Explaining the energetics of walking and running has been difficult because the distribution of energy use among individual muscles has not been known. We estimated energy use by measuring blood flow to the hindlimb muscles in guinea fowl. Blood flow to skeletal muscles is controlled locally and varies directly with metabolic rate. We estimate that the swing-phase muscles consume 26% of the energy used by the limbs and the stance-phase muscles consume the remaining 74%, independent of speed. Thus, contrary to some previous suggestions, swinging the limbs requires an appreciable fraction of the energy used during terrestrial legged locomotion. Models integrating the energetics and mechanics of running will benefit from more detailed information on the distribution of energy use by the muscles.

, a value approximately two thirds of the maximal expected value for skeletal muscle. The muscle-mass-specific peak power output during jumping was nearly 800·W·kg -1 , which is more than twice the peak isotonic power estimated for guinea fowl leg muscles. To account for high power outputs, we concluded that energy has to be stored early in the jumps and released later during peak power production, presumably using mechanisms similar to those found in more specialized jumpers.

Running and walking are mechanically complex activities. Leg muscles must exert forces to support weight and provide stability, do work to accelerate the limbs and body centre of mass, and absorb work to act as brakes. Current understanding of energy use during legged locomotion has been limited by the lack of measurements of energy use by individual muscles. Our study is based on the correlation between blood flow and aerobic energy expenditure in active skeletal muscle during locomotion. This correlation is strongly supported by the available evidence concerning control of blood flow to active muscle, and the relationship between blood flow and the rate of muscle oxygen consumption. We used injectable microspheres to measure the blood flow to the hind-limb muscles, and other body tissues, in guinea fowl (Numida meleagris) at rest, and across a range of walking and running speeds. Combined with data concerning the various mechanical functions of the leg muscles, this approach has enabled the first direct estimates of the energetic costs of some of these functions. Cardiac output increased from 350 ml min −1 at rest, to 1700 ml min −1 at a running speed (∼2.6 m s −1 ) eliciting aV O 2 of 90% ofV O 2 ,max . The increase in cardiac output was achieved via approximately equal factorial increases in heart rate and stroke volume. Approximately 90% of the increased cardiac output was directed to the active muscles of the hind limbs, without redistribution of blood flow from the viscera. Values of mass-specific blood flow to the ventricles, ∼15 ml min −1 g −1 , and one of the hind-limb muscles, ∼9 ml min −1 g −1 , were the highest yet recorded for blood flow to active muscle. The patterns of increasing blood flow with increasing speed varied greatly among different muscles. The increases in flow correlated with the likely fibre type distribution of the muscles. Muscles expected to have many high-oxidative fibres preferentially increased flow at low exercise intensities. We estimated substantial energetic costs associated with swinging the limbs, co-contraction to stabilize the knee and work production by the hind-limb muscles. Our data provide a basis for evaluating hypotheses relating the mechanics and energetics of legged locomotion.

Explosive dehiscence ballistically disperses seeds in a number of plant species. During dehiscence, mechanical energy stored in specialized tissues is transferred to the seeds to increase their kinetic and potential energies. The resulting seed dispersal patterns have been investigated in some ballistic dispersers, but the mechanical performance of a launch mechanism of this type has not been measured. The properties of the energy storage tissue and the energy transfer efficiency of the launch mechanism were quantified in Impatiens capensis. In this species the valves forming the seed pod wall store mechanical energy. Their mass specific energy storage capacity (124 J kg−1) was comparable with that of elastin and spring steel. The energy storage capacity of the pod tissues was determined by their level of hydration, suggesting a role for turgor pressure in the energy storage mechanism. During dehiscence the valves coiled inwards, collapsing the pod and ejecting the seeds. Dehiscence took 4.2±0.4 ms (mean ±SEM, n=13). The estimated efficiency with which energy was transferred to the seeds was low (0.51±0.26%, mean ±SEM, n=13). The mean seed launch angle (17.4±5.2, mean ±SEM, n=45) fell within the range predicted by a ballistic model to maximize dispersal distance. Low ballistic dispersal efficiency or effectiveness may be characteristic of species that also utilize secondary seed dispersal mechanisms.

SUMMARY The pectoralis muscles are the main source of mechanical power for avian flight. The power output of these muscles must be modulated to meet the changing power requirements of flight across a range of speeds. This can be achieved at the muscle level by manipulation of strain trajectory and recruitment patterns, and/or by intermittent flight strategies. We have measured the in vitro power outputs of pectoralis muscle fascicles from budgerigars Melopsittacus undulatus and zebra finches Taeniopygia guttata under conditions replicating those previously measured in vivo during flight. This has allowed us to quantify the extent to which different power modulation mechanisms control flight muscle power output. Intermittent flight behaviour is a more important determinant of flight power in zebra finches than budgerigars. This behaviour accounts for 25–62% of power modulation relative to the maximum available mechanical power output in zebra finch, compared to 0–38% in budgerigars. Muscle level changes in fascicle strain trajectory and motor unit recruitment, rather than intermittent flight behaviours, are the main determinants of pectoralis muscle power output in budgerigars at all speeds, and in zebra finch at speeds below 14 m s–1.