The gaits of five Ostriches Struthio camelus, seven Emus Dromaius novaehollandiae, two Greater Rheas Rhea americana, two Southern Cassowaries Casuarius casuarius and one Brown h w i Apteryx australis were filmed a t zoological parks. Locomotor parameters were measured using footprints on sandy tracks and video records. Osteological measurements
SUMMARYOnly a few studies on quadrupedal locomotion have investigated symmetrical and asymmetrical gaits in the same framework because the mechanisms underlying these two types of gait seem to be different and it took a long time to identify a common set of parameters for their simultaneous study. Moreover, despite the clear importance of the spatial dimension in animal locomotion, the relationship between temporal and spatial limb coordination has never been quantified before. We used anteroposterior sequence (APS) analysis to analyse 486 sequences from five malinois (Belgian shepherd) dogs moving at a large range of speeds (from 0.4 to 10.0·m·s -1 ) to compare symmetrical and asymmetrical gaits through kinematic and limb coordination parameters. Considerable continuity was observed in cycle characteristics, from walk to rotary gallop, but at very high speeds an increase in swing duration reflected the use of sagittal flexibility of the vertebral axis to increase speed. This change occurred after the contribution of the increase in stride length had become the main element driving the increase in speed -i.e. when the dogs had adopted asymmetrical gaits. As the left and right limbs of a pair are linked to the same rigid structure, spatial coordination within pairs of limbs reflected the temporal coordination within pairs of limbs whatever the speed. By contrast, the relationship between the temporal and spatial coordination between pairs of limb was found to depend on speed and trunk length. For trot and rotary gallop, this relationship was thought also to depend on the additional action of trunk flexion and leg angle at footfall.
SUMMARYTake-off mechanics are fundamental to the ecology and evolution of flying animals. Recent research has revealed that initial takeoff velocity in birds is driven mostly by hindlimb forces. However, the contribution of the wings during the transition to air is unknown. To investigate this transition, we integrated measurements of both leg and wing forces during take-off and the first three wingbeats in zebra finch (Taeniopygia guttata, body mass 15g, N7) and diamond dove (Geopelia cuneata, body mass 50g, N3). We measured ground reaction forces produced by the hindlimbs using a perch mounted on a force plate, whole-body and wing kinematics using high-speed video, and aerodynamic forces using particle image velocimetry (PIV). Take-off performance was generally similar between species. When birds were perched, an acceleration peak produced by the legs contributed to 85±1% of the whole-body resultant acceleration in finch and 77±6% in dove. At lift-off, coincident with the start of the first downstroke, the percentage of hindlimb contribution to initial flight velocity was 93.6±0.6% in finch and 95.2±0.4% in dove. In finch, the first wingbeat produced 57.9±3.4% of the lift created during subsequent wingbeats compared with 62.5±2.2% in dove. Advance ratios were <0.5 in both species, even when taking self-convection of shed vortices into account, so it was likely that wing-wake interactions dominated aerodynamics during wingbeats 2 and 3. These results underscore the relatively low contribution of the wings to initial take-off, and reveal a novel transitional role for the first wingbeat in terms of force production. Supplementary material available online at
The locomotion of early tetrapods has long been a subject of great interest in the evolutionary history of vertebrates. However, we still do not have a precise understanding of the evolutionary radiation of their locomotory strategies. We present here the first palaeohistological study based on theoretical biomechanical considerations among a highly diversified group of early tetrapods, the temnospondyls. Based on the quantification of microanatomical and histological parameters in the humerus and femur of nine genera, this multivariate analysis provides new insights concerning the adaptations of temnospondyls to their palaeoenvironments during the Early Permian, and clearly after the Permo‐Triassic crisis. This study therefore presents a methodology that, if based on a bigger sample, could contribute towards a characterization of the behaviour of species during great evolutionary events.
Most birds use at least two modes of locomotion: flying and walking (terrestrial locomotion). Whereas the wings and tail are used for flying, the legs are mainly used for walking. The role of other body segments remains, however, poorly understood. In this study, we examine the kinematics of the head, the trunk, and the legs during terrestrial locomotion in the quail (Coturnix coturnix). Despite the trunk representing about 70% of the total body mass, its function in locomotion has received little scientific interest to date. This prompted us to focus on its role in terrestrial locomotion. We used high-speed video fluoroscopic recordings of quails walking at voluntary speeds on a trackway. Dorso-ventral and lateral views of the motion of the skeletal elements were recorded successively and reconstructed in three dimensions using a novel method based on the temporal synchronisation of both views. An analysis of the trajectories of the body parts and their coordination showed that the trunk plays an important role during walking. Moreover, two sub-systems participate in the gait kinematics: (i) the integrated 3D motion of the trunk and thighs allows for the adjustment of the path of the centre of mass; (ii) the motion of distal limbs transforms the alternating forward motion of the feet into a continuous forward motion at the knee and thus assures propulsion. Finally, head bobbing appears qualitatively synchronised to the movements of the trunk. An important role for the thigh muscles in generating the 3D motion of the trunk is suggested by an analysis of the pelvic anatomy.
Transitions to and from the air are critical for aerial locomotion and likely shaped the evolution of flying animals. Research on take-off demonstrates that legs generate greater body accelerations compared with wings, and thereby contribute more to initial flight velocity. Here, we explored coordination between wings and legs in two species with different wingbeat styles, and quantified force production of these modules during the final phase of landing. We used the same birds that we had previously studied during take-off: zebra finch (Taeniopygia guttata, N=4) and diamond dove (Geopelia cuneata, N=3). We measured kinematics using high-speed video, aerodynamics using particle image velocimetry, and ground-reaction forces using a perch mounted on a force plate. In contrast with the first three wingbeats of take-off, the final four wingbeats during landing featured ~2 times greater force production. Thus, wings contribute proportionally more to changes in velocity during the last phase of landing compared with the initial phase of take-off. The two species touched down at the same velocity (~1 m s −1 ), but they exhibited significant differences in the timing of their final wingbeat relative to touchdown. The ratio of average wing force to peak leg force was greater in diamond doves than in zebra finches. Peak ground reaction forces during landing were ~50% of those during take-off, consistent with the birds being motivated to control landing. Likewise, estimations of mechanical energy flux for both species indicate that wings produce 3-10 times more mechanical work within the final wingbeats of flight compared with the kinetic energy of the body absorbed by legs during ground contact.KEY WORDS: Zebra finch, Taeniopygia guttata, Diamond dove, Geopelia cuneata, Locomotion, Hindlimb, Forelimb, Particle image velocimetry, PIV INTRODUCTIONLanding allows the transition from the air to the ground or other substrate. This phase is fundamental to flight, but is often overlooked in studies compared with take-off or cruising flight. Landing places a unique selective pressure upon aerial animals, because a controlled deceleration and descent must be accomplished to avoid injury (Bonser, 1999;Paskins et al., 2007). Animals that do not decelerate properly risk broken wings or legs, or damaged integument: collisions often lead to mortality (Klem, 1990 birds, and even the engineering of autonomous flying vehicles. Herein, we explored the coordination of force production between both legs and wings during landing in birds.Several lines of evidence indicate that birds carefully control their landing and generally seek to minimize their kinetic energy at touchdown (Warrick et al., 2002) even if some species are eminently capable of targeting in fast flight [for example, a peregrine falcon stooping on a pigeon at relatively fast speed (Alerstam, 1987)]. During landing, birds modulate both wing and tail kinematics to decelerate prior to arriving at the perch (Berg and Biewener, 2010). The distal muscles of the wing, associ...
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