrupedal gait kinematics in mouse lemurs (Microcebus murinus). Mouse lemurs highly preferred asymmetrical to symmetrical gaits as they moved across a flat board and poles of three diameters (2.5, 1.0, and 0.5cm), set at horizontal, 30° inclined, and 30° declined orientations. During symmetrical gaits, mouse lemurs used diagonal sequence walking and ambling gaits on the same substrates and at the same duty factors for which some similarly sized nonprimate mammals use lateral sequence gaits, suggesting that reliance on diagonal sequence walking in primates may not be explicitly a response to body size relative to substrate diameter. When using asymmetrical gaits, kinematic adjustments to small diameter and/or nonhorizontal substrates included a preference for transverse gallops over other gaits, the avoidance of whole-body suspensions, increases in limb contact duration, and increases in the time interval between the landing of trailing and leading limbs. All of these adjustments are consistent with increasing locomotor stability by dampening center of mass movements and reducing the forces imparted to the substrate. Like mouse lemurs, small-bodied ancestral primates likely used symmetrical gaits occasionally, but more frequently used asymmetrical gaits that were adjusted in response to challenging substrates. Therefore, asymmetrical gait dynamics should be incorporated into hypotheses addressing early primate locomotor evolution.

Patterns of quadrupedal locomotion in a vertical clinging and leaping primate (Propithecus coquereli) with implications for understanding the functional demands of primate quadrupedal locomotion.

Many primates exhibit a suite of characteristics that distinguish their quadrupedal gaits from non-primate mammals including the use of a diagonal sequence gait, a relatively protracted humerus at touchdown, and relatively high peak vertical forces on the hindlimbs compared to the forelimbs. These characteristics are thought to have evolved together in early, small-bodied primates possibly in response to the mechanical demands of navigating and foraging in a complex arboreal environment. It remains unclear, however, whether primates that employ quadrupedalism only rarely demonstrate the common primate pattern of quadrupedalism or instead use the common non-primate pattern or an entirely different mechanical pattern from either group. This study compared the kinematics and kinetics of two habitually quadrupedal primates (Lemur catta and Varecia variegata) to those of a dedicated vertical clinger and leaper (Propithecus coquereli) during bouts of quadrupedal walking. All three species employed diagonal sequence gaits almost exclusively, displayed similar degrees of humeral protraction, and exhibited lower vertical peak forces in the forelimbs compared to the hindlimb. From the data in this study, it is possible to reject the idea that P. coquereli uses a non-primate pattern of quadrupedal walking mechanics. Nor do they use an entirely different mechanical pattern from either most primates or most non-primates during quadrupedal locomotion. These findings provide support for the idea that this suite of characteristics is adaptive for the challenges of arboreal locomotion in primates and that these features of primate locomotion may be basal to the order or evolved independently in multiple lineages including indriids. Am J Phys Anthropol 160:644-652, 2016. © 2016 Wiley Periodicals, Inc.

A human quadrupedal gait following poliomyelitis: From the Dercum-Muybridge collaboration (1885).

Beginning in the late 1870s, before the invention of movie cameras or projectors, pioneering English American photographer Eadweard Muybridge photographed iconic image sequences of people and animals in motion using arrays of sequentially triggered single-image cameras. In 1885, Philadelphia neurologist Francis Dercum initiated a collaborative relationship with Muybridge at the University of Pennsylvania to photograph sequential images of patients with various neurologic disorders of movement, including an acquired pathologic quadrupedal gait in a young boy that developed as a consequence of poliomyelitis. This pathologic human quadrupedal gait was compared with other quadrupedal gaits filmed by Muybridge, including a toddler girl and an adult woman crawling on hands and knees, an adult woman bear crawling on hands and feet, and a baboon walking. All of the human quadrupedal gaits were lateral sequence gaits, whereas the baboon's walking gait was a diagonal sequence gait. Modern studies have confirmed the nonpathologic quadrupedal gait sequences of humans and nonhuman primates. Despite Dercum's assertion to the contrary, the limb placement pattern of the boy with a pathologic quadrupedal gait after poliomyelitis was not the typical gait of a primate quadruped, but rather was the typical gait sequence for normal human developmental and volitional quadrupedal gaits.

Quadrupedal movement training improves markers of cognition and joint repositioning

IntroductionExercise, and in particular balance and coordination related activities such as dance, appear to have positive effects on cognitive function, as well as neurodegenerative conditions such as dementia and Parkinson’s disease. Quadrupedal gait training is a movement system requiring coordination of all four limbs that has previously been associated with cognitive development in children. There is currently little research into the effect of complex QDP movements on cognitive function in adults. PurposeTo determine the effects of a novel four-week quadrupedal gait training programme on markers of cognitive function and joint reposition sense in healthy adults. MethodsTwenty-two physically active sports science students (15 male and 7 female) were divided into two groups: a training group (TG) and a control group (CG). All participants completed the Wisconsin Card Sorting Task (WCST) and were tested for joint reposition sense before and after a four-week intervention, during which time the TG completed a series of progressive and challenging quadrupedal movement training sessions. ResultsParticipants in the TG showed significant improvements in the WCST, with improvements in perseverative errors, non-perseverative errors, and conceptual level response. This improvement was not found in the CG. Joint reposition sense also improved for the TG, but only at 20degrees of shoulder flexion. ConclusionsPerformance of a novel, progressive, and challenging task, requiring the coordination of all 4 limbs, has a beneficial impact on cognitive flexibility, and in joint reposition sense, although only at the specific joint angle directly targeted by the training. The findings are consistent with other studies showing improvements in executive function and joint reposition sense following physical activity.

Passive Dynamics Explain Quadrupedal Walking, Trotting, and Tölting.

This paper presents a simplistic passive dynamic model that is able to create realistic quadrupedal walking, tölting, and trotting motions. The model is inspired by the bipedal spring loaded inverted pendulum (SLIP) model and consists of a distributed mass on four massless legs. Each of the legs is either in ground contact, retracted for swing, or is ready for touch down with a predefined angle of attack. Different gaits, that is, periodic motions differing in interlimb coordination patterns, are generated by choosing different initial model states. Contact patterns and ground reaction forces (GRFs) evolve solely from these initial conditions. By identifying appropriate system parameters in an optimization framework, the model is able to closely match experimentally recorded vertical GRFs of walking and trotting of Warmblood horses, and of tölting of Icelandic horses. In a detailed study, we investigated the sensitivity of the obtained solutions with respect to all states and parameters and quantified the improvement in fitting GRF by including an additional head and neck segment. Our work suggests that quadrupedal gaits are merely different dynamic modes of the same structural system and that we can interpret different gaits as different nonlinear elastic oscillations that propel an animal forward.

A galloping quadruped model using left–right asymmetry in touchdown angles

Among quadrupedal gaits, the galloping gait has specific characteristics in terms of locomotor behavior. In particular, it shows a left–right asymmetry in gait parameters such as touchdown angle and the relative phase of limb movements. In addition, asymmetric gait parameters show a characteristic dependence on locomotion speed. There are two types of galloping gaits in quadruped animals: the transverse gallop, often observed in horses; and the rotary gallop, often observed in dogs and cheetahs. These two gaits have different footfall sequences. Although these specific characteristics in quadrupedal galloping gaits have been observed and described in detail, the underlying mechanisms remain unclear. In this paper, we use a simple physical model with a rigid body and four massless springs and incorporate the left–right asymmetry of touchdown angles. Our simulation results show that our model produces stable galloping gaits for certain combinations of model parameters and explains these specific characteristics observed in the quadrupedal galloping gait. The results are then evaluated in comparison with the measured data of quadruped animals and the gait mechanisms are clarified from the viewpoint of dynamics, such as the roles of the left–right touchdown angle difference in the generation of galloping gaits and energy transfer during one gait cycle to produce two different galloping gaits.

Humanoids can take Advantage of Crab-Walking Gaits

Recent advances in control of humanoid robots have resulted in bipedal gaits that are dynamically stable on moderately rough terrain but are still limited to a small range of slopes. Humanoid robots, like humans, can take advantage of quadruped gaits to greatly extend this range. Cleverly designed gaits can provide robustness to rough terrain without requiring extensive feedback. In this paper, we present a robust crab-walking framework that includes forward and backward crawling patterns, rotation patterns, and sit-down and recovery sequences. The latter are activated autonomously once the robot detects that it tipped over. The performance and robustness of each locomotion pattern are investigated over a wide range of slopes. Crab-walking is shown to be especially adept at crawling forward on steep downward slopes (up to -54°) and crawling backward on steep upward slopes (up to 18°). Finally, we demonstrate the framework's autonomous capabilities by crossing the rough terrain in DARPA's virtual robotics challenge.

Reachability-based control for the active SLIP model

One of the key topics in robotics is legged locomotion, taking inspiration from walking and bouncing gaits of bipeds and quadrupeds. A fundamental tool for analyzing running and hopping is the spring loaded inverted pendulum (SLIP), thanks to its simplicity but also effectiveness in modeling bouncing gaits. Being completely passive, the SLIP model does not have the ability to modify its net energy, which, for example, can be a prohibitive obstacle when traveling on a terrain with varying heights. The actuated version of the SLIP model, the active SLIP, considered in this paper, includes a series actuator that allows energy variations via compressing or extending the spring. In this work, we investigate how different actuator motions can affect the system’s state, and we propose a control strategy, based on graphical and numerical studies of the reachability space, and updated throughout the stance phase, to drive the system to a desired state. We quantify its performance benefits, particularly in serving as an error-recovering method. The objective of our control strategy is not to replace any leg-placement approach proposed by other works, but rather to be paired with any other leg-placement or path planning method. Its main advantage is the ability to reduce the effects of sensing errors and disturbances happening at landing as well as during the stance phase.

A simple rule for quadrupedal gait generation determined by leg loading feedback: a modeling study.

We discovered a specific rule for generating typical quadrupedal gaits (the order of the movement of four legs) through a simulated quadrupedal locomotion, in which unprogrammed gaits (diagonal/lateral sequence walks, left/right-lead canters, and left/right-lead transverse gallops) spontaneously emerged because of leg loading feedbacks to the CPGs hard-wired to produce a default trot. Additionally, all gaits transitioned according to speed, as seen in animals. We have therefore hypothesized that various gaits derive from a trot because of posture control through leg loading feedback. The body tilt on the two support legs of each diagonal pair during trotting was classified into three types (level, tilted up, or tilted down) according to speed. The load difference between the two legs led to the phase difference between their CPGs via the loading feedbacks, resulting in nine gaits (32: three tilts to the power of two diagonal pairs) including the aforementioned.

First Report of Small Ornithopodichnus Trackways from the Lower Cretaceous of Sichuan, China

Trackways of ornithopods are well-known from the Lower Cretaceous of East Asia, particularly in Korea and China. However, most morphotypes have been identified as Caririchnium which is characterized by moderate mesaxony, the ratio length = width, or length > width and, in some cases, by a quadrupedal gait. In 2009 the ichnogenus Ornithopodichnus, which exhibits distinctively weak mesaxony and a broader, transverse pes imprint with the ratio length < width, was identified in Korea, on the basis of material with suboptimal preservation. The trackmaker was a large ornithopod. Subsequently, in 2009 a second report recognized Ornithopodichnus in China, the trackmaker being a moderately large quadruped. In 2012, Ornithopodichnus was described on the basis of well-preserved material from the Upper Cretaceous of Korea, in this case left by a small, gregarious biped. Herein we document, a fourth occurrence of Ornithopodichnus in the Lower Cretaceous Feitianshan Formation of Sichuan, China which is a further example of trackways that can be attributed to small, bipedal and gregarious ornithopods. Morphology, size, trackway pattern and speed of locomotion correspond well with trackways known from the Korean Hwasun tracksite. Possibly, the occurrence of Ornithopodichnus which is restricted to East Asia thus far, is related to ecological and biogeographical pecularities in this area.

The invisible gorilla! A visual note of inattentional blindness

Fig. 1. The image displays 33 whole body volume renderings in a 3D skeletal view. The postmortem scan of a gorilla, “invisible gorilla”, is hidden in the last row, 3rd skeletal whole body volume rendering from the right. Gorillas are the largest living primates and other than humans who have biped gait, gorillas do knuckle walking (quadrupedal gait). Therefore the pelvis in gorillas is relatively narrow compared to humans and long arms, bent knees and flat, wide feet support shuffling on all fours. In addition, the face is larger than the cranium, which is vice versa in humans and the skull hangs forward and is not supported on the top of the vertebral column. Gorillas are also primarily vegetarians.

Dynamics of locomotor transitions from arboreal to terrestrial substrates in Verreaux's sifaka (Propithecus verreauxi).

Most primates are able to move with equal facility on the ground and in trees, but most use the same quadrupedal gaits in both environments. A few specialized primates, however, use a suspensory or leaping mode of locomotion when in the trees but a bipedal gait while on the ground. This is a rare behavioral pattern among mammals, and the extent to which the bipedal gaits of these primates converge and are constrained by the anatomical and neurological adaptations associated with arboreal locomotion is poorly understood. Sifakas (Propithecus), primates living only in Madagascar, are highly committed vertical clingers and leapers that also spend a substantial amount of time on the ground. When moving terrestrially sifakas use a unique bipedal galloping gait seen in no other mammals. Little research has examined the mechanics of these gaits, and most of that research has been restricted to controlled captive conditions. The energetic costs associated with leaping and bipedal galloping are unknown. This study begins to fill that gap using triaxial accelerometry to characterize and compare the dynamics of sifakas' leaping and bipedal galloping behavior. As this is a relatively novel approach, the first goal of this article is to explore the feasibility of collecting such data on free-roaming animals and attempt to automate the identification of leaping and bipedal behavior within the output. The second goal is to compare the overall accelerations of the body and to use that as an approximation of aspects of energetic costs during leaping and bipedalism. To achieve this, a lightweight accelerometer was mounted on freely moving sifakas. The resulting acceleration profiles were processed, and sequences of leaps (bouts) were automatically extracted from the waveforms with 85% accuracy. Both vector dynamic body acceleration and overall dynamic body acceleration (ODBA) were used to characterize locomotor patterns and energy expenditure during leaping and bipedalism. The unique kinematics of the gait of sifakas, and the mechanics of bouts involving a string of successive leaps or gallops, appear to minimize redirections of the center of mass as well as the number of acceleration peaks and ODBAs. These results suggest that bipedal galloping is not only a reflection of the unique anatomical configuration of a leaping primate, but it may also provide a musculoskeletal and an energetic advantage to sifakas. In that sense, bipedal galloping represents an advantageous way for sifakas to move when transitioning from arboreal leaping to terrestrial locomotion.

Gait analysis methods for rodent models of osteoarthritis.

Patients with osteoarthritis (OA) primarily seek treatment due to pain and disability, yet the primary endpoints for rodent OA models tend to be histological measures of joint destruction. The discrepancy between clinical and preclinical evaluations is problematic, given that radiographic evidence of OA in humans does not always correlate to the severity of patient-reported symptoms. Recent advances in behavioral analyses have provided new methods to evaluate disease sequelae in rodents. Of particular relevance to rodent OA models are methods to assess rodent gait. While obvious differences exist between quadrupedal and bipedal gait sequences, the gait abnormalities seen in humans and in rodent OA models reflect similar compensatory behaviors that protect an injured limb from loading. The purpose of this review is to describe these compensations and current methods used to assess rodent gait characteristics, while detailing important considerations for the selection of gait analysis methods in rodent OA models.

Kinematics and spatiotemporal parameters of infant-carrying in olive baboons.

In the field of biomechanics of quadrupedal locomotion in primates, infant-carrying has received little attention. This study presents the first biomechanical study of infant-carrying in captive female olive baboons (Papio anubis). We test whether females carrying infants conform 1) to the Support Polygon Model (Rollinson and Martin: Symp Zool Soc Lond 48 (1981) 377-427) of gait selection, according to which diagonality should decrease when the infant is carried cranially and increase when the infant is carried dorsally and caudally; 2) to Biewener's (Biewener: Science 245 () 45-48) theory of limb postures, according to which females should extend their hind limbs more due to infant load, especially in the later stages when the infant is not fully autonomous but relatively heavy. This study focuses on the sagittal kinematics of quadrupedal gaits (joint angles and spatiotemporal parameters) of four females with and without infant loads at the CNRS Primatology Station (France). High-speed video recordings were made using the technical platform "Motion Analysis of Primates" available in the animals' place of life. Regarding diagonality, our results do not fully conform to those predicted by the Support Polygon Model of gait selection; however, the model cannot be rejected at this stage in experiment. With regard to limb posture, our results do not support Biewener's (Biewener: Science 245 () 45-48) theory: loaded females do not extend their hind limbs more as predicted; on the contrary, the hind limbs tend to be more flexed when the infant they carry is relatively heavy.

Human quadrupeds, primate quadrupedalism, and Uner Tan Syndrome.

Since 2005, an extensive literature documents individuals from several families afflicted with “Uner Tan Syndrome (UTS),” a condition that in its most extreme form is characterized by cerebellar hypoplasia, loss of balance and coordination, impaired cognitive abilities, and habitual quadrupedal gait on hands and feet. Some researchers have interpreted habitual use of quadrupedalism by these individuals from an evolutionary perspective, suggesting that it represents an atavistic expression of our quadrupedal primate ancestry or “devolution.” In support of this idea, individuals with “UTS” are said to use diagonal sequence quadrupedalism, a type of quadrupedal gait that distinguishes primates from most other mammals. Although the use of primate-like quadrupedal gait in humans would not be sufficient to support the conclusion of evolutionary “reversal,” no quantitative gait analyses were presented to support this claim. Using standard gait analysis of 518 quadrupedal strides from video sequences of individuals with “UTS”, we found that these humans almost exclusively used lateral sequence–not diagonal sequence–quadrupedal gaits. The quadrupedal gait of these individuals has therefore been erroneously described as primate-like, further weakening the “devolution” hypothesis. In fact, the quadrupedalism exhibited by individuals with UTS resembles that of healthy adult humans asked to walk quadrupedally in an experimental setting. We conclude that quadrupedalism in healthy adults or those with a physical disability can be explained using biomechanical principles rather than evolutionary assumptions.

Central pattern generator based reflexive control of quadruped walking robots using a recurrent neural network

This paper presents a novel Central Pattern Generator (CPG) model for controlling quadruped walking robots. The improvement of this model focuses on generating any desired waveforms along with accurate online modulation. In detail, a well-analyzed Recurrent Neural Network is used as the oscillators to generate simple harmonic periodic signals that exhibit limit cycle effects. Then, an approximate Fourier series is employed to transform those mentioned simple signals into arbitrary desired outputs under the phase constraints of several primary quadruped gaits. With comprehensive closed-form equations, the model also allows the user to modulate the waveform, the frequency and the phase constraint of the outputs online by directly setting the inner parameters without the need for any manual tuning. In addition, an associated controller is designed using leg coordination Cartesian position as the control state space based on which stiffness control is performed at sub-controller level. In addition, several reflex modules are embedded to transform the feedback of all sensors into the CPG space. This helps the CPG recognize external disturbances and utilize inner limit cycle effect to stabilize the robot motion. Finally, experiments with a real quadruped robot named AiDIN III performing several dynamic trotting tasks on several unknown natural terrains are presented to validate the effectiveness of the proposed CPG model and controller.

Roles of carbonic anhydrase 8 in neuronal cells and zebrafish

BackgroundCarbonic anhydrase 8 (CA8) is an isozyme of α-carbonic anhydrases (CAs). Previous studies showed that CA8 can be detected in human adult brain, with more intense expression in the cerebellum. Single mutations in CA8 were reported to cause novel syndromes like ataxia, mild mental retardation or the predisposition to quadrupedal gait. MethodsIn the present study, we examine the functions of CA8 in neuronal cell lines, mouse cerebellar granule neurons and zebrafish. Results and conclusionsWe demonstrated that overexpression of CA8 in neuronal cells significantly decreased cell death under staurosporine treatment. Moreover, CA8 overexpression significantly increased cell migration and invasion ability in neuronal cells and in mouse cerebellar granule neurons, implicating that CA8 may be involved in neuron motility and oncogenesis. By using zebrafish as an animal model, motor reflection of 3dpf zebrafish embryos was significantly affected after the down-regulation of CA8 through ca8 morpholino. ConclusionsWe concluded that CA8 overexpression desensitizes neuronal cells to STS induced apoptotic stress and increases cell migration and invasion ability in neuronal cells. In addition, down-regulated CA8 decreases neuron mobility in neuronal cells and leads to abnormal calcium release in cerebellar granule neurons. Knockdown of the ca8 gene results in an abnormal movement pattern in zebrafish. General significanceOur findings provide evidence to support that the impaired protective function of CA8 contributes to human neuropathology, and to suggest that zebrafish can be used as an animal model to study the biological functions of human CA8 in vivo.

A comparative collision-based analysis of human gait

This study compares human walking and running, and places them within the context of other mammalian gaits. We use a collision-based approach to analyse the fundamental dynamics of the centre of mass (CoM) according to three angles derived from the instantaneous force and velocity vectors. These dimensionless angles permit comparisons across gait, species and size. The collision angle Φ, which is equivalent to the dimensionless mechanical cost of transport CoTmech, is found to be three times greater during running than walking of humans. This threefold difference is consistent with previous studies of walking versus trotting of quadrupeds, albeit tends to be greater in the gaits of humans and hopping bipeds than in quadrupeds. Plotting the collision angle Φ together with the angles of the CoM force vector Θ and velocity vector Λ results in the functional grouping of bipedal and quadrupedal gaits according to their CoM dynamics-walking, galloping and ambling are distinguished as separate gaits that employ collision reduction, whereas trotting, running and hopping employ little collision reduction and represent more of a continuum that is influenced by dimensionless speed. Comparable with quadrupedal mammals, collision fraction (the ratio of actual to potential collision) is 0.51 during walking and 0.89 during running, indicating substantial collision reduction during walking, but not running, of humans.