Suspensory locomotion of Lagothrix lagothricha and Ateles belzebuth in Yasuní National Park,Ecuador

A comparative field study of the locomotion of woolly monkeys (Lagothrix lagothricha) and spider monkeys (Ateles belzebuth) in undisturbed rainforest of northeastern Ecuador reveals substantial differences in their use of suspensory modes. Ateles performed both more brachiation (by forelimbs and tail, with trunk rotation), and forelimb swing (similar to brachiation, but without trunk rotation) than Lagothrix. In contrast, in Lagothrix 20% of suspensory movement was by pronograde forelimb swing, which resembles forelimb swing except that the body is held in a pronograde orientation due to the tail and/or feet intermittently grasping behind the trailing forelimb. Ateles never exhibited this mode. Both brachiation and forelimb swing by Ateles were more dynamic than in Lagothrix, consisting of higher proportions of full-stride bouts (versus single-step). Both species used smaller supports for suspensory than for quadrupedal locomotion, and Ateles used both smaller and larger supports for suspension than did Lagothrix. Analysis of support inclination shows that both species tended to perform more above-support movement on horizontal supports and more below-support (suspensory) movement from oblique supports. Our attempt to elucidate the aspects of canopy structure that favor suspension suggests the need for additional kinds of observational data, focusing on the "immediate structural context" of positional events.     

Substrate alters forelimb to hindlimb peak force ratios in primates

It is often claimed that the walking gaits of primates are unusual because, unlike most other mammals, primates appear to have higher vertical peak ground reaction forces on their hindlimbs than on their forelimbs. Many researchers have argued that this pattern of ground reaction force distribution is part of a general adaptation to arboreal locomotion. This argument is frequently used to support models of primate locomotor evolution. Unfortunately, little is known about the force distribution patterns of primates walking on arboreal supports, nor do we completely understand the mechanisms that regulate weight distribution in primates. We collected vertical peak force data for seven species of primates walking quadrupedally on instrumented terrestrial and arboreal supports. Our results show that, when walking on arboreal vs. terrestrial substrates, primates generally have lower vertical peak forces on both limbs but the difference is most extreme for the forelimb. We found that force reduction occurs primarily by decreasing forelimb and, to a lesser extent, hindlimb stiffness. As a result, on arboreal supports, primates experience significantly greater functional differentiation of the forelimb and hindlimb than on the ground. These data support long-standing theories that arboreal locomotion was a critical factor in the differentiation of the forelimbs and hindlimbs in primates. This change in functional role of the forelimb may have played a critical role in the origin of primates and facilitated the evolution of more specialized locomotor behaviors.     

Limb kinematics during locomotion in the two-toed sloth (Choloepus didactylus,Xenarthra) and its implications for the evolution of the sloth locomotor apparatus

In order to gain insight into the function of the extant sloth locomotion and its evolution, we conducted a detailed videoradiographic analysis of two-toed sloth locomotion (Xenarthra: Choloepus didactylus). Both unrestrained as well as steady-state locomotion was analyzed. Spatio-temporal gait parameters, data on interlimb coordination, and limb kinematics are reported. Two-toed sloths displayed great variability in spatio-temporal gait parameters over the observed range of speeds. They increase speed by decreasing the durations of contact and swing phases, as well as by increasing step length. Gait utilization also varies with no strict gait sequence or interlimb timing evident in slow movements, but a tendency to employ diagonal sequence, diagonal couplet gaits in fast movements. In contrast, limb kinematics were highly conserved with respect to ‘normal’ pronograde locomotion. Limb element and joint angles at touch down and lift off, element and joint excursions, and contribution to body progression of individual elements are similar to those reported for non-cursorial mammals of small to medium size. Hands and feet are specialized to maintain firm connection to supports, and do not contribute to step length or progression. In so doing, the tarsometatarsus lost its role as an individual propulsive element during the evolution of suspensory locomotion. Conservative kinematic behavior of the remaining limb elements does not preclude that muscle recruitment and neuromuscular control for limb pro- and retraction are also conserved. The observed kinematic patterns of two-toed sloths improve our understanding of the convergent evolution of quadrupedal suspensory posture and locomotion in the two extant sloth lineages.     

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.     

Reflex component of the swing phase: A comparative analysis of rat locomotion before and after deafferentation

A comparative analysis was made of the kinematics of movement and EMG activity during different types of locomotion before and after bilateral deafferentation of segments L₁-S₂ of the rat spinal cord. It was found that deafferentation is accompanied by a reduction in the amplitude of locomotor movements and by a delay in both the initiation and increase in duration of flexion in the knee and ankle joints during the swing phase, without producing much effect on the time course of hip joint flexion. An increase in the F period of the swing phase, at its lowest during swimming and highest during stepping, was also discovered, which accordingly rose in step with increasingly deficient afferent inflow. Flexor activity rose especially noticeably during dragging on the limb in the swing phase post-deafferentation. The role of peripheral afferent influence in shaping the F (swing) phase is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 19, No. 5, pp. 654–659, September–October, 1987.     

Locomotion patterns in two South American gymnophthalmid lizards: Vanzosaura rubricauda and Procellosaurinus tetradactylus

We quantified gait and stride characteristics (velocity, frequency, stride length, stance and swing duration, and duty factor) in the bursts of locomotion of two small, intermittently moving, closely related South American gymnophthalmid lizards: Vanzosaura rubricauda and Procellosaurinus tetradactylus. They occur in different environments: V. rubricauda is widely distributed in open areas with various habitats and substrates, while P. tetradactylus is endemic to dunes in the semi-arid Brazilian Caatinga. Both use trot or walking trot characterised by a lateral sequence. For various substrates in a gradient of roughness (perspex, cardboard, sand, gravel), both species have low relative velocities in comparison with those reported for larger continuously moving lizards. To generate velocity, these animals increase stride frequency but decrease relative stride length. For these parameters, P. tetradactylus showed lower values than V. rubricauda. In their relative range of velocities, no significant differences in stride length and frequency were recorded for gravel. However, the slopes of a correlation between velocity and its components were lower in P. tetradactylus on cardboard, whereas on sand this was only observed for velocity and stride length. The data showed that the difference in rhythmic parameters between both species increased with the smoothness of the substrates. Moreover, P. tetradactylus shows a highly specialised locomotor strategy involving lower stride length and frequency for generating lower velocities than in V. rubricauda. This suggests the evolution of a central motor pattern generator to control slower limb movements and to produce fewer and longer pauses in intermittent locomotion.     

Trip recovery strategies following perturbations of variable duration

Appropriately responding to mechanical perturbations during gait is critical to maintain balance and avoid falls. Tripping perturbation onset during swing phase is strongly related to the use of different recovery strategies; however, it is insufficient to fully explain how strategies are chosen. The dynamic interactions between the foot and the obstacle may further explain observed recovery strategies but the relationship between such contextual elements and strategy selection has not been explored. In this study, we investigated whether perturbation onset, duration and side could explain strategy selection for all of swing phase. We hypothesized that perturbations of longer duration would elicit lowering and delayed-lowering strategies earlier in swing phase than shorter perturbations. We developed a custom device to trip subjects multiple times while they walked on a treadmill. Seven young, healthy subjects were tripped on the left or right side at 10% to 80% of swing phase for 150 ms, 250 ms or 350 ms. Strategies were characterized by foot motion post-perturbation and identified by an automated algorithm. A multinomial logistic model was used to investigate the effect of perturbation onset, side, and the interaction between duration and onset on recovery strategy selection. Side perturbed did not affect strategy selection. Perturbation duration interacted with onset, limiting the use of elevating strategies to earlier in swing phase with longer perturbations. The choice between delayed-lowering and lowering strategies was not affected by perturbation duration. Although these variables did not fully explain strategy selection, they improved the prediction of strategy used in response to tripping perturbations throughout swing phase.     

The effect of branch diameter on primate gait sequence pattern

Most mammals use lateral sequence gaits during quadrupedal locomotion, a pattern characterized by the touchdown of a forelimb directly following the ipsilateral hind limb in a given stride cycle. Primates, however, tend to use diagonal sequence (DS) gaits, whereby it is the touchdown of a contralateral forelimb that follows that of a given hind limb most closely in time. A number of scenarios have been offered to explain why primates favor DS gaits, most of them relating to the use of the arboreal habitat and, in particular, the exploitation of a narrow branch niche. This experimental study explores the potential explanation for the use of DS gaits by examining the relationship between branch diameter and gait patterns in 360 strides collected from six species of quadrupedal strepsirrhine primates on broad and narrow diameter supports. Gait sequence is quantified using limb phase, or the percentage of time during a stride cycle that a forelimb touchdown follows an ipsilateral hind limb touchdown. Although Loris, Nycticebus and Eulemur rubriventer individuals in this study did exhibit significantly lower locomotor velocities on narrower supports (P<0.01 in all three species), analyses of covariance showed no significant differences in limb phase values between broad and narrow diameter supports. Hence, results indicate surprisingly little evidence to suggest that alterations in gait sequence pattern provide a specific advantage for negotiating narrow supports.     

Serratus ventralis function in vervet monkeys (Cercopithecus aethiops): are primate quadrupeds unique?

The serratus ventralis in mammals is a fan-shaped scapulo-thoracic muscle that is believed by most morphologists both to support body weight and to rotate the scapula during quadrupedal locomotion. Electromyographic studies of this muscle in cats, dogs and opossums confirm the dual supportive and rotatory roles of the serratus ventralis. Although this muscle has been studied in several primate species, the concentration on arboreal locomotion has resulted in an inadequate data set to permit direct comparisons to non-primate terrestrial quadrupeds. In order to provide a more comparable data set, we examined cranial, mid- and caudal thoracic regions of the serratus ventralis during terrestrial quadrupedalism in the vervet monkey, Cereopithecus aethiops. Our results indicate that the serratus ventralis does support the body during the stance phase of quadrupedalism in this primate. However, unlike several non-primate mammals, it plays a relatively insignificant rotatory role during swing phase.     

How locomotion sub-functions can control walking at different speeds?

Inspired from template models explaining biological locomotory systems and Raibert׳s pioneering legged robots, locomotion can be realized by basic sub-functions: elastic axial leg function, leg swinging and balancing. Combinations of these three can generate different gaits with diverse properties. In this paper we investigate how locomotion sub-functions contribute to stabilize walking at different speeds. Based on this trilogy, we introduce a conceptual model to quantify human locomotion sub-functions in walking. This model can produce stable walking and also predict human locomotion sub-function control during swing phase of walking. Analyzing experimental data based on this modeling shows different control strategies which are employed to increase speed from slow to moderate and moderate to fast gaits.     

Modulation of limb dynamics in the swing phase of locomotion

A method was presented for quantifying cat (Felis catus) hind limb dynamics during swing phase of locomotion using a two-link rigid body model of leg and paw, which highlighted the dynamic interactions between segments. Comprehensive determination was made of cat segment parameters necessary for dynamic analysis, and regression equations were formulated to predict the inertial parameters of any comparable cat. Modulations in muscle and non-muscle components of knee and ankle joint moments were examined at two treadmill speeds using three gaits: (a) pace-like walk and trot-like walk, at 1.0 ms-1, and (b) gallop, at 2.1 ms-1. Results showed that muscle and segment interactive moments significantly effected limb trajectories during swing. Some moment components were greater in galloping than in walking, but net joint maxima were not significantly different between speeds. Moment magnitudes typically were greater for pace-like walking than for trot-like walking at the same speed. Generally, across gaits, the net and muscle moments were in phase with the direction of distal joint motion, and these same moments were out of phase with proximal joint motion. Intersegmental dynamics were not modulated exclusively by speed of locomotion, but interactive moments were also influenced significantly by gait mode.     

Interactive effects of leg autotomy and incline on locomotor performance and kinematics of the cellar spider,Pholcus manueli

Leg autotomy can be a very effective strategy for escaping a predation attempt in many animals. In spiders, autotomy can be very common (5–40% of individuals can be missing legs) and has been shown to reduce locomotor speeds, which, in turn, can reduce the ability to find food, mates, and suitable habitat. Previous work on spiders has focused mostly on the influence of limb loss on horizontal movements. However, limb loss can have differential effects on locomotion on the nonhorizontal substrates often utilized by many species of spiders. We examined the effects of leg autotomy on maximal speed and kinematics while moving on horizontal, 45° inclines, and vertical (90°) inclines in the cellar spider Pholcus manueli, a widespread species that is a denizen of both natural and anthropogenic, three‐dimensional microhabitats, which frequently exhibits autotomy in nature. Maximal speeds and kinematic variables were measured in all spiders, which were run on all three experimental inclines twice. First, all spiders were run at all inclines prior to autotomization. Second, half of the spiders had one of the front legs removed, while the other half was left intact before all individuals were run a second time on all inclines. Speeds decreased with increasing incline and following autotomy at all inclines. Autotomized spiders exhibited a larger decrease in speed when moving horizontally compared to on inclines. Stride length decreased at 90° but not after autotomy. Stride cycle time and duty factor increased after autotomy, but not when moving uphill. Results show that both incline and leg autotomy reduce speed with differential effects on kinematics with increasing incline reducing stride length, but not stride cycle time or duty factor, and vice versa for leg autotomy. The lack of a significant influence on a kinematic variable could be evidence for partial compensation to mitigate speed reduction.     

In vivo bone strain on the dog tibia during locomotion

Rosette and single-element strain gauges were implanted on the tibia in 2 dogs and recordings were made during locomotion on a treadmill. At foot contact and during the swing phase of locomotion, bone strains were low and directions of the principal strains were variable. There was a large shift in the directions of the principal strains at the beginning of the stance phase and bone strains were considerably higher. Peak strain occurred midway through the stance phase. At that time, the maximum principal strain (tension) was directed upwards and anteriorly between 30 and 60 degrees with respect to the long axis of the tibia. These bone strain patterns in the dog are similar to those found in sheep while both differ markedly from those found in humans.     

Locomotor behavior of wild orangutans (Pongo pygmaeus wurmbii) in disturbed peat swamp forest, Sabangau, Central Kalimantan, Indonesia

This study examined the locomotor behavior of wild Bornean orangutans (P. p. wurmbii) in an area of disturbed peat swamp forest (Sabangau Catchment, Indonesia) in relation to the height in the canopy, age-sex class, behavior (feeding or traveling), and the number of supports used to bear body mass. Backward elimination log-linear modeling was employed to expose the main influences on orangutan locomotion. Our results showed that the most important distinctions with regard to locomotion were between suspensory and compressive, or, orthograde (vertical trunk) and pronograde (horizontal trunk) behavior. Whether orangutans were traveling or feeding had the most important influence on locomotion whereby compressive locomotion had a strong association with feeding, suspensory locomotion had a strong association with travel in the peripheral strata using multiple supports, whereas vertical climb/descent and oscillation showed a strong association with travel on single supports in the core stratum. In contrast to theoretical predictions on positional behavior and body size, age-sex category had a limited influence on locomotion. The study revealed that torso orthograde suspension dominates orangutan locomotion, concurring with previous studies in dipterocarp forest. But, orangutans in the Sabangau exhibited substantially higher frequencies of oscillatory locomotion than observed at other sites, suggesting this behavior confers particular benefits for traversing the highly compliant arboreal environment typical of disturbed peat swamp forest. In addition, torso pronograde suspensory locomotion was observed at much lower levels than in the Sumatran species. Together these results highlight the necessity for further examination of differences between species, which control for habitat.     

Kinetic gait analysis in English Bulldogs

Background

Canine breed conformation may interfere with locomotion and may predispose to orthopedic disease. Bulldogs have a high incidence of orthopedic diseases such as hip dysplasia. Kinetic gait analysis provides an objective way to assess and analyze locomotion. The aim of this study was to study the vertical forces of English Bulldogs during walk using a pressure sensitive walkway. We hypothesize that Bulldogs affected by orthopedic diseases have decreased weight bearing and asymmetric locomotion in the limbs despite having mild to no sings during clinical examination. Thirty English Bulldogs were tested. Peak vertical force, vertical impulse, rate of loading, stance phase duration, symmetry index, goniometry and incidence of orthopedic diseases were recorded.

Results

Although none of the dogs showed signs of pain or discomfort upon manipulation of the hip joints, all dogs had radiographic evidences of hip dysplasia and lack of significant peak vertical force, vertical impulse and stance time differences. The dogs had a mean hind limb symmetry index of 19.8 ± 19.5% and rates of loading ranged from 1.0 to 3.1.

Conclusions

Despite the lack of evident decrease in weight bearing, subclinical lameness can be inferred. The examined dogs had a mean hind limb symmetry index of 19.8 ± 19.5%. Symmetry indices reported in dogs free from orthopedic diseases range from 0.3 to 9.6%. Given non-lame dogs are expected to have a symmetry index close to 0%, data from this study suggests that Bulldogs have gait dysfunctions, which translates into hind limb asymmetries and rate of loading was consistent with severe hip dysplasia despite no visible signs of gait dysfunction. Future studies comparing lame and non-lame Bulldogs are warranted.

     

Optimum walking techniques for idealized animals

The vertical component of the force exerted by a foot on the ground, in the course of a step, may rise to a single maximum and decline again (as in human running) or may show two distinct maxima (as in human walking). A foot may remain on the ground for a large or small fraction of the duration of a stride. Mathematical models are used to investigate the effects of these differences of technique on the energy cost of locomotion. The optimum technique for a biped at a given speed is different from the optimum for a hypothetical many-legged animal. The optima for quadrupedal walking are likely to lie between these extremes.
The walking techniques adopted by men at different speeds are close to the optima indicated by the bipedal model. The two maxima of the force exerted by a foot are higher, and have a lower minimum between them, at higher speeds of walking. The techniques adopted by a sheep are close to the optima indicated by the many-legged model but dogs use techniques rather closer to the optima for bipeds.
The limitations of the models are discussed.     

Locomotor energetics in primates: gait mechanics and their relationship to the energetics of vertical and horizontal locomotion

All primates regularly move within three-dimensional arboreal environments and must often climb, but little is known about the energetic costs of this critical activity. Limited previous work on the energetics of incline locomotion suggests that there may be differential selective pressures for large compared to small primates in choosing to exploit a complex arboreal environment. Necessary metabolic and gait data have never been collected to examine this possibility and biomechanical mechanisms that might explain size-based differences in the cost of arboreal movement. Energetics and kinematics were collected for five species of primate during climbing and horizontal locomotion. Subjects moved on a treadmill with a narrow vertical substrate and one with a narrow horizontal substrate at their maximum sustainable speed for 10–20 min while oxygen consumption was monitored. Data during climbing were compared to those during horizontal locomotion and across size. Results show that climbing energetic costs were similar to horizontal costs for small primates (<0.5 kg) but were nearly double for larger species. Spatio-temporal gait characteristics suggest that the relationship between the cost of locomotion and the rate of force production changes between the two locomotor modes. Thus, the main determinants of climbing costs are fundamentally different from those during horizontal locomotion. These new results combining spatiotemporal and energetic data confirm and expand on our previous argument (Hanna et al.: Science 320 (2008) 898) that similar costs of horizontal and vertical locomotion in small primates facilitated the successful occupation of a fine-branch arboreal milieu by the earliest primates.     

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.