Symmetrical and asymmetrical gaits in the mouse: patterns to increase velocity

The gaits of the adult SWISS mice during treadmill locomotion at velocities ranging from 15 to 85 cm s^–1 have been analysed using a high-speed video camera combined with cinefluoroscopic equipment. The sequences of locomotion were analysed to determine the various space and time parameters of limb kinematics. We found that velocity adjustments are accounted for differently by the stride frequency and the stride length if the animal showed a symmetrical or an asymmetrical gait. In symmetrical gaits, the increase of velocity is provided by an equal increase in the stride length and the stride frequency. In asymmetrical gaits, the increase in velocity is mainly assured by an increase in the stride frequency in velocities ranging from 15 to 29 cm s^–1. Above 68 cm s^–1, velocity increase is achieved by stride length increase. In velocities ranging from 29 to 68 cm s^–1, the contribution of both variables is equal as in symmetrical gaits. Both stance time and swing time shortening contributed to the increase of the stride frequency in both gaits, though with a major contribution from stance time decrease. The pattern of locomotion obtained in a normal mouse should be used as a template for studying locomotor control deficits after lesions or in different mutations affecting the nervous system.     

The movement patterns of lacertid lizards: speed, gait and pauses in Lacerta vivipara

Lacerta vivipara moving across an open space at their normal activity temperature alternate bursts of locomotion with short pauses which tend to occur at the extremes of the limb cycle, i.e. when individual limbs are maximally adducted or retracted and the spinal cord is maximally flexed in the lateral plane. The movement bursts and pauses in adult lizards have mean durations of 0–30 and 012 s, respectively, and within bursts the lizards move at a mean speed of 14–6 cm s^-1. Movement in juvenile lizards is 2–5 times faster (relative to body length) and the pauses are of longer duration (mean = 019 s), giving the locomotion of juveniles a more jerky appearance. Lizards which are chasing crickets increase the speed and the duration of locomo-tory bursts, although the pauses persist. Lizards which are searching for a previously perceived cricket increase pause duration (mean = 0–40 s). Lizards which are fleeing from a sudden disturbance move at almost twice (juveniles) or 3–7 times (adults) the speed of foraging animals: the pauses persist, although at much reduced frequency. Increases in speed result from increases in both stride length (Λ) and stride frequency (n); the ratio Λ/ n appears to remain constant at 006. The significance of these observations is discussed, although the functions of the pauses cannot yet be explained.     

Stride length and its determinants in humans, early hominids, primates, and mammals

Primate stride lengths during quadrupedal locomotion are very long when compared to those of nonprimate quadrupedal mammals at the speed of trot/gallop transition. These exceptional lengths are a consequence of the relatively long limbs of primates and the large angular excursions of their limbs during quadrupedalism. When quadrupedal primates employ bipedal gaits they exhibit much lower angular excursions. Consequently their bipedal stride lengths do not appear to be exceptional in length when compared to other mammals. Angular excursions of the lower limbs of modern humans are not exceptionally large. However, when running, humans exhibit relatively long periods of flight (i.e., they have low duty factors) when compared to other mammals including primates. Because of these long periods of flight and their relative long lower limbs, humans have running stride lengths that are at the lower end of the range of stride lengths of quadrupedal primates. The stride length of the Laetoli hominid trails are evaluated in this context.     

Effects of Microhabitat‐Dependent Predation Risk on Vigilance during Intermittent Locomotion in Psammodromus algirus Lizards

Animals should be able to adjust their behavior by tracking changes in predation risk level continuously. Many animals show a pattern of intermittent locomotion with short pauses that may increase detection and vigilance of predators. These locomotor patterns may depend on the microhabitat structure, which affect predation risk levels. We examined in detail in the laboratory the characteristics of spontaneous locomotion, scanning behavior, and the escape performance of Psammodromus algirus lizards moving in two different microhabitats (leaf litter patches and open sand areas). Results showed that in leaf litter, lizards moved at slower speed and had shorter bursts of locomotion both in distance and duration, than in sand substrates. This locomotor pattern allowed lizards to increase scanning rate and total time spent in vigilance behavior. When lizards were forced to flee, they escaped to longer distances and during more time in open sand areas, but lizards were able to attain similar escape speed in the two substrates. Lizards may be able to compensate the cost of moving between different microhabitats with different predation risk by behaviorally changing their locomotor and vigilance patterns. However, complex interactions between the visibility of lizards to predators and the ability of lizards to detect predators, together with the need of attending simultaneously to other conflicting demands, may lead to apparently non‐intuitive solutions in locomotor patterns and the rate of vigilance behavior.     

Variation in morphology, gait characteristics and speed of locomotion in two populations of lizards

Locomotor behaviour varies between two subspecies of the Spanish wall lizard Podarcis hispanica. One subspecies inhabits the Columbretes islands, the other lives on the Spanish mainland. Size standardized voluntary speeds (as measured in unrestrained laboratory conditions) are lower in the island population (P. h. atrata) than in the mainland population (P. h. hispanica). Maximal running performance (when chased) is much higher in the mainland population than in the island population. High speed video recordings show that subspecies differ in gait characteristics: individuals from the mainland modulate running velocity primarily by modifying stride length, individuals from the island primarily by altering stride frequency. P. h. hispanica's strategy for modulating speed probably allows this mainland subspecies to attain higher maximal speeds than the island subspecies P. h. atrata. Theoretical considerations suggest that at high speeds, P. h. hispanica's running style is energetically more favourable, but this hypothesis awaits experimental verification. We suggest that the differences in locomotion efficiency between the subspecies result from differences in predation pressure between the mainland and the island. The mainland study site has a higher predator diversity and offers less hiding opportunities to the lizards.     

How to climb a tree: lizards accelerate faster,but pause more,when escaping on vertical surfaces

Many species of lizards effectively traverse both two and three‐dimensional habitats. However, few studies have examined maximum locomotor performance on different inclines. Do maximum acceleration and velocity differ on a level and inclined surface? Do lizards pause more on an inclined surface? To address these questions, Sceloporus woodi lizards (N = 12) were run in the laboratory on a level trackway and a vertical tree trunk. This species is known to frequently utilize both vertical and horizontal aspects of its habitat. Average maximum acceleration on the vertical surface exceeded that on the level surface, although average maximum velocity exhibited the opposite pattern. The average number of pauses during level locomotion was lower compared to vertical locomotion. In addition, the average location of the first pause on the level surface was 0.51 m, which is farther than the average for vertical locomotion where the first pause was at 0.35 m. The combination of performance and pause data suggests that the relative lack of pausing during level locomotion allows individuals to reach higher maximum velocities on level surfaces because they accelerate over greater distances. The increased pausing when moving vertically could be a result of high energetic demands of vertical locomotion, or greater microhabitat complexity as a result of branching and/or refuges. The faster acceleration exhibited during vertical locomotion by S. woodi likely offsets the frequent pauses. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 83–90.     

Oxygen delivery does not limit peak running speed during incremental downhill running to exhaustion

Oxygen consumption (VO2), ventilation (VI), respiratory exchange ratio (R), stride frequency and blood lactate concentrations were measured continuously in nine trained athletes during two continuous incremental treadmill runs to exhaustion on gradients of either 0 degree or -3 degrees. Compared to the run at 0 degree gradient, the athletes reached significantly higher maximal treadmill velocities but significantly lower VO2, VI, R and peak blood lactate concentrations (P less than 0.001) during downhill running. These lower VO2 and blood lactate concentrations at exhaustion indicated that factors other than oxygen delivery limited maximal performance during the downhill run. In contrast, stride frequencies were similar at each treadmill velocity; the higher maximal speed during the downhill run was achieved with a significantly longer stride length (P less than 0.001); maximal stride frequency was the same between tests. Equivalent maximal stride frequencies suggested that factors determining the rate of lower limb stride recovery may have limited maximal running speed during downhill running and, possibly, also during horizontal running.     

Pedestrian locomotion energetics and gait characteristics of a diving bird, the great cormorant, Phalacrocorax carbo

Great cormorants Phalacrocorax carbo are foot propelled diving birds that seem poorly suited to locomotion on land. They have relatively short legs, which are presumably adapted for the generation of high forces during the power stroke of aquatic locomotion, and walk with a pronounced "clumsy waddle". We hypothesise (1) that the speed, independent minimum cost of locomotion (C min, ml O2 m(-1)) will be high for cormorants during treadmill exercise, and (2) that cormorants will have a relatively limited speed range in comparison to more cursorial birds. We measured the rate of oxygen consumption (V02) of cormorants during pedestrian locomotion on a treadmill, and filmed them to determine duty factor (the fraction of stride period that the foot is in contact with the ground), foot contact time (tc), stride frequency (f), swing phase duration and stride length. C min was 2.1-fold higher than that predicted by their body mass and phylogenetic position, but was not significantly different from the C min of runners (Galliformes and Struthioniformes). The extrapolated gamma-intercept of the relationship between V02 and speed was 1.9-fold higher than that predicted by allometry. Again, cormorants were not significantly different from runners. Contrary to our hypothesis, we therefore conclude that cormorants do not have high pedestrian transport costs. Cormorants were observed to use a grounded gait with two double support phases at all speeds measured, and showed an apparent gait transition between 0.17 and 0.25 m s(-1). This transition occurs at a Froude number between 0.016 and 0.037, which is lower than the value of approximately 0.5 observed for many other species. However, despite the use of a limited speed range, and a gait transition at relatively low speed, we conclude that the pedestrian locomotion of these foot propelled diving birds is otherwise generally similar to that of cursorial birds at comparable relative velocities.     

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.     

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.     

Explaining the scaling of transport costs: the role of stride frequency and stride length

The mechanisms which enable large animals to transport a unit of body mass through a unit distance at a lower metabolic cost than smaller animals have been the subject of numerous studies. Recent investigations have concluded that stride frequency is a main determinant. We examine the role of both stride frequency and stride length in determining the scaling of the cost of transport.
Slopes for regressions between stride frequency and speed and stride length and speed were determined in four species of rodents. These data were pooled with literature values for the slopes of stride frequency, stride length and cost of locomotion (all vs. speed) for a total of 17 species ranging in size from 30 g to 250 kg. Interspecific equations were calculated for each of these slopes versus body mass, and residuals from these allometric lines were calculated. Residuals were compared to see if variation in the rate of cost increase at a given size is related to variation in the rates of stride frequency and/or stride length increase.
The residual analysis revealed that the variation in transport cost is explicable only in terms of the interaction of stride frequency and stride length slopes. The product of the scaling exponents for stride frequency slope and stride length slope is not significantly different from the scaling exponent for the cost of transport. A model seeking to explain the scaling of the cost of transport must therefore consider the influence of both stride length and stride frequency.
We propose that absolutely longer limbs allow large animals to minimize the rate of increase of stride frequency and stride length with speed, and that this allows utilization of muscles with lower intrinsic rates of contraction, which in turn results in a lower mass-specific cost of transport.     

Hemiplegic gait: a kinematic analysis using walking speed as a basis.

The kinematics of treadmill ambulation of stroke patients (N = 9) and healthy subjects (N = 4) was studied at a wide range of different velocities (i.e. 0.25-1.5 m s-1), with a focus on the transverse rotations of the trunk. Video recordings revealed, for both stroke patients and healthy subjects, similar relations between walking speed and stride length as well as stride frequency. The phase difference between pelvic and thoracic rotations (i.e. trunk rotation) and the total range of trunk rotation were almost linearly related to the walking speed. Healthy subjects showed a marked increase in pelvic rotation from 1 to 1.5 m s-1. Using dimensional analysis in a comparison between stroke patients and healthy subjects, invariances in the coordination of gait were found for stride length, stride frequency, pelvic rotation, and trunk rotation. Constant relations were obtained between, on the one hand, dimensionless velocity and, on the other, dimensionless stride length as well as stride frequency. Transitions were found between the velocities 0.75 and 1 m s-1 for dimensionless pelvic rotation and trunk rotation, indicating that, from this velocity range onwards, pelvic swing lengthens the stride: rotations of pelvis, thorax and trunk become tightly coordinated. On the basis of the dimensionless stride length, stride frequency, pelvic rotation and trunk rotation, deficits in the gait of stroke patients could be quantified. It is concluded that walking speed is an important control parameter, which should be used as a basic variable in the evaluation of the gait of stroke patients.     

Stride lengths and frequencies of arboreal walking in seven species of didelphid marsupials

Didelphid marsupials differ in their use of the forest strata, with corresponding differences in morphology and arboreal walking performances. Similar performances may be reached by different combinations of stride length and frequency, but it has been suggested that arboreal walkers increase velocity by longer strides. Our objective was to determine how stride length and frequency contribute to the velocity in the arboreal walking of seven species of didelphid marsupials of the Atlantic Forest of Brazil. Animals were stimulated to cross five 3-m long horizontal supports of different diameters. The cycle of maximum velocity was chosen to measure relative stride length, frequency, and relative velocity. Except forCaluromys philander, the more arboreal species were faster than the terrestrial species, but maximum velocity of arboreal species was reached by two strategies, increasing stride frequency (Gracilinanus microtarsus, Micoureus demerarae, andDidelphis aurita), or reducing frequency and increasing stride length (Marmosops incanus andC. philander). Increasing velocity in arboreal walking by more frequent strides may reduce oscillations of the body, whereas longer strides may reduce branch swaying. Among the terrestrial species,Philander frenatus performed similarly to more arboreal species, suggesting a potential ability to use the canopy, undetected in field observations.     

Hindlimb morphology and locomotor performance in waders: an evolutionary approach

Locomotion performance (measured as stride frequency and stride length) was studied in 16 species of waders. Differences in hindlimb morphology (osteology and myology) were analysed among species. Evolutionary changes in both locomotion and morphological variables were analysed using comparative methods revealing the existence of some ecomorphological patterns relating these two sets of characters. Evolutionary changes in stride frequency were correlated with changes in the muscles M. iliotibialis cranialis, M. iliotibiales lateralis and M. gastrocnemius, whereas changes in stride length showed correlated evolution with changes in the length of distal segments of the leg. We identify two different evolutionary strategies in locomotion of waders. One is a change in distal leg segments (skeletal system), an adaptive modification that increases stride length; the second is a change in the skeletal-muscular system, providing an increase in muscular performance (force or speed of contraction) in several muscles, and is an adaptation that increases stride frequency.     

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.     

The relationship between limb morphology, kinematics, and force during running: the evolution of locomotor dynamics in lizards

Terrestrial locomotion occurs via the hierarchical links between morphology, kinematics, force, and center-of-mass mechanics. In a phylogenetically broad sample of seven lizard species, we show that morphological variation drives kinematic variation, which, in turn, drives force variation. Species with short limbs use a short stride–high frequency strategy when running at steady-speed and to change speeds. This link between morphology and kinematics results in relatively small vertical forces during the support phase of the stride cycle. Conversely, species with long limbs use a long stride–low frequency strategy, resulting in large vertical forces during the support phase. In view of these findings, we suggest that limb length may predict locomotor energetics in lizards because energetics are largely determined by vertical forces and stride frequency. Additionally, we propose an energetic trade-off with both long- and short-limbed species paying the most energy to move, whereas intermediate-limbed species move using less energy. Finally, when these traits are mapped onto a lizard phylogeny, we show that locomotor functional morphology exhibits both deep phylogenetic effects and contemporary patterns of evolutionary convergence. Overall, the present study provides a foundation for testing hypotheses regarding the integration and evolution of functional traits in lizards and animals in general.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 634–651.     

Quantitative Comparison of the Walk and Trot of Border Collies and Labrador Retrievers,Breeds with Different Performance Requirements

Introduction

We hypothesized that breed differences of Border Collies and Labrador Retrievers would be reflected in the temporospatial characteristics of the walk and trot.

Materials and Methods

Twenty healthy Border Collies and 20 healthy Labrador Retrievers made three passes across a pressure sensing walkway system that recorded quantitative temporospatial information at a walk and a trot. The following variables were measured for each dog: velocity, total pressure index percentage (TPI%), ratio of weight borne on the thoracic vs. pelvic limbs (T/P TPI%), stance time percentage (ST%), and thoracic limb stride length (TSrL).

Results

The mean T/P TPI% for Border Collies at a walk and at a trot were significantly lower than for Labrador Retrievers (p = 0.0007 and p = 0.0003). Border Collies had a significantly lower ST% than Labrador Retrievers for the thoracic limbs and pelvic limbs at a walk (p = 0.0058 and 0.0003) and the trot (p = 0.0280 and 0.0448). There was no relationship between ST% and TSrL in Border Collies and an inverse correlation between ST% and TSrL in Labrador Retrievers (p = 0.0002).

Discussion

Key quantitative gait differences were identified in Border Collies and Labrador Retrievers, which could potentially provide each breed with an advantage for their working function.     

Patterns of axial and appendicular movements during aquatic walking in the salamander Siren lacertina

Most studies of salamander locomotion have focused either on swimming or terrestrial walking, but some salamanders also use limb-based locomotion while submerged under water (aquatic walking). In this study we used video motion analysis to describe the aquatic walking gait of Siren lacertina, an elongate salamander with reduced forelimbs and no hindlimbs. We found that S. lacertina uses a bipedal-undulatory gait, which combines alternating use of the forelimbs with a traveling undulatory wave. Each forelimb is in contact with the substrate for about 50% of the stride cycle and forelimbs have little temporal overlap in contact intervals. We quantified the relative timing and frequency of limb and tail movements and found that, unlike the terrestrial gaits of most salamanders, axial and appendicular movements are decoupled during aquatic walking. We found no significant relationship between stride frequency and aquatic walking velocity, but we did find a statistically significant relationship between tailbeat frequency and aquatic walking velocity, which suggests that aquatic walking speed is mainly modulated by axial movements. By comparing axial wavespeed and distance traveled per tailbeat during swimming (forelimbs not used) and aquatic walking (forelimbs used), we found lower wavespeed and greater distance traveled per tailbeat during aquatic walking. These findings suggest that the reduced forelimbs of S. lacertina contribute to forward propulsion during aquatic walking.     

The evolution of human running: effects of changes in lower-limb length on locomotor economy

Previous studies have differed in expectations about whether long limbs should increase or decrease the energetic cost of locomotion. It has recently been shown that relatively longer lower limbs (relative to body mass) reduce the energetic cost of human walking. Here we report on whether a relationship exists between limb length and cost of human running. Subjects whose measured lower-limb lengths were relatively long or short for their mass (as judged by deviations from predicted values based on a regression of lower-limb length on body mass) were selected. Eighteen human subjects rested in a seated position and ran on a treadmill at 2.68 ms(-1) while their expired gases were collected and analyzed; stride length was determined from videotapes. We found significant negative relationships between relative lower-limb length and two measures of cost. The partial correlation between net cost of transport and lower-limb length controlling for body mass was r=-0.69 (p=0.002). The partial correlation between the gross cost of locomotion at 2.68 ms(-1) and lower-limb length controlling for body mass was r=-0.61 (p=0.009). Thus, subjects with relatively longer lower limbs tend to have lower locomotor costs than those with relatively shorter lower limbs, similar to the results found for human walking. Contrary to general expectation, a linear relationship between stride length and lower-limb length was not found.