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.

Gait parameters of treadmill versus overground locomotion in mouse.

Many studies of interest in motor behaviour and motor impairment in mice use equally treadmill or track as a routine test. However, the literature in mammals shows a wide difference of results between the kinematics of treadmill and overground locomotion. To study these discrepancies, we analyzed the locomotion of adult SWISS-OF1 mice over a large range of velocities using treadmill and overground track. The use of a high-speed video camera combined with cinefluoroscopic equipment allowed us to quantify in detail the various space and time parameters of limb kinematics. The results show that mice maintain the same gait pattern in both conditions. However, they also demonstrate that during treadmill exercise mice always exhibit higher stride frequency and consequently lower stride length. The relationship of the stance time and the swing time against the stride frequency are still the same in both conditions. We conclude that the conflict related to the discrepancy between the proprioceptive, vestibular, and visual inputs contribute to an increase in the stride frequency during the treadmill locomotion.

An alternative approach to normalization and evaluation for gait patterns: Procrustes analysis applied to the cyclograms of sprinters and middle-distance runners.

In order to compare gait patterns, a common procedure is to normalize strides both in time and magnitude. The stride duration is usually normalized to a time percentage before averaging curves. As the timing of event occurrences may shift across strides, the shape of the averaged curves is distorted and therefore the standard deviation is overvalued. Stride magnitude normalization is performed by means of dimensionless numbers. However, there is little agreement on which body size correction methods should be used. The Procrustes method describes curve shape and shape change in a mathematical and statistical framework, independently of time and size factors. The present study aims to explore how this technique may be used for time- and magnitude-stride normalization to reflect individual and group mean responses. The Procrustes method, which combines quantitative and visual features, is applied to the shape of the ankle and knee cyclograms. Superimposition of 25 cyclograms (10 for sprinters (SP) and 15 for middle-distance runners (MDR)) was supplemented by statistical procedures (principal component analysis, discriminant function) to extract the main key events, which vary according to the athletic specialities. In comparison with the MDR (poulaine-shaped cyclogram), the ovoid cyclogram of SP reveals the following gait indicators: a short braking phase, a rapid initial lower limb swing in the forward direction, a fast upward movement of the knee and ankle, and an active foot contact. The Procrustes approach could be used to describe other quasi-periodic movements through relative motion plots (e.g., cyclograms, angle-angle diagrams, phase plane portraits).

Comparative study on the forefoot and hindfoot intrinsic muscles of some cavioidea rodents (Mammalia, Rodentia).

The present study compares the forefoot and hindfoot musculature of five representative species of Cavioidea rodents. In all species, the musculature of both forefeet and hindfeet have the same array regardless of the absence of digit I in the manus of Hydrochaeris hydrochaeris and Cavia porcellus. Our results suggest a tendency in these species towards a three-digit system, with a functional loss of digit V and a predominance of digit III in their forefeet. In the same way, the muscular reduction of digit I in the other rodents analyzed indicates a four-digit system with predominance of digit II in Myoprocta acouchy and Dasyprocta leporina and of digit V in Agouti paca. There seems to be an association between the muscular arrangement and functional axis of the foot, raising the general question why this axis runs between the third and forth digit, or along the third digit.

Contribution to the analysis of gaits: practical elements to complement the Hildebrand method.

This paper suggests three additions to the Hildebrand method for gait-pattern specification. The first allows an extension of this method by the use of the forelimb as reference. Thus, dataset could be gathered indifferently from fore or hindlimbs cycles to identify a symmetrical gait or compare gaits of diverse species. On the basis of Hildebrand's definitions, the second suggestion permits to check the proportion of symmetrical and asymmetrical gaits adopted by different individuals of one or different species. The third addition makes a graphical clear-cut between four basic modes of gallops: rotary, transverse, half-bound and bound. These additions will facilitate extensive data comparison either to evaluate the range of variation within a single species or to specify the preferred gaits in diverse species.

Evidence of force exchanges during the six-legged walking of the bottom-dwelling fish, Chelidonichthys lucerna.

Locomotion in terrestrial vertebrates is supposed to be derived from preadaptation in bottom-dwelling fish. A few fish species have been assumed to walk on the substratum, on the basis of coordinated movements of their paired fins. However, the validity of this assumption has remained uncertain, because of a lack of evidence that their fin rays actually exert a force on the substratum. Here, we provide the first conclusive evidence that a benthic teleost fish, the gurnard, Chelidonichthys lucerna (Triglidae), exerts forces on the substratum during its temporary bottom-dwelling hexapod locomotion. This demonstration was achieved by the use of a photoelastic gel technique combined with a force calibration device. The movement patterns of the three first pairs of rays of the pectoral fins were analysed in relation to the forces exerted on the substratum, by measuring deformations of the photoelastic gel substratum produced by the rays. The rays were shown to produce a force pattern that confirmed the existence of a hexapod locomotion in a vertebrate that was consistent with body propulsion and voluntary substratum walking.

Geometric morphometrics of the shoulder girdle in extant turtles (Chelonii).

The aim of this study was to identify shape patterns of the shoulder girdle in relation to different functional and environmental behaviours in turtles. The Procrustes method was used to compare the shoulder girdles (scapula and coracoid) of 88 adult extant turtles. The results indicate that four shape patterns can be distinguished. The shoulder girdles of (1) terrestrial (Testudinidae), (2) highly aquatic freshwater (Trionychidae, Carettochelyidae) and (3) marine turtles (Cheloniidae, Dermochelyidae) correspond to three specialized morphological patterns, whereas the shoulder girdle of (4) semi-aquatic freshwater turtles (Bataguridae, Chelidae, Chelydridae, Emydidae, Kinosternidae, Pelomedusidae, Platysternidae, Podocnemididae) is more generalized. In terrestrial turtles, the long scapular prong and the short coracoid are associated with a domed shell and a mode of locomotion in which walking is predominant. By contrast, highly aquatic freshwater turtles share traits with marine turtles. In both, the short scapular prong and the long coracoid are associated with a flat shell, and swimming locomotion. The enlarged attachment sites of the biceps, coracobrachialis magnus, and supracoracoideus also give these strong swimmers a mechanical advantage during adduction and retraction of the arm. Increasing size leads to allometrical shape changes that emphasize mechanical efficiency both in terrestrial and in aquatic turtles.

Comments on the pelvic appendicular vestiges in an amphisbaenian: Blanus cinereus (Reptilia, squamata).

A detailed description of muscular and skeletal features of the pelvi-cloacal region of the amphisbaenian Blanus cinereus reveals that the species has real hind limbs articulating with a real pelvic girdle. Arguments support this homology: 1) a link with the vertebral column; 2) a movable articulation, showing all the features of a diarthrosis, between the femur and the acetabulum; 3) all the long bone characteristics for the femur which distally bears a horny element. The morphological peculiarities of the amphisbaenian pelvic girdle are generally close to those of lizards, but the pubis seems to be more reduced. For the same number of precloacal vertebrae, the amphisbaenians have more appendicular elements than lizards have, and this composition recalls that of the Leptotyphlopidae, Aniliidae, and Boidae. The account provides more information concerning the aponeuro-tendinous system associated to the skeleton of the girdle and the hindlimb, the musculature, and the interrelations between the different structures of the pelvi-cloacal region.

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.