title: The energetic costs of trunk and distal-limb loading during walking and running in guinea fowl Numida meleagris: I. Organismal metabolism and biomechanics.
abstract: We examined the energetic cost of loading the trunk or distal portion of the leg in walking and running guinea fowl (Numida meleagris). These different loading regimes were designed to separately influence the energy use by muscles used during the stance and swing phases of the stride. Metabolic rate, estimated from oxygen consumption, was measured while birds locomoted on a motorized treadmill at speeds from 0.5 to 2.0 m s-1, either unloaded, or with a mass equivalent to 23% of their body mass carried on their backs, or with masses equal to approximately 2.5% of their body mass attached to each tarsometatarsal segment. In separate experiments, we also measured the duration of stance and swing in unloaded, trunk-loaded, or limb-loaded birds. In the unloaded and limb-loaded birds, we also calculated the mechanical energy of the tarsometatarsal segment throughout the stride. Trunk and limb loads caused similar increases in metabolic rate. During trunk loading, the net metabolic rate (gross metabolic rate-resting metabolic rate) increased by 17% above the unloaded value across all speeds. This percentage increase is less than has been found in most studies of humans and other mammals. The economical load carriage of guinea fowl is consistent with predictions based on the relative cost of the stance and swing phases of the stride in this species. However, the available comparative data and considerations of the factors that determine the cost of carrying extra mass lead us to the conclusion that the cost of load carrying is unlikely to be a reliable indicator of the distribution of energy use in stance and swing. Both loading regimes caused small changes in the swing and/or stance durations, but these changes were less than 10%. Loading the tarsometatarsal segment increased its segmental energy by 4.1 times and the segmental mechanical power averaged over the stride by 3.8 times. The increases in metabolism associated with limb loading appear to be linked to the increases in mechanical power. The delta efficiency (change in mechanical power divided by the change in metabolic power) of producing this power increased from 11% in walking to approximately 25% in running. Although tarsometatarsal loading was designed to increase the mechanical energy during swing phase, 40% of the increase in segmental energy occurred during late stance. Thus, the increased energy demand of distal limb loading in guinea fowl is predicted to cause increases in energy use by both stance- and swing-phase muscles.
authors.affiliation: Department of Biology, Northeastern University, 360 Huntington Avenue, Boston, MA 02115, USA. r.marsh@neu.edu
authors.forename: Richard L
authors.initials: RL
authors.lastname: Marsh
authors.forename: David J
authors.initials: DJ
authors.lastname: Ellerby
authors.forename: Havalee T
authors.initials: HT
authors.lastname: Henry
authors.forename: Jonas
authors.initials: J
authors.lastname: Rubenson
mesh_terms: D000818:Animals; D001696:Biomechanical Phenomena; D004734:Energy Metabolism / Q000502:physiology*; D046094:Galliformes / Q000378:metabolism / Q000502:physiology*; D006614:Hindlimb / Q000502:physiology*; D012420:Running / Q000502:physiology*; D016138:Walking / Q000502:physiology*; D016474:Weight-Bearing / Q000502:physiology*