Insects were the first animals to evolve active flight, and remain unsurpassed in many features of aerodynamic performance (Dudley, 2000
). In recent years, studies using a variety of methods including physical modeling, flow visualization, and computational fluid mechanics have converged to provide a more detailed understanding of flapping flight aerodynamics (Ellington et al., 1996; Dickinson et al., 1999b; Srygley and Thomas, 2002; Sane, 2003). Although critical issues remain, the basic means by which insects generate enough lift to stay aloft have been resolved and new research can more easily focus on the challenging question of how insects actively manipulate aerodynamic forces to steer and maneuver.
When considering the control of complex aerial behaviors, it is impossible to disentangle the aerodynamics of flapping from the mechanics of the wing hinge, the physiology of the flight muscles, or the properties of sensory-motor circuits in the brain. Thus, an integrative approach that addresses the functional interactions among these various components is essential for identifying the organizational principles of the flight control system. In this review, I will describe a single stereotyped flight maneuver of the fruit fly, Drosophila melanogaster, at several levels of analysis— from the sensory signals that trigger the behavior to the aerodynamic forces generated by the wings. The results show how the rules governing the physical interaction of an animal with the external world strongly influence the evolution of the neural circuits that control locomotion.
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