Functional morphology of the vanes of the flight feathers of the pigeon Columba livia.

The functional morphology of feather vanes was examined by combining morphological examination with mechanical tests. A geometrical model was derived which related the in-plane behaviour of the vane to the orientation of the barbs and barbules. This predicted that the small branching angles of both barbs and proximal barbules should result in a vane which is easier to move distally than proximally. These predictions were verified by mechanical tests on primary and secondary feathers of the pigeon Columba livia. A further prediction, that the inclination of the barbs' cross section should make the vanes more resistant to forces from below than those from above, was also confirmed by mechanical tests. Differences in the mechanical behaviour of feathers are related to differences in their morphology and function. The vanes of outer primaries are more resistant to out-of-plane forces than those of the inner primaries and secondaries, particularly towards their tip, a property which will help them withstand the larger aerodynamic forces to which they may be subjected in flight. The outermost primary vane also showed the least asymmetry to out-of-plane forces as a result of the more vertical orientation of its barbs. This may help it to act as a reversible aerofoil during take-off.