Elastic deformation and energy loss of flapping fly wings

During flight, the wings of many insects undergo considerable shape changes in spanwise and chordwise directions. We determined the origin of spanwise wing deformation by combining measurements on segmental wing stiffness of the blowfly Calliphora vicina in the ventral and dorsal directions with numerical modelling of instantaneous aerodynamic and inertial forces within the stroke cycle using a two-dimensional unsteady blade elementary approach. We completed this approach by an experimental study on the wing's rotational axis during stroke reversal. The wing's local flexural stiffness ranges from 30 to 40 nN m(2) near the root, whereas the distal wing parts are highly compliant (0.6 to 2.2 nN m(2)). Local bending moments during wing flapping peak near the wing root at the beginning of each half stroke due to both aerodynamic and inertial forces, producing a maximum wing tip deflection of up to 46 deg. Blowfly wings store up to 2.30 μJ elastic potential energy that converts into a mean wing deformation power of 27.3 μW. This value equates to approximately 5.9 and 2.3% of the inertial and aerodynamic power requirements for flight in this animal, respectively. Wing elasticity measurements suggest that approximately 20% or 0.46 μJ of elastic potential energy cannot be recovered within each half stroke. Local strain energy increases from tip to root, matching the distribution of the wing's elastic protein resilin, whereas local strain energy density varies little in the spanwise direction. This study demonstrates a source of mechanical energy loss in fly flight owing to spanwise wing bending at the stroke reversals, even in cases in which aerodynamic power exceeds inertial power. Despite lower stiffness estimates, our findings are widely consistent with previous stiffness measurements on insect wings but highlight the relationship between local flexural stiffness, wing deformation power and energy expenditure in flapping insect wings.