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Published November 2014 | Published
Journal Article Open

Hovering Flight in the Honeybee Apis mellifera: Kinematic Mechanisms for Varying Aerodynamic Forces


During hovering flight, animals can increase the wing velocity and therefore the net aerodynamic force per stroke by increasing wingbeat frequency, wing stroke amplitude, or both. The magnitude and orientation of aerodynamic forces are also influenced by the geometric angle of attack, timing of wing rotation, wing contact, and pattern of deviation from the primary stroke plane. Most of the kinematic data available for flying animals are average values for wing stroke amplitude and wingbeat frequency because these features are relatively easy to measure, but it is frequently suggested that the more subtle and difficult-to-measure features of wing kinematics can explain variation in force production for different flight behaviors. Here, we test this hypothesis with multicamera high-speed recording and digitization of wing kinematics of honeybees (Apis mellifera) hovering and ascending in air and hovering in a hypodense gas (heliox: 21% O_2, 79% He). Bees employed low stroke amplitudes (86.7° ± 7.9°) and high wingbeat frequencies (226.8 ± 12.8 Hz) when hovering in air. When ascending in air or hovering in heliox, bees increased stroke amplitude by 30%–45%, which yielded a much higher wing tip velocity relative to that during simple hovering in air. Across the three flight conditions, there were no statistical differences in the amplitude of wing stroke deviation, minimum and stroke-averaged geometric angle of attack, maximum wing rotation velocity, or even wingbeat frequency. We employed a quasi-steady aerodynamic model to estimate the effects of wing tip velocity and geometric angle of attack on lift and drag. Lift forces were sensitive to variation in wing tip velocity, whereas drag was sensitive to both variation in wing tip velocity and angle of attack. Bees utilized kinematic patterns that did not maximize lift production but rather maintained lift-to-drag ratio. Thus, our data indicate that, at least for honeybees, the overall time course of wing angles is generally preserved and modulation of wing tip velocity is sufficient to perform a diverse set of vertical flight behaviors.

Additional Information

© 2014 University of Chicago Press. Accepted 8/31/2014; Electronically Published 11/10/2014. We thank David Shelton for the loan of equipment during filming and Nicholas Kostreski and J. Sean Humbert (University of Maryland, Department of Aerospace Engineering) for constructing the three-dimensional honey bee model and simulation videos used for the analysis of digitizing error. Funding was provided by a Nevada NASA Space Grant Fellowship (to J.T.V.), National Institutes of Health Fellowship F32 NS46221 (to D.L.A.), Office of Naval Research grant N00014-03-1-0604 (to M.H.D.), Packard Foundation grant 2001-17741A (to M.H.D.), and National Science Foundation grants IBN-0217229 (to M.H.D.) and IOS-0725030 (to S.P.R).

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Published - Vance-2014-Hovering-flight-in-the-honeybee-api.pdf


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August 20, 2023
August 20, 2023