A model of bipedal locomotion on compliant legs.

Simple mathematical models capable of walking or running are used to compare the merits of bipedal gaits. Stride length, duty factor (the fraction of the stride, for which the foot is on the ground) and the pattern of force on the ground are varied, and the optimum gait is deemed to be the one that minimizes the positive work that the muscles must perform, per unit distance travelled. Even the simplest model, whose legs have neither mass nor elastic compliance, predicts the changes of duty factor and force pattern that people make as they increase their speed of walking. It predicts a sudden change to running at a critical speed, but this is much faster than the speed at which people make the change. When elastic compliance is incorporated in the model, unnaturally fast walking becomes uncompetitive. However, a slow run with very brief foot contact becomes the optimum gait at low speeds, at which people would walk, unless severe energy dissipation occurs in the compliance. A model whose legs have mass as well as elastic compliance predicts well the relationship between speed and stride length in human walking.     

The Effect of Speed on the Hindlimb Kinematics of the Reeves' Butterfly Lizard,Leiolepis reevesii(Agamidae)

We recorded locomotor performance of Reeves' butterfly lizards(Leiolepis reevesii) on a racetrack and to describe hindlimb kinematic patterns and to evaluate the effect of speed on hindlimb kinematics. The studied lizards predominantly used quadrupedal locomotion at relatively low speeds, but ran bipedally with a digitigrade posture at high speeds. Speed was positively correlated with both stride length and stride frequency, and was negatively correlated with duty factor. Lizards modulated speed probably by a combination of changing frequency and amplitude of limb movements. Within the range of standardized speeds from 50 to 150 cm/s, speed effects on 28 out of a total of 56 kinematic variables were significant. The hip height at footfall increased as speed increased, whereas the amplitude of vertical oscillations of the hip did not vary with speed. The total longitudinal and dorsoventral movements relative to the hip varied with speed for all parts of the limb that were distal to the knee, whereas the lateral movements did not. The knee and ankle angle at footfall varied with speed, but did not at the end of stance. The degree of pelvis rotation during the entire stride cycle did not vary with speed. Our results suggest that pelvic rotation and femoral protraction/retraction have a minor role in modulating speed in L. reevesii.     

The effect of rider weight and additional weight in Icelandic horses in tölt: part II. Stride parameters responses

This study investigated the effects of rider weight in the BW ratio (BWR) range common for Icelandic horses (20% to 35%), on stride parameters in tölt in Icelandic horses. The kinematics of eight experienced Icelandic school horses were measured during an incremental exercise test using a high-speed camera (300 frames/s). Each horse performed five phases (642 m each) in tölt at a BWR between rider (including saddle) and horse starting at 20% (BWR₂₀) and increasing to 25% (BWR₂₅), 30% (BWR₃₀), 35% (BWR₃₅) and finally 20% (BWR_20b) was repeated. One professional rider rode all horses and weight (lead) was added to saddle and rider as needed. For each phase, eight strides at speed of 5.5 m/s were analyzed for stride duration, stride frequency, stride length, duty factor (DF), lateral advanced placement, lateral advanced liftoff, unipedal support (UPS), bipedal support (BPS) and height of front leg action. Stride length became shorter (Y=2.73−0.004x; P<0.01) and more frequent (Y=2.56+0.002x; P<0.001) with added weight. Duty factor and BPS increased with increased BWR (P<0.001), whereas UPS decreased (P<0.001). Lateral advanced timing of limb placement and liftoff and height of front leg action were not affected by BWR (P>0.05). In conclusion, increased BWR decreased stride length and increased DF proportionally to the same extent in all limbs, whereas BPS increased at the expense of decreased UPS. These changes can be expected to decrease tölt quality when subjectively evaluated according to the breeding goals for the Icelandic horse. However, beat, symmetry and height of front leg lifting were not affected by BWR.     

Monkey locomotion on inclined surfaces

Motion pictures were taken of a subadult, female squirrel monkey moving on level and inclined surfaces. The gaits for the various degrees of incline were compared. The noncontact periods were not affected by changes in inclination. The contact periods, which were inversely related to speed, were affected, by changes in slope and intersect constants. When the incline was steep the animal was forced to jump and zig-zag to retain its progression. The affect of inclination was advantageous at 16° and disadvantageous at higher and lower inclines. At the 16° inclination the animal's short fore legs and long hind legs held the body horizontal, cancelling the effect of the incline. It is reasoned that limb proportions may be adaptive for progression on inclined surfaces and that the locomotor niches for the various arboreal primates may be usefully categorized in terms of the inclinations of their substrates.     

Predictors of sprint start speed: the effects of resistive ground-based vs. inclined treadmill training

There is currently no consensus with regard to the most effective method to train for improved acceleration, or with regard to which kinematic variable provides the greatest opportunity for improvement in this important performance characteristic. The purpose of this study was to determine the effects of resistive ground-based speed training and incline treadmill speed training on speed-related kinematic measures and sprint start speed. The hypothesis tested was that incline treadmill training would improve sprint start time, while the ground-based resistive training would not. Corollary hypotheses were that treadmill training would increase stride frequency and ground-based training would not affect kinematics during the sprint start. Thirty-one high school female soccer players (15.7 +/- 0.5 years) were assigned to either treadmill (n = 17) or ground-based (n = 14) training groups and trained 2 times a week for 6 weeks. The treadmill group utilized incline speed training on a treadmill, while the ground-based group utilized partner band resistance ground-based techniques. Three-dimensional motion analysis was used (4.5 m mark) before and after training to quantify kinematics during the fastest of 3 recorded sprint starts (9.1 m). Both groups decreased average sprint start time from 1.75 +/- 0.12 to 1.68 +/- 0.08 seconds (p < 0.001). Training increased stride frequency (p = 0.030) but not stride length. After training, total vertical pelvic displacement and stride length predicted 62% of the variance in sprint start time for the resistive ground-based group, while stride length and stride frequency accounted for 67% prediction of the variance in sprint start time for the treadmill group. The results of this study indicate that both incline treadmill and resistive ground-based training are effective at improving sprint start speed, although they potentially do so through differing mechanisms.     

Bipedal locomotion: effects of speed, size and limb posture in birds and humans

Seven species of ground-dwelling birds (body mass range: 0.045-90 kg) were filmed while walking and running on a treadmill. High-speed light films were also taken of humans to compare kinematic patterns of avian with human bipedalism. Consistent patterns of stride frequency, stride length, step length, duty factor and limb excursion were observed in all species, with most of the variation among species being due to differences in body size. In general, smaller bipeds have higher stride frequencies (α M ^−0.18), shorter stride lengths (α M ^0.38) and more limited ranges of speed within each gait than large bipeds. After normalizing for size (based on Froude number, after Alexander, 1977), remaining kinematic variation is largely due to interspecific differences in posture and relative limb segment lengths. For their size, smaller bipeds have greater step lengths, limb excursion angles and duty factors than large bipeds because of their more crouched posture and greater effective limb length. The most notable differences in limb kinematics between birds and humans occur at the walk-run transition and are maintained as running speed increases. Change of gait is smooth and difficult to discern in birds, but distinct in humans, involving abrupt decreases in step length and duty factor (time of contact) and a corresponding increase in limb swing time. These differences appear to reflect a spring-like run that is stiff in humans (favouring elastic energy recovery) but more compliant in birds (increasing time of ground contact). Differences between birds and humans in balance of the body's centre of mass not only affect femoral orientation and motion, but also affect pattern of limb excursion with speed.     

Locomotion in the quail (Coturnix japonica): the kinematics of walking and increasing speed

Hindlimb segmental kinematics and stride characteristics are quantified in several quail locomoting on a treadmill over a six-fold increase in speed. These data are used to describe the kinematics of a walking stride and to identify which limb elements are used to change stride features as speed increases. In quail, the femur does not move during locomotion and the tarsometatarsus-phalangeal joint is a major moving joint; thus, quail have lost the most proximal moving joint and added one distally. The tibiotarsus and tarsometatarsus act together as a fixed strut swinging from the knee during stance phase (the ankle angle remains constant at a given speed) and the tarsometatarsus-phalangeal joint appears to have a major role in increasing limb length during the propulsive phase of the stride. Speed is increased with greater knee extension and by lengthening the tibiotarsus/tarsometatarsus via increased ankle extension at greater speeds. Because the femur is not moved and three distal elements are, quail move the limb segments through a stride and increase speed in a way fundamentally different from other nonavian vertebrates. However, the three moving joints in quail (the knee, ankle, and tarsometatarsophangeal joint) have strikingly similar kinematics to the analogous moving joints (the hip, knee, and ankle) in other vertebrates. Comparisons to other vertebrates indicate that birds appear to have two modes of limb function (three- and four-segment modes) that vary with speed and locomotory habits.     

The effect of uphill stride manipulation on race walking gait

Stride length analysis represents an easy method for assessing race walking kinematics. However, the stride parameters emerging from such an analysis have never been used to design a training protocol aimed at increasing stride length. With this aim, we investigated the effects of stride frequency manipulation during three weeks of uphill (2%) training on stride length at iso-efficiency speed. Twelve male race walkers were randomly allocated to one of two training groups: stride frequency manipulation (RWM, n=6) and free stride frequency (RWF, n=6). Results. Kinematic parameters measured before and after the 3-week training in RWM showed increased stride length (4.54%; p<0.0001) and contact time (4.58%; p<0.001); inversely, a decreased stride frequency (4.44%; p<0.0001) and internal work (7.09%; p<0.05) were found. In RWF the effect of the training showed a decrease in stride length (1.18%; p<0.0001) and contact time (<1%; p<0.0001) with respect to baseline conditions and an increased stride frequency and internal work of 1.19% (p<0.0001). These results suggest that using slopes (2%) as RWM could help coaches to provide some training methods that would improve an athlete''s performance, through increasing stride length without altering his or her race walking technique or metabolic demands.     

Spatio-temporal gait characteristics of the hind-limb cycles during voluntary bipedal and quadrupedal walking in bonobos (Pan paniscus)

Spatio-temporal gait characteristics (step and stride length, stride frequency, duty factor) were determined for the hind-limb cycles of nine bonobos (Pan paniscus) walking quadrupedally and bipedally at a range of speeds. The data were recalculated to dimensionless quantities according to the principle of dynamic similarity. Lower leg length was used as the reference length. Interindividual variability in speed modulation strategy of bonobos appears to be low. Compared to quadrupedal walking, bipedal bonobos use smaller steps to attain a given speed (differences increase with speed), resulting in shorter strides at a higher frequency. In the context of the ("hybrid") dynamic pattern approach to locomotion (Latach, 1998) we argue that, despite these absolute differences, intended walking speed is the basic control variable which elicits both quadrupedal and bipedal walking kinematics in a similar way. Differences in the initial status of the dynamic system may be responsible for the differences in step length between both gaits. Comparison with data deduced from the literature shows that the effects of walking speed on stride length and frequency are similar in bonobos, common chimpanzees, and humans. This suggests that (at least) within extant homininae, spatio-temporal gait characteristics are highly comparable, and this in spite of obvious differences in mass distribution and bipedal posture.     

Crawling at High Speeds: Steady Level Locomotion in the Spider Cupiennius salei—Global Kinematics and Implications for Centre of Mass Dynamics

Spiders are an old yet very successful predatory group of arthropods. Their locomotor system differs from those of most other arthropods by the lack of extensor muscles in two major leg joints. Though specific functional characteristics can be expected regarding the locomotion dynamics of spiders, this aspect of movement physiology has been only scarcely examined so far.This study presents extensive analyses of a large dataset concerning global kinematics and the implications for dynamics of adult female specimens of the large Central American spider Cupiennius salei (Keyserling). The experiments covered the entire speed-range of straight runs at constant speeds. The analyses revealed specific characteristics of velocity dependent changes in the movements of the individual legs, as well as in the translational and rotational degrees of freedom of both the centre of mass and the body.In contrast to many other fast moving arthropods, C. salei avoid vertical fluctuations of their centre of mass during fast locomotion. Accordingly, aerial phases were not observed here. This behaviour is most likely a consequence of optimising energy expenditure with regard to the specific requirements of spiders'' leg anatomy. A strong synchronisation of two alternating sets of legs appears to play only a minor role in the locomotion of large spiders. Reduced frequency and low centre of mass amplitudes as well as low angular changes of the body axes, in turn, seems to be the result of relatively low leg coordination.     

Effects of tail loss on the movement patterns of the lizard, Psammodromus algirus

1. Many lizards use caudal autotomy as a defensive strategy. However, subsequent costs related to the alteration of locomotor abilities might decrease the fitness of individuals. In this paper, the movement patterns of spontaneously moving Psammodromus algirus lizards and their escape performance running at high speed were compared before and after tail loss. A control tailed group was also studied to assess the repeatability of locomotor patterns between trials.
2. Tail loss had a significant effect on spontaneous movement patterns. Tailless individuals moved at significantly slower speeds during bursts of locomotion, and distances moved within bursts were significantly reduced. The overall time spent pausing increased, and, as a result, overall speeds decreased to an even greater extent than burst speeds. However, mean durations of individual locomotor bursts and mean pause durations did not change significantly after tail loss.
3. Loss of the tail decreased mean stride length, although the positive relation between stride length and speed was retained.
4. Escape performance was also greatly affected; loss of the tail resulted in substantially reduced attained, maximal and overall escape speeds. These changes resulted in shorter escape distances (the time of the first pause after the initiation of the escape response) because the mean duration of escape responses did not change.
5. The relevance of these alterations for the ecology of this species, and how individuals may compensate for the costs of tail loss, favouring autotomy as an escape strategy, are discussed.     

Autotomy and its Effects on Wolf Spider Foraging Success

Few studies have attempted to determine how physical injury affects predators. One of the ways that physical injury can be expressed is by autotomy or the voluntary loss of a body part. Here, we examined whether the loss of specific legs affects the foraging success of the wolf spider Rabidosa santrita (predator) on another species, Pardosa valens (prey). We also wanted to identify whether the loss of legs in both the predator and prey would impact the outcome of a predation event. Both predator and prey were collected from a creek bed at Portal, AZ, in 2012. Predators were randomly assigned groups where all prey items were intact or all prey had one randomly chosen leg IV removed. Within these groups, predators were organized into a control, leg I autotomy, or leg IV autotomy treatment. All predators had their pre‐ and post‐foraging running speed determined. Predators were introduced into chambers with five prey items and allowed to forage for 1 h. The leg position autotomized or the comparison of pre‐ and post‐foraging trials had no effect on predator running speed. Additionally, there was no significant effect of either predator or prey leg treatment on the total proportion of prey items captured by the end of the foraging trials. Survival analyses indicated that intact prey items tended to have a higher survival rate when predators were missing a leg IV than when predators were intact. When both the predator and prey were missing legs, no significant difference in prey survival rates was detected. We suggest that for predators that inhabit complex, heterogeneous habitats and are classified as ambush predators, the loss of a limb may affect prey capture success, especially when the prey is intact, but that increased sample size is necessary to determine whether this trend is significant.     

The integration of lateral gastrocnemius muscle function and kinematics in running turkeys

Animals commonly move over a range of speeds, and encounter considerable variation in habitat structure, such as inclines. Hindlimb kinematics and muscle function in diverse groups of vertebrates are affected by these changes in behavior and habitat structure, providing a fruitful source of variation for studying the integration of kinematics and muscle function. While it has been observed in a variety of vertebrates that muscle length change can be minimal during locomotion, it is unclear how, and to what degree, in vivo muscle length change patterns are integrated with kinematics. We tested the hypothesis that the length of the turkey lateral gastrocnemius (LG), a biarticular muscle that has moments at the ankle and knee, is not solely affected by changes in joint kinematics. We recorded in vivo muscle length changes (using sonomicrometry) and hindlimb movements (using high-speed video) of wild turkeys running on various inclines, and at different speeds. We quantified the relationship between joint angle (knee and ankle separately) and muscle length in freshly euthanized specimens, and then applied an empirically derived correction for changes in pennation angle and tendon strain during locomotion to improve the accuracy of our predicted lengths. We estimated muscle length at four points during each stride and then compared these values with those measured directly. Other than during swing, the predicted changes in muscle length calculated from the changes in joint kinematics did not correspond with our measured values of LG length. Therefore, the lengths at which the LG operates in turkeys are not determined entirely by kinematics. In addition to strain in series elastic components, we hypothesize that heterogeneous strain within muscles, interactions between muscles and muscle pennation angle all contribute to the nonlinear relationship between muscle length changes and kinematics.     

Jumping kinematics in the wandering spider Cupiennius salei

Spiders use hemolymph pressure to extend their legs. This mechanism should be challenged when required to rapidly generate forces during jumping, particularly in large spiders. However, effective use of leg muscles could facilitate rapid jumping. To quantify the contributions of different legs and leg joints, we investigated jumping kinematics by high-speed video recording. We observed two different types of jumps following a disturbance: prepared and unprepared jumps. In unprepared jumps, the animals could jump in any direction away from the disturbance. The remarkable directional flexibility was achieved by flexing the legs on the leading side and extending them on the trailing side. This behaviour is only possible for approximately radial-symmetric leg postures, where each leg can fulfil similar functions. In prepared jumps, the spiders showed characteristic leg positioning and the jumps were directed anteriorly. Immediately after a preliminary countermovement in which the centre of mass was moved backwards and downwards, the jump was executed by extending first the fourth and then the second leg pair. This sequence provided effective acceleration to the centre of mass. At least in the fourth legs, the hydraulic and the muscular mechanism seem to interact to generate ground reaction forces.     

Morphological evolution of spiders predicted by pendulum mechanics

Background

Animals have been hypothesized to benefit from pendulum mechanics during suspensory locomotion, in which the potential energy of gravity is converted into kinetic energy according to the energy-conservation principle. However, no convincing evidence has been found so far. Demonstrating that morphological evolution follows pendulum mechanics is important from a biomechanical point of view because during suspensory locomotion some morphological traits could be decoupled from gravity, thus allowing independent adaptive morphological evolution of these two traits when compared to animals that move standing on their legs; i.e., as inverted pendulums. If the evolution of body shape matches simple pendulum mechanics, animals that move suspending their bodies should evolve relatively longer legs which must confer high moving capabilities.

Methodology/Principal Findings

We tested this hypothesis in spiders, a group of diverse terrestrial generalist predators in which suspensory locomotion has been lost and gained a few times independently during their evolutionary history. In spiders that hang upside-down from their webs, their legs have evolved disproportionately longer relative to their body sizes when compared to spiders that move standing on their legs. In addition, we show how disproportionately longer legs allow spiders to run faster during suspensory locomotion and how these same spiders run at a slower speed on the ground (i.e., as inverted pendulums). Finally, when suspensory spiders are induced to run on the ground, there is a clear trend in which larger suspensory spiders tend to run much more slowly than similar-size spiders that normally move as inverted pendulums (i.e., wandering spiders).

Conclusions/Significance

Several lines of evidence support the hypothesis that spiders have evolved according to the predictions of pendulum mechanics. These findings have potentially important ecological and evolutionary implications since they could partially explain the occurrence of foraging plasticity and dispersal constraints as well as the evolution of sexual size dimorphism and sociality.     

Walking at the preferred stride frequency maximizes local dynamic stability of knee motion

Healthy humans display a preference for walking at a stride frequency dependent on the inertial properties of their legs. Walking at preferred stride frequency (PSF) is predicted to maximize local dynamic stability, whereby sensitivity to intrinsic perturbations arising from natural variability inherent in biological motion is minimized. Previous studies testing this prediction have employed different variability measures, but none have directly quantified local dynamic stability by computing maximum finite-time Lyapunov exponent (λ^Max), which quantifies the rate of divergence of nearby trajectories in state space. Here, ten healthy adults walked 45 m overground while sagittal motion of both knees was recorded via electrogoniometers. An auditory metronome prescribed 7 different frequencies relative to each individual's PSF (PSF; ±5, ±10, ±15 strides/min). Stride frequencies were performed under both freely adopted speed (FS) and controlled speed (CS: set at the speed of PSF trials) conditions. Local dynamic stability was maximal (λ^Max was minimal) at the PSF, becoming less stable for higher and lower stride frequencies. This occurred under both FS and CS conditions, although controlling speed further reduced local dynamic stability at non-preferred stride frequencies. In contrast, measures of variability revealed effects of stride frequency and speed conditions that were distinct from λ^Max. In particular, movement regularity computed by approximate entropy (ApEn) increased for slower walking speeds, appearing to depend on speed rather than stride frequency. The cadence freely adopted by humans has the benefit of maximizing local dynamic stability, which can be interpreted as humans tuning to their resonant frequency of walking.     

Three-dimensional kinematics of capuchin monkey bipedalism

Capuchin monkeys are known to use bipedalism when transporting food items and tools. The bipedal gait of two capuchin monkeys in the laboratory was studied with three-dimensional kinematics. Capuchins progress bipedally with a bent-hip, bent-knee gait. The knee collapses into flexion during stance and the hip drops in height. The knee is also highly flexed during swing to allow the foot which is plantarflexed to clear the ground. The forefoot makes first contact at touchdown. Stride frequency is high, and stride length and limb excursion low. Hind limb retraction is limited, presumably to reduce the pitch moment of the forward-leaning trunk. Unlike human bipedalism, the bipedal gait of capuchins is not a vaulting gait, and energy recovery from pendulum-like exchanges is unlikely. It extends into speeds at which humans and other animals run, but without a human-like gait transition. In this respect it resembles avian bipedal gaits. It remains to be tested whether energy is recovered through cyclic elastic storage and release as in bipedal birds at higher speeds. Capuchin bipedalism has many features in common with the facultative bipedalism of other primates which is further evidence for restrictions on a fully upright striding gait in primates that transition to bipedalism. It differs from the facultative bipedalism of other primates in the lack of an extended double-support phase and short aerial phases at higher speeds that make it a run by kinematic definition. This demonstrates that facultative bipedalism of quadrupedal primates need not necessarily be a walking gait.     

Descending control of body attitude in the cockroach <Emphasis Type="Italic">Blaberus discoidalis</Emphasis> and its role in incline climbing

Unlike intact animals, cockroaches with bilateral circumoesophageal connective lesions have difficulty climbing up smooth inclines. Typically, they slipped badly or even fell over backward before traveling more than 1.5 body lengths up the incline. The major problem involves increased slippage of the front and sometimes middle legs. Periods of front leg slipping are correlated with excess body elevation that pushes the height of the head to 11 mm above the substrate. Intact animals control body attitude very well on the incline, only rarely rearing above 11 mm. Cockroaches with bilateral circumoesophageal connective lesions spend considerable time above this critical amplitude and slipping increases with amplitude above that value. We conclude that circumoesophageal lesion compromises the insects ability to control body attitude on the incline and this deficit contributes to the inability of lesioned cockroaches to climb steep inclines. A separate body attitude deficit was noted on steps. Upon climbing to the top of a barrier, intact animals bend their body between the first and second thoracic segments. This action maintains good leg mechanics throughout the climb. Cockroaches with bilateral circumoesophageal connective lesions fail to perform this downward flexion. A beneficial role for similar body flexion in simple robots is described in the Discussion.     

Sprawling locomotion in the lizard Sceloporus clarkii: the effects of speed on gait, hindlimb kinematics, and axial bending during walking

Although the hindlimb is widely considered to provide the propulsive force in lizard locomotion, no study to date has analysed kinematic patterns of hindlimb movements for more than one stride for a single individual and no study has considered limb and axial kinematics together. In this study, kinematic data from several individuals of the Sceloporus clarkii are used to describe the movement patterns of the axial skeleton and hindlimb at different speeds, to analyse how kinematics change with speed, and to compare and contrast these findings with the inferred effects of speed cited in the literature. Angular limb movements and axial bending patterns (standing wave with nodes on the girdles) did not change with speed. Only the relative speed of retracting the femur and flexing the knee during limb retraction changes with speed. Based on these data and similar results from a recent study of salamanders, it appears that, over a range of speeds involving a walking trot, sprawling vertebrates increase speed by simply retracting the femur relatively faster, thus this simple functional adjustment may be a general mechanism to increase speed in tetrapods. The demonstration that femoral retraction alone is the major speed effector in Sceloporus clarkii lends strong functional support to ecomorphological implications of limb length (and especially femur length and caudifemoralis size) in locomotory ecology and performance in phrynosomatid lizards. It also lends support to inferences about the caudifemoralis muscle as a preadaptation to terrestrial locomotion and as a key innovation in the evolution of bipedalism.