Circulatory variables and the flight performance of birds

When considering the 'burst' flight performance of birds, such as during take-off, one of the most important structural variables is the ratio of the mass of the flight muscle myofibrils with respect to body mass. However, when considering 'prolonged' flight performance the variable of interest should be the body mass ratio of the mass of the flight muscle myofibrils that can be perfused sustainably with metabolites via the blood supply. The latter variable should be related to blood flow (ml min(-1)), which in turn has been shown to be a function of heart muscle mass, the value of which is more easily obtainable for different species than that for the mass of perfused muscle. The limited empirical evidence available suggests that for birds and mammals the rate of maximum oxygen consumption scales with heart mass (M-h) as Mh(0.88) and that for birds Mh scales with body mass (M-b) as Mb(0.92), leading to the conclusion that the rate of maximum oxygen consumption in birds scales with an exponent of around Mb(0.82). A similar exponent would be expected for the rate of maximum oxygen consumption with respect to the flight muscle mass of birds. This suggests that the sustainable power output from the flight muscles may ultimately be limiting the flight performance of very large flying animals, but as a result of circulatory constraints rather than biomechanical considerations of the flight muscles per se. Under the particular circumstances of sustainable flight performance, calculations of rates of metabolic energy consumed by the flight muscles can be compared directly with the estimates of biomechanical power output required, as calculated using various aerodynamic models. The difference between these calculated values for rates of energy input and output from the muscles is equivalent to the 'apparent' mechanochemical conversion efficiency. The results of one such analysis, of the maximum sustainable flight performance of migratory birds, leads to the conclusion that the efficiency of the flight muscles appears to scale as M-b(0.14). However, much of this apparent scaling may be an artefact of the application and assumptions of the models. The resolution of this issue is only likely to come from studying bird species at either extreme of the size range.