The DOI matches its Crossref record; 57 of the 57 references it cites were found in a registry or an index.
Flexible clap and fling in tiny insect flight
Open accessAbstract
Of the insects that have been filmed in flight, those that are 1 mm in length or less often clap their wings together at the end of each upstroke and fling them apart at the beginning of each downstroke. This ;clap and fling' motion is thought to augment the lift forces generated during flight. What has not been highlighted in previous work is that very large forces are required to clap the wings together and to fling the wings apart at the low Reynolds numbers relevant to these tiny insects. In this paper, we use the immersed boundary method to simulate clap and fling in rigid and flexible wings. We find that the drag forces generated during fling with rigid wings can be up to 10 times larger than what would be produced without the effects of wing-wing interaction. As the horizontal components of the forces generated during the end of the upstroke and beginning of the downstroke cancel as a result of the motion of the two wings, these forces cannot be used to generate thrust. As a result, clap and fling appears to be rather inefficient for the smallest flying insects. We also add flexibility to the wings and find that the maximum drag force generated during the fling can be reduced by about 50%. In some instances, the net lift forces generated are also improved relative to the rigid wing case.
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“The immersed boundary method has been applied to a wide variety of problems in biological fluid dynamics for intermediate Re including insect flight [26, 27, 28], aquatic locomotion [16, 15], and ciliary driven flows [18].”
From Reynolds number limits for jet propulsion: a numerical study of simplified jellyfish (2011) · cites this paper as [28]
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“For these low Re , leading and trailing edge vortices are diffuse and do not separate from the wings.”
From Reynolds number limits for jet propulsion: a numerical study of simplified jellyfish (2011) · cites this paper as [28]
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“flapping performance (defined as the ratio of lift to drag) drastically decreases for Re < 10 [28].”
From Reynolds number limits for jet propulsion: a numerical study of simplified jellyfish (2011) · cites this paper as [28]
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Cites
Peskin, C. S. (2002) · Acta Numerica · doi:10.1017/s0962492902000077 · Cited by 4,603
Self-citationWeis‐Fogh, T. (1973) · Journal of Experimental Biology · doi:10.1242/jeb.59.1.169 · Cited by 1,373
The aerodynamics of insect flight
Sane, S. P. (2003) · Journal of Experimental Biology · doi:10.1242/jeb.00663 · Cited by 1,269
The Aerodynamics of Hovering Insect Flight. III. Kinematics
Ellington, C. (1984) · Philosophical transactions of the Royal Society of London. Series B, Biological sciences · doi:10.1098/rstb.1984.0051 · Cited by 840
The novel aerodynamics of insect flight: applications to micro-air vehicles
Ellington, C. P. (1999) · Journal of Experimental Biology · doi:10.1242/jeb.202.23.3439 · Cited by 768
The Biomechanics of Insect Flight
Dudley, R. (2000) · Princeton University Press eBooks · doi:10.1515/9780691186344 · Cited by 758
(2000) · doi:10.1093/aesa/93.5.1195f · Cited by 710
Unsteady Aerodynamic Performance of Model Wings at Low Reynolds Numbers
Dickinson, M. H., & Götz, K. (1993) · Journal of Experimental Biology · doi:10.1242/jeb.174.1.45 · Cited by 671
Wang, Z. J. (2003) · Annual Review of Fluid Mechanics · doi:10.1146/annurev.fluid.36.050802.121940 · Cited by 640
The aerodynamics of hovering insect flight. IV. Aerodynamic mechanisms
Ellington, C. P. (1984) · Philosophical transactions of the Royal Society of London. Series B, Biological sciences · doi:10.1098/rstb.1984.0052 · Cited by 533
Flagged first, then by citations
Research works printed but not linked: 1
- Tanaka, S. (1995). Thrips' flight. Part 1. In Symposia '95 of Exploratory Research for Advanced Technology, Japan Science and Technology Corporation, Abstracts (ed. K. Kawachi),pp. 27-34. Tokyo: Japan Science and Technology Corporation.
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Metadata
- Authors
- Miller, L. A., & Peskin, C. S.
- Affiliations
- University of North Carolina at Chapel Hill (US)
- Corresponding author
- Miller, Laura A.
- Venue
- Journal of Experimental Biology
- Publisher
- The Company of Biologists
- Published
- 2009-10-01
- Volume
- 212
- Issue
- 19
- Pages
- 3076-3090
- ISSN
- 0022-0949 (ISSN-L)
- 1477-9145
- OpenAlex
- W2141248119
- Cited by
- 179
- References
- 57
- Citations by year
- 2026: 5 · 2025: 10 · 2024: 9 · 2023: 5 · 2022: 8 · 2021: 17 · 2020: 13 · 2019: 20 · 2018: 11 · 2017: 22 · 2016: 11 · 2015: 12 · 2014: 13 · 2013: 5 · 2012: 7
- FWCI
- 11.63
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Journal of Experimental Biology (publishedVersion)
- PubMed (publishedVersion)
- Carolina Digital Repository (University of North Carolina at Chapel Hill) (submittedVersion)
- UNC Libraries
- Topics
- Physical Sciences › Engineering › Aerospace Engineering › Biomimetic flight and propulsion mechanisms
- Physical Sciences › Engineering › Computational Mechanics › Fluid dynamics and aerodynamics studies
- Physical Sciences › Engineering › Aerospace Engineering › Aeroelasticity and Vibration Control
- Keywords
- clap-and-fling, insect flight, wing flexibility, wing-wing interaction, drag force, lift generation, low Reynolds number aerodynamics, upstroke and downstroke, thrust generation, immersed boundary method
- MeSH terms
- Animals, Biomechanical Phenomena, Flight, Animal, Insecta, Wings, Animal, Body Size
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