Abstract
The complexity of low speed maneuvering flight is apparent from the combination of two critical aspects of this behavior: high power and precise control. To understand how such control is achieved, we examined the underlying kinematics and resulting aerodynamic mechanisms of low speed turning flight in the pigeon (Columba livia). Three birds were trained to perform 90 deg level turns in a stereotypical fashion and detailed three-dimensional (3D) kinematics were recorded at high speeds. Applying the angular momentum principle, we used mechanical modeling based on time-varying 3D inertia properties of individual sections of the pigeon's body to separate angular accelerations of the torso based on aerodynamics from those based on inertial effects. Directly measured angular accelerations of the torso were predicted by aerodynamic torques, justifying inferences of aerodynamic torque generation based on inside wing versus outside wing kinematics. Surprisingly, contralateral asymmetries in wing speed did not appear to underlie the 90 deg aerial turns, nor did contralateral differences in wing area, angle of attack, wingbeat amplitude or timing. Instead, torso angular accelerations into the turn were associated with the outside wing sweeping more anteriorly compared with a more laterally directed inside wing. In addition to moving through a relatively more retracted path, the inside wing was also more strongly pronated about its long axis compared with the outside wing, offsetting any difference in aerodynamic angle of attack that might arise from the observed asymmetry in wing trajectories. Therefore, to generate roll and pitch torques into the turn, pigeons simply reorient their wing trajectories toward the desired flight direction. As a result, by acting above the center of mass, the net aerodynamic force produced by the wings is directed inward, generating the necessary torques for turning.
Highlights
To gain insight into the control of flight maneuvers, an understanding of the mechanics and aerodynamics involved in turning is needed
Reorientation of the torso in flight can be achieved using two physical principles: (i) aerodynamic forces that induce torques about the center of mass (CM), and (ii) inertia-based changes in angular momentum of body segments that result in net changes in body orientation (Frohlich, 1980; Hedrick et al, 2007)
We found that aerodynamic torques, and not inertia-based angular momentum exchanges between the wings and the rest of the bird, predicted observed torso angular roll and pitch accelerations (Fig. 4)
Summary
To gain insight into the control of flight maneuvers, an understanding of the mechanics and aerodynamics involved in turning is needed. The task of turning consists of two conceptually separate components: (1) re-direction of the animal’s flight path and (2) reorientation of the animal’s body. Re-direction of the flight path (involving changes in translational movement) requires centripetal force production. Flying vertebrates, including horseshoe bats, pigeons, fruit bats and cockatoos, all bank to orient the net aerodynamic force into the turn, analogous to how fixed-wing aircraft turn (Aldridge, 1986; Warrick, 1998; Hedrick and Biewener, 2007; Iriarte-Díaz and Swartz, 2008; Ros et al, 2011). Reorientation of the torso (involving rotational movement) in flight can be achieved using two physical principles: (i) aerodynamic forces that induce torques about the center of mass (CM), and (ii) inertia-based changes in angular momentum of body segments that result in net changes in body orientation (Frohlich, 1980; Hedrick et al, 2007). Three-dimensional (3D) analysis of the mass distributions and time-varying configurations of the head, torso and wings can be used to determine both aerodynamics- and inertiabased maneuvering in flying birds
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