Aerodynamics of tip-reversal upstroke in a revolving pigeon wing

During slow flight, bird species vary in their upstroke kinematics using either a 'flexed wing' or a distally supinated 'tip-reversal' upstroke. Two hypotheses have been presented concerning the function of the tip-reversal upstroke. The first is that this behavior is aerodynamically inactive and serves to minimize drag. The second is that the tip-reversal upstroke is capable of producing significant aerodynamic forces. Here, we explored the aerodynamic capabilities of the tip-reversal upstroke using a well-established propeller method. Rock dove (Columba livia, N=3) wings were spread and dried in postures characteristic of either mid-upstroke or mid-downstroke and spun at in vivo Reynolds numbers to simulate forces experienced during slow flight. We compared 3D wing shape for the propeller and in vivo kinematics, and found reasonable kinematic agreement between methods (mean differences 6.4% of wing length). We found that the wing in the upstroke posture is capable of producing substantial aerodynamic forces. At in vivo angles of attack (66 deg at mid-upstroke, 46 deg at mid-downstroke), the upstroke wings averaged for three birds produced a lift-to-drag ratio of 0.91, and the downstroke wings produced a lift-to-drag ratio of 3.33. Peak lift-to-drag ratio was 2.5 for upstroke and 6.3 for downstroke. Our estimates of total force production during each half-stroke suggest that downstroke produces a force that supports 115% of bodyweight, and during upstroke a forward-directed force (thrust) is produced at 36% of body weight.