Andrew A. Biewener

Bio: Andrew A. Biewener is an academic researcher from Harvard University. The author has contributed to research in topics: Isometric exercise & Motor unit recruitment. The author has an hindex of 71, co-authored 210 publications receiving 14592 citations. Previous affiliations of Andrew A. Biewener include Field Museum of Natural History & University of Bristol.

Scaling body support in mammals: limb posture and muscle mechanics.

Abstract: The scaling of bone and muscle geometry in mammals suggests that peak stresses (ratio of force to cross-sectional area) acting in these two support elements increase with increasing body size. Observations of stresses acting in the limb bones of different sized mammals during strenuous activity, however, indicate that peak bone stress is independent of size (maintaining a safety factor of between 2 and 4). It appears that similar peak bone stresses and muscle stresses in large and small mammals are achieved primarily by a size-dependent change in locomotor limb posture: small animals run with crouched postures, whereas larger species run more upright. By adopting an upright posture, large animals align their limbs more closely with the ground reaction force, substantially reducing the forces that their muscles must exert (proportional to body mass) and hence, the forces that their bones must resist, to counteract joint moments. This change in limb posture to maintain locomotor stresses within safe limits, however, likely limits the maneuverability and accelerative capability of large animals.

Biomechanics of mammalian terrestrial locomotion.

Abstract: Mammalian skeletons experience peak locomotor stresses (force per area) that are 25 to 50% of their failure strength, indicating a safety factor of between two and four. The mechanism by which animals achieve a constant safety factor varies depending on the size of the animal. Over much of their size (0.1 to 300 kilograms), larger mammals maintain uniform skeletal stress primarily by having a more upright posture, which decreases mass-specific muscle force by increasing muscle mechanical advantage. At greater sizes, increased skeletal allometry and decreased locomotor performance likely maintain stresses constant. At smaller sizes, skeletal stiffness may be more critical than strength. The decrease in mass-specific muscle force in mammals weighing 0.1 to 300 kilogram indicates that peak muscle stresses are also constant and correlates with a decrease in mass-specific energy cost of locomotion. The consistent pattern of locomotor stresses developed in long bones at different speeds and gaits within a species may have important implications for how bones adaptively remodel to changes in stress.

Bipedal locomotion: effects of speed, size and limb posture in birds and humans

Abstract: Seven species of ground-dwelling birds (body mass range: 0.045-90 kg) were filmed while walking and running on a treadmill. High-speed light films were also taken of humans to compare kinematic patterns of avian with human bipedalism. Consistent patterns of stride frequency, stride length, step length, duty factor and limb excursion were observed in all species, with most of the variation among species being due to differences in body size. In general, smaller bipeds have higher stride frequencies (αM−0.18), shorter stride lengths (αM0.38) and more limited ranges of speed within each gait than large bipeds. After normalizing for size (based on Froude number, after Alexander, 1977), remaining kinematic variation is largely due to interspecific differences in posture and relative limb segment lengths. For their size, smaller bipeds have greater step lengths, limb excursion angles and duty factors than large bipeds because of their more crouched posture and greater effective limb length. The most notable differences in limb kinematics between birds and humans occur at the walk-run transition and are maintained as running speed increases. Change of gait is smooth and difficult to discern in birds, but distinct in humans, involving abrupt decreases in step length and duty factor (time of contact) and a corresponding increase in limb swing time. These differences appear to reflect a spring-like run that is stiff in humans (favouring elastic energy recovery) but more compliant in birds (increasing time of ground contact). Differences between birds and humans in balance of the body's centre of mass not only affect femoral orientation and motion, but also affect pattern of limb excursion with speed.

Allometry of quadrupedal locomotion: the scaling of duty factor, bone curvature and limb orientation to body size

Abstract: Measurements of the chord length (alpha M0.31) and diameter (alpha M0.35) of the femora, tibiae, humeri and radii from 32 species of mammals, ranging in approximate body mass from 0.020-3500 kg, support previous data which show that mammalian long bones scale close to geometric similarity. Scaling of peak stresses based on these measurements of limb bone geometry predicts that peak stress increases alpha M0.28, assuming that the forces acting on a bone are directly proportional to an animal's weight. Peak locomotory stresses measured in small and large quadrupeds contradict this scaling prediction, however, showing that the magnitude of peak bone stress is similar over a range of size. Consequently, a uniform safety factor is maintained. Bone curvature (alpha M-0.09) and limb bone angle relative to the direction of ground force (alpha M-0.07) exhibit a slight, but significant, decrease with increasing body mass. Duty factor measured at the animal's trot--gallop transition speed does not change significantly with body size. The moment arm ratio of ground force to muscular force exerted about a joint was found to decrease dramatically for horses as compared to ground squirrels and chipmunks. This six-fold decrease (alpha M-0.23) provides preliminary data which appear to explain, along with the decrease in bone curvature and angle, the similar magnitudes of peak bone stress developed during locomotion in different sized animals. The crouched posture adopted by small quadrupeds while running may allow greater changes in momentum (when accelerating or decelerating) or a decrease in the forces exerted on their limbs.

Energetics and mechanics of human running on surfaces of different stiffnesses

Abstract: Mammals use the elastic components in their legs (principally tendons, ligaments, and muscles) to run economically, while maintaining consistent support mechanics across various surfaces. To examine how leg stiffness and metabolic cost are affected by changes in substrate stiffness, we built experimental platforms with adjustable stiffness to fit on a force-plate-fitted treadmill. Eight male subjects [mean body mass: 74.4 +/- 7.1 (SD) kg; leg length: 0.96 +/- 0.05 m] ran at 3.7 m/s over five different surface stiffnesses (75.4, 97.5, 216.8, 454.2, and 945.7 kN/m). Metabolic, ground-reaction force, and kinematic data were collected. The 12.5-fold decrease in surface stiffness resulted in a 12% decrease in the runner's metabolic rate and a 29% increase in their leg stiffness. The runner's support mechanics remained essentially unchanged. These results indicate that surface stiffness affects running economy without affecting running support mechanics. We postulate that an increased energy rebound from the compliant surfaces studied contributes to the enhanced running economy.

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How Animals Move: An Integrative View

Abstract: Recent advances in integrative studies of locomotion have revealed several general principles. Energy storage and exchange mechanisms discovered in walking and running bipeds apply to multilegged locomotion and even to flying and swimming. Nonpropulsive lateral forces can be sizable, but they may benefit stability, maneuverability, or other criteria that become apparent in natural environments. Locomotor control systems combine rapid mechanical preflexes with multimodal sensory feedback and feedforward commands. Muscles have a surprising variety of functions in locomotion, serving as motors, brakes, springs, and struts. Integrative approaches reveal not only how each component within a locomotor system operates but how they function as a collective whole.