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1 Department of Biology, Northeastern University, 360 Huntington Avenue, Boston, MA 02115, USA
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 a 
of 90% of 
. 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.
(Received 13 January 2005;
accepted after revision 20 February 2005;
first published online 24 February 2005)
Corresponding author R. L. Marsh: Department of Biology, Northeastern University, 360 Huntington Avenue, Boston, MA 02115, USA. Email: r.marsh{at}neu.edu
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