{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,11]],"date-time":"2026-02-11T20:36:10Z","timestamp":1770842170194,"version":"3.50.1"},"reference-count":57,"publisher":"The Company of Biologists","license":[{"start":{"date-parts":[[2020,1,1]],"date-time":"2020-01-01T00:00:00Z","timestamp":1577836800000},"content-version":"am","delay-in-days":0,"URL":"http:\/\/www.biologists.com\/user-licence-1-1\/"}],"funder":[{"DOI":"10.13039\/100007602","name":"University of California, Riverside","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100007602","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:p>Why do hummingbirds hum and insects whine when their wings flap in flight? Gutin proposed that a spinning propeller produces tonal sound because the location of the center of aerodynamic pressure on each blade oscillates relative to an external receiver. Animal wings also move, and in addition, aerodynamic force produced by animal wings fluctuates in magnitude and direction over the course of the wingbeat. Here we model animal wing tone as the equal, opposite reaction to aerodynamic forces on the wing, using Lowson's equation for the sound field produced by a moving point force. Two assumptions of Lowson's equation are met: animal flight is low (&amp;lt; 0.3) Mach and animals from albatrosses to mosquitoes are acoustically compact, meaning they have a small spatial extent relative to the wavelength of their wingbeat frequency. This model predicts the acoustic waveform of a hovering Costa's Hummingbird (Calypte costae), which varies in the X, Y, and Z directions around the animal. We modeled the wing forces of a hovering animal as a sinusoid with an amplitude equal to body weight. This model predicts wing sound pressure levels below a hovering hummingbird and mosquito to within 2 dB; and that far-field mosquito wing tone attenuates to 20 dB within about 0.2 m of the animal, while hummingbird humming attenuates to 20 dB at about 10 m. Wing tones play a role in communication of certain insects, such as mosquitoes, and influences predator-prey interactions, because it potentially reveals the predator's presence to its intended prey.<\/jats:p>","DOI":"10.1242\/jeb.214965","type":"journal-article","created":{"date-parts":[[2020,8,25]],"date-time":"2020-08-25T13:10:14Z","timestamp":1598361014000},"source":"Crossref","is-referenced-by-count":7,"title":["Humming hummingbirds, insect flight tones, and a model of animal flight sound"],"prefix":"10.1242","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7943-9291","authenticated-orcid":false,"given":"Christopher J.","family":"Clark","sequence":"first","affiliation":[{"name":"Department of Evolution, Ecology, and Organismal Biology, University of California, Riverside, Riverside CA 92521, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Emily A.","family":"Mistick","sequence":"additional","affiliation":[{"name":"Department of Evolution, Ecology, and Organismal Biology, University of California, Riverside, Riverside CA 92521, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"237","published-online":{"date-parts":[[2020,1,1]]},"reference":[{"key":"2021042700360548100_JEB214965C1","doi-asserted-by":"publisher","first-page":"4070","DOI":"10.1242\/jeb.075044","article-title":"Wingbeat kinematics and motor control of yaw turns in Anna's Hummingbirds (Calypte anna)","volume":"215","author":"Altshuler","year":"2012","journal-title":"J. Exp. Biol."},{"key":"2021042700360548100_JEB214965C2","doi-asserted-by":"publisher","first-page":"933","DOI":"10.1121\/1.4861233","article-title":"Mosquito (Aedes aegypti) flight tones: frequency, harmonicity, spherical spreading, and phase relationships","volume":"135","author":"Arthur","year":"2014","journal-title":"J. Acoust. Soc. Am."},{"key":"2021042700360548100_JEB214965C3","doi-asserted-by":"publisher","first-page":"72","DOI":"10.1121\/1.2932340","article-title":"Aerodynamic sound generation of flapping wing","volume":"124","author":"Bae","year":"2008","journal-title":"J. Acoust. Soc. Am."},{"key":"2021042700360548100_JEB214965C4","doi-asserted-by":"crossref","first-page":"505","DOI":"10.1016\/B978-0-12-809274-3.00006-4","article-title":"Chapter 6 - noise from rotating machinery","volume-title":"Mechanics of Flow-Induced Sound and Vibration","author":"Blake","year":"2017","edition":"2nd edn."},{"key":"2021042700360548100_JEB214965C5","volume-title":"Mechanics of Flow-Induced Sound and Vibration","author":"Blake","year":"2017"},{"key":"2021042700360548100_JEB214965C6","doi-asserted-by":"publisher","first-page":"1524","DOI":"10.1242\/jeb.128231","article-title":"Determination of the wingsnap sonation mechanism of the golden-collared manakin (Manacus vitellinus)","volume":"219","author":"Bodony","year":"2016","journal-title":"J. Exp. Biol."},{"key":"2021042700360548100_JEB214965C7","doi-asserted-by":"publisher","first-page":"484","DOI":"10.1016\/j.beproc.2018.06.011","article-title":"The sounds of silence: Barn owl noise in landing and taking off","volume":"157","author":"Boonman","year":"2018","journal-title":"Behav. Process."},{"key":"2021042700360548100_JEB214965C8","doi-asserted-by":"publisher","first-page":"icaa085","DOI":"10.1093\/icb\/icaa085","article-title":"Wing-beat frequency and its acoustics in birds and bats","author":"Boonman","year":"2020","journal-title":"Integr. Comp. Biol."},{"key":"2021042700360548100_JEB214965C9","doi-asserted-by":"publisher","first-page":"118","DOI":"10.26786\/1920-7603(2011)17","article-title":"Comparative bioacoustical studies on flight and buzzing of neotropical bees","volume":"6","author":"Burkart","year":"2011","journal-title":"J. Pollination Ecol."},{"key":"2021042700360548100_JEB214965C10","doi-asserted-by":"publisher","first-page":"1077","DOI":"10.1126\/science.1166541","article-title":"Harmonic convergence in the love songs of the dengue vector mosquito","volume":"323","author":"Cator","year":"2009","journal-title":"Science"},{"key":"2021042700360548100_JEB214965C11","volume-title":"Raven Pro 1.3 User's Manual","author":"Charif","year":"2008"},{"key":"2021042700360548100_JEB214965C12","doi-asserted-by":"publisher","first-page":"5354","DOI":"10.1038\/s41467-019-13347-3","article-title":"Birds repurpose the role of drag and lift to take off and land","volume":"10","author":"Chin","year":"2019","journal-title":"Nat. Commun."},{"key":"2021042700360548100_JEB214965C13","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-319-27721-9_4","article-title":"Locomotion-induced sounds and sonations: mechanisms, communication function, and relationship with behavior","volume-title":"Vertebrate Sound Production and Acoustic Communication","author":"Clark","year":"2016"},{"key":"2021042700360548100_JEB214965C14","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1016\/j.anbehav.2018.07.009","article-title":"Signal or cue? Locomotion-induced sounds and the evolution of communication","volume":"143","author":"Clark","year":"2018","journal-title":"Anim. Behav."},{"key":"2021042700360548100_JEB214965C15","doi-asserted-by":"publisher","first-page":"jeb173625","DOI":"10.1242\/jeb.173625","article-title":"Kinematic control of male Allen's Hummingbird wing trill over a range of flight speeds","volume":"221","author":"Clark","year":"2018","journal-title":"J. Exp. Biol."},{"key":"2021042700360548100_JEB214965C16","doi-asserted-by":"publisher","first-page":"3395","DOI":"10.1242\/jeb.080317","article-title":"Hummingbird feather sounds are produced by aeroelastic flutter, not vortex-induced vibration","volume":"216","author":"Clark","year":"2013","journal-title":"J. Exp. Biol."},{"key":"2021042700360548100_JEB214965C17","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1016\/j.anbehav.2015.02.002","article-title":"Use of acoustic signals in mating in an eavesdropping frog-biting midge","volume":"103","author":"de Silva","year":"2015","journal-title":"Anim. Behav."},{"key":"2021042700360548100_JEB214965C18","doi-asserted-by":"crossref","DOI":"10.1515\/9780691186344","volume-title":"The Biomechanics of Insect Flight: Form, Function, Evolution","author":"Dudley","year":"2000"},{"key":"2021042700360548100_JEB214965C19","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1098\/rsta.1969.0031","article-title":"Sound generation by turbulence and surfaces in arbitrary motion","volume":"264","author":"Ffowcs Williams","year":"1969","journal-title":"Philos. Trans. R. Soc. Lond. A"},{"key":"2021042700360548100_JEB214965C20","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1242\/jeb.156.1.215","article-title":"Mechanics of a rapid running insect: two-, four- and six-legged locomotion","volume":"156","author":"Full","year":"1991","journal-title":"J. Exp. Biol."},{"key":"2021042700360548100_JEB214965C21","doi-asserted-by":"crossref","first-page":"xi","DOI":"10.1016\/B978-0-12-809651-2.09992-6","article-title":"Chapter 16: Open rotor noise","volume-title":"Aeroacoustics of Low Mach Number Flows","author":"Glegg","year":"2017"},{"key":"2021042700360548100_JEB214965C22","doi-asserted-by":"publisher","first-page":"837","DOI":"10.1017\/S0368393100109915","article-title":"The silent flight of owls","volume":"38","author":"Graham","year":"1934","journal-title":"J. R. Aeronaut. Soc."},{"key":"2021042700360548100_JEB214965C23","first-page":"1","article-title":"Dimensional relationships for flying animals","volume-title":"Smithsonian Miscellaneous Collections","author":"Greenewalt","year":"1962"},{"key":"2021042700360548100_JEB214965C24","volume-title":"On the Sound Field of a Rotating Propeller","author":"Gutin","year":"1948"},{"key":"2021042700360548100_JEB214965C25","doi-asserted-by":"publisher","first-page":"064001","DOI":"10.1088\/1748-3190\/aa7eb2","article-title":"Design and analysis of aerodynamic force platforms for free flight studies","volume":"12","author":"Hightower","year":"2017","journal-title":"Bioinspir. Biomim."},{"key":"2021042700360548100_JEB214965C26","volume-title":"Acoustics of Fluid-Structure Interactions","author":"Howe","year":"2008"},{"key":"2021042700360548100_JEB214965C27","doi-asserted-by":"publisher","first-page":"923","DOI":"10.1007\/s00265-007-0516-x","article-title":"Wait before running for your life: defensive tactics of spiny mice (Acomys cahirinus) in evading barn owl (Tyto alba) attack","volume":"62","author":"Ilany","year":"2008","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2021042700360548100_JEB214965C28","first-page":"437","article-title":"Numerical analysis of sound generation of insect flapping wings","volume":"57","author":"Inada","year":"2009","journal-title":"Theor. Appl. Mech. Jpn."},{"key":"2021042700360548100_JEB214965C29","doi-asserted-by":"crossref","DOI":"10.2174\/9781681083179117010004","article-title":"Sound and sound sources","volume-title":"Comparative Bioacoustics: An Overview","author":"Larsen","year":"2017"},{"key":"2021042700360548100_JEB214965C30","doi-asserted-by":"publisher","first-page":"20141283","DOI":"10.1098\/rsif.2014.1283","article-title":"In vivo recording of aerodynamic force with an aerodynamic force platform: from drones to birds","volume":"12","author":"Lentink","year":"2015","journal-title":"J. R. Soc. Interface"},{"key":"2021042700360548100_JEB214965C31","doi-asserted-by":"publisher","first-page":"icaa045","DOI":"10.1093\/icb\/icaa045","article-title":"Evidence that the dorsal velvet of Barn Owl wing feathers decreases rubbing sounds during flapping flight","author":"LePiane","year":"2020","journal-title":"Integr. Comp. Biol."},{"key":"2021042700360548100_JEB214965C32","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1098\/rspa.1962.0090","article-title":"The Bakerian lecture, 1961 Sound generated aerodynamically","volume":"267","author":"Lighthill","year":"1962","journal-title":"Proc. R. Soc. A"},{"key":"2021042700360548100_JEB214965C33","doi-asserted-by":"publisher","first-page":"559","DOI":"10.1098\/rspa.1965.0164","article-title":"The sound field for singularities in motion","volume":"286","author":"Lowson","year":"1965","journal-title":"Proc. R. Soc. Math. Phys. Eng. Sci. A"},{"key":"2021042700360548100_JEB214965C34","first-page":"1","article-title":"Propeller and propfan noise","volume-title":"Aeroacoustics of Flight Vehicles: Theory and Practice","author":"Magliozzi","year":"1981"},{"key":"2021042700360548100_JEB214965C35","doi-asserted-by":"publisher","first-page":"284","DOI":"10.1111\/joa.12313","article-title":"Structure and function of the mammalian middle ear. I: Large middle ears in small desert mammals","volume":"228","author":"Mason","year":"2016","journal-title":"J. Anat."},{"key":"2021042700360548100_JEB214965C36","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1007\/BF00290824","article-title":"Sound and vibrational signals in the dance language of the honeybee, Apis mellifera","volume":"18","author":"Michelsen","year":"1986","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2021042700360548100_JEB214965C37","doi-asserted-by":"publisher","first-page":"393","DOI":"10.1017\/S0094837300019801","article-title":"Supersonic sauropods?","volume":"23","author":"Myhrvold","year":"1997","journal-title":"Tail Dyn. Diplodocids Paleobiol."},{"key":"2021042700360548100_JEB214965C38","doi-asserted-by":"publisher","first-page":"366","DOI":"10.1016\/j.jsv.2018.11.014","article-title":"Effects of flapping wing kinematics on the aeroacoustics of hovering flight","volume":"442","author":"Nedunchezian","year":"2019","journal-title":"J. Sound Vib."},{"key":"2021042700360548100_JEB214965C39","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1242\/jeb.150.1.171","article-title":"Predicting wingbeat frequency and wavelength of birds","volume":"150","author":"Pennycuick","year":"1990","journal-title":"J. Exp. Biol."},{"key":"2021042700360548100_JEB214965C40","doi-asserted-by":"crossref","first-page":"2607","DOI":"10.1242\/jeb.204.15.2607","article-title":"The control of flight force by a flapping wing: lift and drag production","volume":"204","author":"Sane","year":"2001","journal-title":"J. Exp. Biol."},{"key":"2021042700360548100_JEB214965C41","doi-asserted-by":"publisher","first-page":"769","DOI":"10.2514\/1.J050703","article-title":"Silent owl flight: bird flyover noise measurements","volume":"49","author":"Sarradj","year":"2011","journal-title":"AIAA J."},{"key":"2021042700360548100_JEB214965C42","first-page":"65","article-title":"Rotor noise","volume-title":"Aeroacoustics of Flight Vehicles: Theory and Practice","author":"Schmitz","year":"1981"},{"key":"2021042700360548100_JEB214965C43","doi-asserted-by":"publisher","first-page":"016008","DOI":"10.1088\/1748-3190\/ab54fc","article-title":"Mechanism and scaling of wing tone generation in mosquitoes","volume":"15","author":"Seo","year":"2019","journal-title":"Bioinspir. Biomim."},{"key":"2021042700360548100_JEB214965C44","doi-asserted-by":"publisher","first-page":"20140541","DOI":"10.1098\/rsif.2014.0541","article-title":"Three-dimensional flow and lift characteristics of a hovering ruby-throated hummingbird","volume":"11","author":"Song","year":"2014","journal-title":"J. R. Soc. Interface"},{"key":"2021042700360548100_JEB214965C45","doi-asserted-by":"publisher","first-page":"1057","DOI":"10.1038\/1701057a0","article-title":"Flight-tone and wing-stroke frequency of insects and the dynamics of insect flight","volume":"170","author":"Sotavalta","year":"1952","journal-title":"Nature"},{"key":"2021042700360548100_JEB214965C46","doi-asserted-by":"publisher","first-page":"385","DOI":"10.1146\/annurev.en.19.010174.002125","article-title":"Courtship behavior in Drosophila","volume":"19","author":"Spieth","year":"1974","journal-title":"Annu. Rev. Entomol."},{"key":"2021042700360548100_JEB214965C47","doi-asserted-by":"publisher","first-page":"853","DOI":"10.2307\/1369527","article-title":"Behavioral, ecological and morphological correlates of foraging for arthropods by the hummingbirds of a tropical wet forest","volume":"97","author":"Stiles","year":"1995","journal-title":"Condor"},{"key":"2021042700360548100_JEB214965C48","doi-asserted-by":"publisher","first-page":"530","DOI":"10.1121\/1.1932227","article-title":"Sound radiation around a flying fly","volume":"118","author":"Sueur","year":"2005","journal-title":"J. Acoust. Soc. Am."},{"key":"2021042700360548100_JEB214965C49","doi-asserted-by":"publisher","first-page":"jeb211862","DOI":"10.1242\/jeb.211862","article-title":"Hearing in caterpillars of the monarch butterfly (Danaus plexippus)","volume":"222","author":"Taylor","year":"2019","journal-title":"J. Exp. Biol."},{"key":"2021042700360548100_JEB214965C50","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1242\/jeb.201.3.403","article-title":"Gliding flight: speed and acceleration of ideal falcons during diving and pull out","volume":"201","author":"Tucker","year":"1998","journal-title":"J. Exp. Biol."},{"key":"2021042700360548100_JEB214965C51","doi-asserted-by":"publisher","first-page":"1483","DOI":"10.1111\/ele.13331","article-title":"Flowers respond to pollinator sound within minutes by increasing nectar sugar concentration","volume":"22","author":"Veits","year":"2019","journal-title":"Ecol. Lett."},{"key":"2021042700360548100_JEB214965C52","doi-asserted-by":"crossref","first-page":"3783","DOI":"10.1242\/jeb.205.24.3783","article-title":"Rotational lift: something different or more of the same?","volume":"205","author":"Walker","year":"2002","journal-title":"J. Exp. Biol."},{"key":"2021042700360548100_JEB214965C53","doi-asserted-by":"crossref","first-page":"761","DOI":"10.1242\/jeb.64.3.761","article-title":"Acoustical properties of the flight activities of the Caribbean fruit fly","volume":"64","author":"Webb","year":"1976","journal-title":"J. Exp. Biol."},{"key":"2021042700360548100_JEB214965C54","doi-asserted-by":"publisher","first-page":"248","DOI":"10.1086\/physzool.35.3.30152809","article-title":"A function of the enlarged middle-ear cavities of the kangaroo rat, Dipodomys","volume":"35","author":"Webster","year":"1962","journal-title":"Physiol. Zool."},{"key":"2021042700360548100_JEB214965C55","doi-asserted-by":"publisher","first-page":"310","DOI":"10.1159\/000125441","article-title":"Adaptive value of hearing and vision in kangaroo rat predator avoidance","volume":"4","author":"Webster","year":"1971","journal-title":"Brain Behav. Evol."},{"key":"2021042700360548100_JEB214965C56","doi-asserted-by":"publisher","first-page":"247","DOI":"10.1093\/icb\/20.1.247","article-title":"Morphological adaptations of the ear in the rodent family Heteromyidae","volume":"20","author":"Webster","year":"1980","journal-title":"Am. Zool."},{"key":"2021042700360548100_JEB214965C57","doi-asserted-by":"publisher","first-page":"300","DOI":"10.2307\/1538745","article-title":"Oscilloscopic and stroboscopic analysis of the flight sounds of Drosophila","volume":"99","author":"Williams","year":"1950","journal-title":"Biol. Bull."}],"container-title":["Journal of Experimental Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.1242\/jeb.214965","content-type":"unspecified","content-version":"vor","intended-application":"syndication"},{"URL":"http:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.214965\/1978721\/jeb214965.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"http:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.214965\/2067818\/jeb_214965v1.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"http:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.214965\/2067818\/jeb_214965v1.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,5,18]],"date-time":"2021-05-18T22:13:35Z","timestamp":1621376015000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/jeb\/article\/doi\/10.1242\/jeb.214965\/267724\/Humming-hummingbirds-insect-flight-tones-and-a"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,1,1]]},"references-count":57,"URL":"https:\/\/doi.org\/10.1242\/jeb.214965","relation":{},"ISSN":["1477-9145","0022-0949"],"issn-type":[{"value":"1477-9145","type":"electronic"},{"value":"0022-0949","type":"print"}],"subject":[],"published":{"date-parts":[[2020,1,1]]},"article-number":"jeb.214965"}}