The scaling of uphill and downhill locomotion in legged animals

Animals must continually respond dynamically as they move through complex environments, and slopes are a common terrain on which legged animals must move. Despite this, non-level locomotion remains poorly understood. In this study, we first review the literature on locomotor mechanics, metabolic cost, and kinematic strategies on slopes. Using existing literature we then performed scaling analyses of kinematic variables, including speed, duty factor, and stride-length across a range of body sizes from ants to horses. The studies that examined locomotion on inclines vastly outnumbered those focusing on declines. On inclines, animals tend to reduce speed and increase duty factor, but a similar consensus could not be reached for declines. Remarkably, stride-length did not differ between locomotion on inclines and on level terrain, but this may have resulted from data only being available for low slopes (<30°). On declines there appears to be a shift in locomotor strategy that is size-dependent. At masses <1-10 kg, animals tended to use shorter strides than on level terrain, and the opposite occurred at larger body masses. Therefore, possibly due to stability issues, body mass plays a significant role in the locomotor strategy used when traveling downhill. Although we currently lack sufficient data, differential leg function is likely to be critical for locomotion on slopes, with mechanical demands differing on limbs during movement on level, inclined, and declined surfaces. Our scaling analysis not only highlights areas that require future work, but also suggests that body size is important for determining the mechanics and strategies animals use to negotiate non-level terrain. It is clear that selection has resulted in an incredible range of body size among animals, both extant and extinct, and it is likely that the ability to move up and down slopes has constrained or relaxed these mechanical pressures. Given the lack of integration of ecological data with laboratory experiments, future work should first determine which inclines animals actually use in nature, as this likely plays a key role in behaviors such as predator-prey interactions.