The energy cost of loaded flight is substantially lower than expected due to alterations in flight kinematics

The effect of experimentally increased wing loading on the energy cost of flight was examined in cockatiels Nyphicus hollandicus. Five individuals were flown for periods of approximately 2 min, while carrying additional payload mass amounting to between 5 and 20% of unloaded body mass. The energy cost of flight was measured using the 13C-labelled bicarbonate technique, which was also calibrated in a separate experiment on resting birds, by comparing the elimination rate of 13C in breath with a simultaneous measurement of oxygen consumption by indirect calorimetry. It was not possible to perform a similar calibration during flight when energy costs were higher, so we extrapolated the relationship from the resting calibration to predict flight cost. Flight cost in the pre-manipulated individuals averaged 16.7+/-1.8 W. Flight cost in the pre-manipulated birds was significantly related to the interaction between downstroke duration and flight speed. There was no significant increase in flight cost with increases in payload mass. The birds responded to payload masses between 5 and 15% of their unloaded body mass by decreasing flight speed relative to unloaded birds, while maintaining wing beat frequency (Fb). At a payload mass equivalent to 20% of body mass, however, the birds flew at higher speeds than unloaded controls, and had a significantly higher Fb, generated by a reduction in both the upstroke and downstroke durations. Wing amplitude was unaffected by the increase in loading. Using the measured flight parameters, the effect of loading was not significantly different than predicted using aerodynamic models.