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Showing papers by "Andrew A. Biewener published in 2004"


Journal ArticleDOI
TL;DR: It is concluded that the decrease in limb mechanical advantage (mean limb extensor EMA) and increase in knee extensor impulse during running likely contribute to the higher metabolic cost of transport in running than in walking.
Abstract: Muscular forces generated during locomotion depend on an animal's speed, gait, and size and underlie the energy demand to power locomotion. Changes in limb posture affect muscle forces by altering ...

286 citations


Journal ArticleDOI
TL;DR: Investigating the variation in instantaneous aerodynamic force production during the wingbeat cycle of birds flying across a range of steady speeds found that the inertial power requirements of downstroke were consistent with the assumption that nearly all wing kinetic energy in downstroke was applied to the production of aerodynamic forces and therefore should not be added separately to the overall power cost of flight.
Abstract: We used a combination of high-speed 3-D kinematics and three-axis accelerometer recordings obtained from cockatiels flying in a low-turbulence wind tunnel to characterize the instantaneous accelerations and, by extension, the net aerodynamic forces produced throughout the wingbeat cycle across a broad range of flight speeds (1-13 m s(-1)). Our goals were to investigate the variation in instantaneous aerodynamic force production during the wingbeat cycle of birds flying across a range of steady speeds, testing two predictions regarding aerodynamic force generation in upstroke and the commonly held assumption that all of the kinetic energy imparted to the wings of a bird in flapping flight is recovered as useful aerodynamic work. We found that cockatiels produce only a limited amount of lift during upstroke (14% of downstroke lift) at slower flight speeds (1-3 m s(-1)). Upstroke lift at intermediate flight speeds (7-11 m s(-1)) was moderate, averaging 39% of downstroke lift. Instantaneous aerodynamic forces were greatest near mid-downstroke. At the end of each half-stroke, during wing turnaround, aerodynamic forces were minimal, but inertial forces created by wing motion were large. However, we found that the inertial power requirements of downstroke (minimum of 0.29+/-0.10 W at 7 m s(-1) and maximum of 0.56+/-0.13 W at 1 m s(-1)) were consistent with the assumption that nearly all wing kinetic energy in downstroke was applied to the production of aerodynamic forces and therefore should not be added separately to the overall power cost of flight. The inertial power requirements of upstroke (minimum of 0.16+/-0.04 W at 7 m s(-1) and maximum of 0.35+/-0.11 W at 1 m s(-1)) cannot be recovered in a similar manner, but their magnitude was such that the power requirements for the upstroke musculature (minimum of 54+/-13 W kg(-1) at 7 m s(-1) and maximum of 122+/-35 W at 1 m s(-1)) fall within the established range for cockatiel flight muscle (<185 W kg(-1)).

121 citations


Journal ArticleDOI
TL;DR: Kassina maculata (red-legged running frog) primarily uses a third type of locomotion during which the hindlimbs alternate, and its distinct locomotory mode is examined to determine whether these frogs walk or run and how their gait may change with speed.
Abstract: SUMMARY Although most frog species are specialized for jumping or swimming, Kassina maculata (red-legged running frog) primarily uses a third type of locomotion during which the hindlimbs alternate. In the present study, we examined Kassina 9s distinct locomotory mode to determine whether these frogs walk or run and how their gait may change with speed. We used multiple methods to distinguish between terrestrial gaits: the existence or absence of an aerial phase, duty factor, relative footfall patterns and the mechanics of the animal9s center of mass (COM). To measure kinematic and kinetic variables, we recorded digital video as the animals moved over a miniature force platform ( N =12 individuals). With respect to footfall patterns, the frogs used a single gait and walked at all speeds examined. Duty factor always exceeded 0.59. Based on COM mechanics, however, the frogs used both walking and running gaits. At slower speeds, the fluctuations in the horizontal kinetic energy ( E k ) and gravitational potential energy ( E p ) of the COM were largely out of phase, indicating a vaulting or walking gait. In most of the trials, Kassina used a combined gait at intermediate speeds, unlike cursorial animals with distinct gait transitions. This combined gait, much like a mammalian gallop, exhibited the mechanics of both vaulting and bouncing gaits. At faster speeds, the E k and E p of Kassina 9s COM were more in phase, indicating the use of a bouncing or running gait. Depending on the definition used to distinguish between walking and running, Kassina either only used a walking gait at all speeds or used a walking gait at slower speeds but then switched to a running gait as speed increased.

115 citations


Journal ArticleDOI
TL;DR: The results indicate that design for spring-like tendon energy savings and economical muscle force generation is key for these two distal muscle-tendon units of the tammar wallaby, and the need to accommodate changes in work associated with level versus incline locomotion is achieved by more proximal muscles of the limb.
Abstract: The goal of our study was to examine whether the in vivo force-length behavior, work and elastic energy savings of distal muscle-tendon units in the legs of tammar wallabies (Macropus eugenii) change during level versus incline hopping. To address this question, we obtained measurements of muscle activation (via electromyography), fascicle strain (via sonomicrometry) and muscle-tendon force (via tendon buckles) from the lateral gastrocnemius (LG) and plantaris (PL) muscles of tammar wallabies trained to hop on a level and an inclined (10°, 17.4% grade) treadmill at two speeds (3.3 m s^(-1) and 4.2 m s^(-1)). Similar patterns of muscle activation, force and fascicle strain were observed under both level and incline conditions. This also corresponded to similar patterns of limb timing and movement (duty factor, limb contact time and hopping frequency). During both level and incline hopping, the LG and PL exhibited patterns of fascicle stretch and shortening that yielded low levels of net fascicle strain [LG: level, -1.0±4.6% (mean ± s.e.m.) vs incline, 0.6±4.5%; PL: level, 0.1±1.0% vs incline, 0.4±1.6%] and muscle work (LG: level, -8.4±8.4 J kg^(-1) muscle vs incline, -6.8±7.5 J kg^(-1) muscle; PL: level, -2.0±0.6 J kg^(-1) muscle vs incline, -1.4±0.7 J kg^(-1) muscle). Consequently, neither muscle significantly altered its contractile dynamics to do more work during incline hopping. Whereas electromyographic (EMG) phase, duration and intensity did not differ for the LG, the PL exhibited shorter but more intense periods of activation, together with reduced EMG phase (P<0.01), during incline versus level hopping. Our results indicate that design for spring-like tendon energy savings and economical muscle force generation is key for these two distal muscle-tendon units of the tammar wallaby, and the need to accommodate changes in work associated with level versus incline locomotion is achieved by more proximal muscles of the limb.

86 citations


Journal ArticleDOI
TL;DR: Even though bone strains in the younger goats were small enough to suggest that they maintain safety factors at least comparable with adults when moving at similar absolute speeds, increased variability of loading in juvenile animals may also favor the more robust dimensions of the radius, and possibly other limb bones, early in growth.
Abstract: As tetrapods increase in size and weight through ontogeny, the limb skeleton must grow to accommodate the increases in body weight and the resulting locomotor forces placed upon the limbs. No study to date, however, has examined how morphological changes in the limb skeleton during growth reflect ontogenetic patterns of limb loading and the resulting stresses and strains produced in the limbs. The goal of this study was to relate forelimb loads to in vivo bone strains in the radius of the domestic goat (Capra hircus) across a range of gaits and speeds through ontogeny while observing how the growth patterns of the bone relate to the mechanics of the limb. In vivo bone strains in the radius were recorded from two groups of juvenile goats (4 kg, 6 weeks and 9 kg, 15 weeks) and compared with previously reported strain data for the radius of adult goats. Ontogenetic strain patterns were examined in relation to peak forelimb ground reaction forces, ontogenetic scaling patterns of cross-sectional geometry and bone curvature, and percentage mineral ash content. Peak principal longitudinal tensile strains on the cranial surface and compressive strains on the caudal surface of the radius increased during ontogeny but maintained a uniform distribution, resulting in the radius being loaded primarily in bending through ontogeny. The increase in strain occurred despite uniform loading (relative to body weight) of the forelimb through ontogeny. Instead, the increase in bone strain resulted from strong negative growth allometry of the cross-sectional area (proportional to M(0.53)) and medio-lateral and cranio-caudal second moments of area (I(ML) proportional to M(1.03), I(CC) proportional to M(0.84)) of the radius and only a small increase (+2.8%) in mineral ash content. Even though bone strains increased with growth and age, strains in the younger goats were small enough to suggest that they maintain safety factors at least comparable with adults when moving at similar absolute speeds. Increased variability of loading in juvenile animals may also favor the more robust dimensions of the radius, and possibly other limb bones, early in growth.

79 citations