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Science 18 June 1999:
Vol. 284. no. 5422, pp. 1954 - 1960
DOI: 10.1126/science.284.5422.1954
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Research Articles

Wing Rotation and the Aerodynamic Basis of Insect Flight

Michael H. Dickinson, 1* Fritz-Olaf Lehmann, 2 Sanjay P. Sane 1

The enhanced aerodynamic performance of insects results from an interaction of three distinct yet interactive mechanisms: delayed stall, rotational circulation, and wake capture. Delayed stall functions during the translational portions of the stroke, when the wings sweep through the air with a large angle of attack. In contrast, rotational circulation and wake capture generate aerodynamic forces during stroke reversals, when the wings rapidly rotate and change direction. In addition to contributing to the lift required to keep an insect aloft, these two rotational mechanisms provide a potent means by which the animal can modulate the direction and magnitude of flight forces during steering maneuvers. A comprehensive theory incorporating both translational and rotational mechanisms may explain the diverse patterns of wing motion displayed by different species of insects.

1 Department of Integrative Biology, University of California, Berkeley, CA 94720, USA.
2 Theodor-Boveri-Institute, Department of Behavioral Physiology and Sociobiological Zoology, University of Würzburg am Hubland, 97074 Würzburg, Germany.
*   To whom correspondence should be addressed. E-mail: flymanmd{at}socrates.berkeley.edu

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   Abstract »    Full Text »    PDF »
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