Flight feather attachment in rock pigeons (Columba livia): covert feathers and smooth muscle coordinate a morphing wing.

Mechanisms for passively coordinating forelimb movements and flight feather abduction and adduction have been described separately from both in vivo and ex vivo studies. Skeletal coordination has been identified as a way for birds to simplify the neuromotor task of controlling flight stroke, but an understanding of the relationship between skeletal coordination and the coordination of the aerodynamic control surface (the flight feathers) has been slow to materialize. This break between the biomechanical and aerodynamic approaches - between skeletal kinematics and airfoil shape - has hindered the study of dynamic flight behaviors. Here I use dissection and histology to identify previously overlooked interconnections between musculoskeletal elements and flight feathers. Many of these structures are well-placed to directly link elements of the passive musculoskeletal coordination system with flight feather movements. Small bundles of smooth muscle form prominent connections between upper forearm coverts (deck feathers) and the ulna, as well as the majority of interconnections between major flight feathers of the hand. Abundant smooth muscle may play a role in efficient maintenance of folded wing posture, and may also provide an autonomically regulated means of tuning wing shape and aeroelastic behavior in flight. The pattern of muscular and ligamentous linkages of flight feathers to underlying muscle and bone may provide predictable passive guidance for the shape of the airfoil during flight stroke. The structures described here provide an anatomical touchstone for in vivo experimental tests of wing surface coordination in an extensively researched avian model species.