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      Centripetal Acceleration Reaction: An Effective and Robust Mechanism for Flapping Flight in Insects

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      PLoS ONE
      Public Library of Science

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          Abstract

          Despite intense study by physicists and biologists, we do not fully understand the unsteady aerodynamics that relate insect wing morphology and kinematics to lift generation. Here, we formulate a force partitioning method (FPM) and implement it within a computational fluid dynamic model to provide an unambiguous and physically insightful division of aerodynamic force into components associated with wing kinematics, vorticity, and viscosity. Application of the FPM to hawkmoth and fruit fly flight shows that the leading-edge vortex is the dominant mechanism for lift generation for both these insects and contributes between 72–85% of the net lift. However, there is another, previously unidentified mechanism, the centripetal acceleration reaction, which generates up to 17% of the net lift. The centripetal acceleration reaction is similar to the classical inviscid added-mass in that it depends only on the kinematics (i.e. accelerations) of the body, but is different in that it requires the satisfaction of the no-slip condition, and a combination of tangential motion and rotation of the wing surface. Furthermore, the classical added-mass force is identically zero for cyclic motion but this is not true of the centripetal acceleration reaction. Furthermore, unlike the lift due to vorticity, centripetal acceleration reaction lift is insensitive to Reynolds number and to environmental flow perturbations, making it an important contributor to insect flight stability and miniaturization. This force mechanism also has broad implications for flow-induced deformation and vibration, underwater locomotion and flows involving bubbles and droplets.

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          Most cited references13

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          Wing rotation and the aerodynamic basis of insect flight.

          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.
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            Spanwise flow and the attachment of the leading-edge vortex on insect wings.

            The flow structure that is largely responsible for the good performance of insect wings has recently been identified as a leading-edge vortex. But because such vortices become detached from a wing in two-dimensional flow, an unknown mechanism must keep them attached to (three-dimensional) flapping wings. The current explanation, analogous to a mechanism operating on delta-wing aircraft, is that spanwise flow through a spiral vortex drains energy from the vortex core. We have tested this hypothesis by systematically mapping the flow generated by a dynamically scaled model insect while simultaneously measuring the resulting aerodynamic forces. Here we report that, at the Reynolds numbers matching the flows relevant for most insects, flapping wings do not generate a spiral vortex akin to that produced by delta-wing aircraft. We also find that limiting spanwise flow with fences and edge baffles does not cause detachment of the leading-edge vortex. The data support an alternative hypothesis-that downward flow induced by tip vortices limits the growth of the leading-edge vortex.
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              The aerodynamics of free-flight maneuvers in Drosophila.

              Using three-dimensional infrared high-speed video, we captured the wing and body kinematics of free-flying fruit flies as they performed rapid flight maneuvers. We then "replayed" the wing kinematics on a dynamically scaled robotic model to measure the aerodynamic forces produced by the wings. The results show that a fly generates rapid turns with surprisingly subtle modifications in wing motion, which nonetheless generate sufficient torque for the fly to rotate its body through each turn. The magnitude and time course of the torque and body motion during rapid turns indicate that inertia, not friction, dominates the flight dynamics of insects.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                2015
                7 August 2015
                : 10
                : 8
                : e0132093
                Affiliations
                [1 ]Department of Mechanical Engineering, the Johns Hopkins University, Baltimore, Maryland, United States of America
                [2 ]Department of Biology, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, United States of America
                Brown University, UNITED STATES
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: CZ TH RM. Performed the experiments: CZ TH. Analyzed the data: CZ TH RM. Contributed reagents/materials/analysis tools: CZ TH RM. Wrote the paper: CZ TH RM.

                Article
                PONE-D-15-13262
                10.1371/journal.pone.0132093
                4529139
                26252016
                a93116e7-c4ad-4117-88d2-f5f6dbda6a46
                Copyright @ 2015

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited

                History
                : 26 March 2015
                : 10 June 2015
                Page count
                Figures: 4, Tables: 1, Pages: 16
                Funding
                This work was supported by Air Force Office of Scientific Research FA9550-10-1-006 RM TH; National Science Foundation CBET-1357819, IOS-1124804, PLR-1246317 RM; and National Science Foundation IOS-0920358 TH. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
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                All relevant data are within the paper and its Supporting Information files.

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