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In Diptera, the forewings power flight, whereas the hindwings have evolved into specialized structures called halteres, which provide rapid mechanosensory feedback for flight stabilization. Although actuated by independent muscles, wing and haltere motion is precisely phase-coordinated at high frequencies. Because wingbeat frequency is a product of wing-thorax resonance, any wear-and-tear of wings or thorax should impair flight ability. How robust is the Dipteran flight system against such perturbations? Here, we show that wings and halteres are independently driven, coupled oscillators. We systematically reduced the wing length in flies and observed how wing-haltere synchronization was affected. The wing-wing system is a strongly coupled oscillator, whereas the wing-haltere system is weakly coupled through mechanical linkages that synchronize phase and frequency. Wing-haltere link acts in a unidirectional manner; altering wingbeat frequency affects haltere frequency, but not vice versa. Exoskeletal linkages are thus key morphological features of the Dipteran thorax that ensure wing-haltere synchrony, despite severe wing damage.<\/jats:p>","DOI":"10.7554\/elife.53824","type":"journal-article","created":{"date-parts":[[2021,11,12]],"date-time":"2021-11-12T10:15:14Z","timestamp":1636712114000},"update-policy":"https:\/\/doi.org\/10.7554\/elife.53824","source":"Crossref","is-referenced-by-count":23,"title":["Wings and halteres act as coupled dual oscillators in flies"],"prefix":"10.7554","volume":"10","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4019-2882","authenticated-orcid":true,"given":"Tanvi","family":"Deora","sequence":"first","affiliation":[{"name":"Department of Biology, University of Washington","place":["Seattle, Washington, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Siddharth S","family":"Sane","sequence":"additional","affiliation":[{"name":"Department of Physics, Azim Premji University","place":["Bangalore, India"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8274-1181","authenticated-orcid":true,"given":"Sanjay P","family":"Sane","sequence":"additional","affiliation":[{"name":"National Centre for Biological Sciences, Tata Institute of Fundamental Research","place":["Bangalore, India"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"4374","published-online":{"date-parts":[[2021,11,16]]},"reference":[{"key":"bib1","doi-asserted-by":"publisher","first-page":"4597","DOI":"10.1242\/jeb.02583","article-title":"A comparison of visual and haltere-mediated feedback in the control of body saccades in Drosophila melanogaster","volume":"209","author":"Bender","year":"2006","journal-title":"The Journal of Experimental Biology"},{"key":"bib2","doi-asserted-by":"publisher","first-page":"200","DOI":"10.2307\/1538368","article-title":"The mechanics of flight movements in diptera","volume":"102","author":"Boettiger","year":"1952","journal-title":"The Biological Bulletin"},{"key":"bib3","doi-asserted-by":"publisher","DOI":"10.2307\/5525","article-title":"Morphological Senescence and Longevity: An Experiment Relating Wing Wear and Life Span in Foraging Wild Bumble Bees","volume":"61","author":"Cartar","year":"1992","journal-title":"The Journal of Animal Ecology"},{"key":"bib4","doi-asserted-by":"publisher","first-page":"405","DOI":"10.1002\/(SICI)1096-9861(19960603)369:3<405::AID-CNE6>3.0.CO;2-9","article-title":"Position-specific central projections of mechanosensory neurons on the haltere of the blow fly, Calliphora vicina","volume":"369","author":"Chan","year":"1996","journal-title":"The Journal of Comparative Neurology"},{"key":"bib5","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1126\/science.280.5361.289","article-title":"Visual input to the efferent control system of a fly\u2019s \u201cgyroscope.\u201d","volume":"280","author":"Chan","year":"1998","journal-title":"Science"},{"key":"bib6","doi-asserted-by":"publisher","first-page":"426","DOI":"10.1098\/rsbl.2009.0915","article-title":"Dynamics of animal movement in an ecological context: dragonfly wing damage reduces flight performance and predation success","volume":"6","author":"Combes","year":"2010","journal-title":"Biology Letters"},{"key":"bib7","doi-asserted-by":"publisher","first-page":"1481","DOI":"10.1073\/pnas.1412279112","article-title":"Biomechanical basis of wing and haltere coordination in flies","volume":"112","author":"Deora","year":"2015","journal-title":"PNAS"},{"key":"bib8","doi-asserted-by":"publisher","first-page":"1382","DOI":"10.1242\/jeb.128363","article-title":"Mechanics of the thorax in flies","volume":"220","author":"Deora","year":"2017","journal-title":"The Journal of Experimental Biology"},{"key":"bib9","article-title":"Coupleddualoscillators_DeoraSane&Sane","volume-title":"Software Heritage","author":"Deora","year":"2021","unstructured":"Deora T. 2021. 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