It has been argued that minimization of metabolic-energy costs is a primary determinant of gait selection in terrestrial animals. This view is based predominantly on data from humans and horses, which have been shown to choose the most economical gait (walking, running, galloping) for any given speed. It is not certain whether a minimization of metabolic costs is associated with the selection of other prevalent forms of terrestrial gaits, such as grounded running (a widespread gait in birds). Using biomechanical and metabolic measurements of four ostriches moving on a treadmill over a range of speeds from 0.8 to 6.7 m s -1 , we reveal here that the selection of walking or grounded running at intermediate speeds also favours a reduction in the metabolic cost of locomotion. This gait transition is characterized by a shift in locomotor kinetics from an inverted-pendulum gait to a bouncing gait that lacks an aerial phase. By contrast, when the ostrich adopts an aerial-running gait at faster speeds, there are no abrupt transitions in mechanical parameters or in the metabolic cost of locomotion. These data suggest a continuum between grounded and aerial running, indicating that they belong to the same locomotor paradigm.

displacement of the lower limb necessary for steering the swinging limb clear of the stance limb and replaces what would otherwise require greater adduction/abduction and/or internal/external rotation, allowing for less complex joints, musculoskeletal geometry and neuromuscular control. Significant rotation about the joints' nonflexion/extension axes nevertheless occurs over the running stride. In particular, hip abduction and knee internal/external and varus/valgus motion may further facilitate limb clearance during the swing phase, and substantial non-flexion/extension movement at the knee is also observed during stance. Measurement of 3-D segment and joint motion in birds will be aided by the use of functionally determined axes of rotation rather than assumed axes, proving important when interpreting the biomechanics and motor control of avian bipedalism.

The purpose of this study was to examine the mechanical adaptations linked to economical locomotion in cursorial bipeds. We addressed this question by comparing mass-matched humans and avian bipeds (ostriches), which exhibit marked differences in limb structure and running economy. We hypothesized that the nearly 50 per cent lower energy cost of running in ostriches is a result of: (i) lower limb-swing mechanical power, (ii) greater stance-phase storage and release of elastic energy, and (iii) lower total muscle power output. To test these hypotheses, we used three-dimensional joint mechanical measurements and a simple model to estimate the elastic and muscle contributions to joint work and power. Contradictory to our first hypothesis, we found that ostriches and humans generate the same amounts of mechanical power to swing the limbs at a similar self-selected running speed, indicating that limb swing probably does not contribute to the difference in energy cost of running between these species. In contrast, we estimated that ostriches generate 120 per cent more stance-phase mechanical joint power via release of elastic energy compared with humans. This elastic mechanical power occurs nearly exclusively at the tarsometatarso-phalangeal joint, demonstrating a shift of mechanical power generation to distal joints compared with humans. We also estimated that positive muscle fibre power is 35 per cent lower in ostriches compared with humans, and is accounted for primarily by higher capacity for storage and release of elastic energy. Furthermore, our analysis revealed much larger frontal and internal/external rotation joint loads during ostrich running than in humans. Together, these findings support the hypothesis that a primary limb structure specialization linked to economical running in cursorial species is an elevated storage and release of elastic energy in tendon. In the ostrich, energy-saving specializations may also include passive frontal and internal/external rotation load-bearing mechanisms.

SUMMARY The alleged high net energy cost of running and low net energy cost of walking in humans have played an important role in the interpretation of the evolution of human bipedalism and the biomechanical determinants of the metabolic cost of locomotion. This study re-explores how the net metabolic energy cost of running and walking (J kg–1m–1) in humans compares to that of animals of similar mass using new allometric analyses of previously published data. Firstly, this study shows that the use of the slope of the regression between the rate of energy expenditure and speed to calculate the net energy cost of locomotion overestimates the net cost of human running. Also, the net energy cost of human running is only 17% higher than that predicted based on their mass. This value is not exceptional given that over a quarter of the previously examined mammals and birds have a net energy cost of running that is 17% or more above their allometrically predicted value. Using a new allometric equation for the net energy cost of walking, this study also shows that human walking is 20%less expensive than predicted for their mass. Of the animals used to generate this equation, 25% have a relatively lower net cost of walking compared with their allometrically predicted value. This new walking allometric analysis also indicates that the scaling of the net energy cost of locomotion with body mass is gait dependent. In conclusion, the net costs of running and walking in humans are moderately different from those predicted from allometry and are not remarkable for an animal of its size.