{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T01:03:35Z","timestamp":1790125415992,"version":"4.0.1"},"reference-count":40,"publisher":"The Royal Society","issue":"242","license":[{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"NWO Vidi","award":["VI.Vidi.193.054"],"award-info":[{"award-number":["VI.Vidi.193.054"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,23]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Landing on a moving target is challenging for flying animals, particularly under dynamic outdoor conditions. For pollinators such as bumblebees, wind-induced flower motion adds substantial complexity to landing manoeuvres. Here, we investigate how bumblebees (Bombus terrestris) land on laterally oscillating flowers. Using high-speed stereoscopic videography, we quantified three-dimensional flight kinematics of 48 bumblebees approaching a sinusoidally moving flower-mimic and analysed their control strategies using system-identification and control-theoretic modelling. Landing dynamics were consistent with combined visual (optic-flow) and angular yaw-rate feedback, enabling precise sideways tracking and gaze stabilization. Sideways motion was captured by a proportional controller, whereas gaze control required a proportional\u2013derivative structure. Both controllers yielded similar case-specific effective delays (approximately 54 and 52\u00a0ms). Surprisingly, optic-target-angle contributed negligibly to gaze control, indicating reliance on velocity-based rather than position-based cues. Lateral acceleration correlated strongly with body roll, consistent with the helicopter model of force redirection. These results reveal a modular visual\u2013inertial control strategy enabling robust landings on moving targets and suggest principles for agile bio-inspired flight.<\/jats:p>","DOI":"10.1098\/rsif.2025.1198","type":"journal-article","created":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:13:06Z","timestamp":1790122386000},"source":"Crossref","is-referenced-by-count":0,"title":["Visual\u2013inertial integration enables bumblebees to land on rapidly moving flowers"],"prefix":"10.1098","volume":"23","author":[{"ORCID":"https:\/\/orcid.org\/0009-0001-1493-5355","authenticated-orcid":false,"given":"Chenyao","family":"Wang","sequence":"first","affiliation":[{"name":"Experimental Zoology Group, Wageningen University 1 , Wageningen, The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lana J.","family":"de Vries","sequence":"additional","affiliation":[{"name":"Experimental Zoology Group, Wageningen University 1 , Wageningen, The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vincent","family":"van de Pas","sequence":"additional","affiliation":[{"name":"Experimental Zoology Group, Wageningen University 1 , Wageningen, The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Steffen","family":"Werner","sequence":"additional","affiliation":[{"name":"Experimental Zoology Group, Wageningen University 1 , Wageningen, The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christophe","family":"De Wagter","sequence":"additional","affiliation":[{"name":"Micro Air Vehicle Laboratory, Delft University of Technology 2 , Delft, The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5668-0653","authenticated-orcid":false,"given":"Florian T.","family":"Muijres","sequence":"additional","affiliation":[{"name":"Experimental Zoology Group, Wageningen University 1 , Wageningen, The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"175","published-online":{"date-parts":[[2026,9,23]]},"reference":[{"key":"2026092220114752700_RSIF20251198C1","doi-asserted-by":"crossref","first-page":"jeb249380","DOI":"10.1242\/jeb.249380","article-title":"Follow the flower: approach-flight behaviour of bumblebees landing on a moving target","volume":"228","author":"de Vries","year":"2025","journal-title":"J. Exp. Biol."},{"key":"2026092220114752700_RSIF20251198C2","doi-asserted-by":"crossref","first-page":"20203051","DOI":"10.1098\/rspb.2020.3051","article-title":"The effect of optic flow cues on honeybee flight control in wind","volume":"288","author":"Baird","year":"2021","journal-title":"Proc. R. Soc. B"},{"key":"2026092220114752700_RSIF20251198C3","doi-asserted-by":"crossref","first-page":"18 686","DOI":"10.1073\/pnas.1314311110","article-title":"A universal strategy for visually guided landing","volume":"110","author":"Baird","year":"2013","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026092220114752700_RSIF20251198C4","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1007\/s00359-022-01610-w","article-title":"Optic flow based spatial vision in insects","volume":"209","author":"Egelhaaf","year":"2023","journal-title":"J. Comp. Physiol. A"},{"key":"2026092220114752700_RSIF20251198C5","doi-asserted-by":"crossref","first-page":"102467","DOI":"10.1016\/j.isci.2021.102407","article-title":"Bumblebees land rapidly and robustly using a sophisticated modular flight control strategy","volume":"24","author":"Goyal","year":"2021","journal-title":"iScience"},{"key":"2026092220114752700_RSIF20251198C6","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1017\/S0952523800004582","article-title":"Visual tracking of moving targets by freely flying honeybees","volume":"4","author":"Zhang","year":"1990","journal-title":"Vis. Neurosci."},{"key":"2026092220114752700_RSIF20251198C7","volume-title":"The perception of the visual world","author":"Gibson","year":"1950","edition":"1st"},{"key":"2026092220114752700_RSIF20251198C8","doi-asserted-by":"crossref","first-page":"3895","DOI":"10.1242\/jeb.01818","article-title":"Visual control of flight speed in honeybees","volume":"208","author":"Baird","year":"2005","journal-title":"J. Exp. Biol."},{"key":"2026092220114752700_RSIF20251198C9","doi-asserted-by":"crossref","first-page":"016004","DOI":"10.1088\/1748-3190\/11\/1\/016004","article-title":"Monocular distance estimation with optical flow maneuvers and efference copies: a stability-based strategy","volume":"11","author":"de Croon","year":"2016","journal-title":"Bioinspir. Biomim."},{"key":"2026092220114752700_RSIF20251198C10","doi-asserted-by":"crossref","first-page":"4608","DOI":"10.1016\/j.cub.2021.09.068","article-title":"Suppression of motion vision during course-changing, but not course-stabilizing, navigational turns","volume":"31","author":"Fenk","year":"2021","journal-title":"Curr. Biol."},{"key":"2026092220114752700_RSIF20251198C11","doi-asserted-by":"crossref","first-page":"4743","DOI":"10.1016\/j.cub.2024.08.036","article-title":"Divergent visual ecology of Drosophila species drives object-tracking strategies matched to landscape sparsity","volume":"34","author":"Rimniceanu","year":"2024","journal-title":"Curr. Biol."},{"key":"2026092220114752700_RSIF20251198C12","doi-asserted-by":"crossref","first-page":"2901","DOI":"10.1016\/j.cub.2017.08.035","article-title":"Drosophila spatiotemporally integrates visual signals to control saccades","volume":"27","author":"Mongeau","year":"2017","journal-title":"Curr. Biol."},{"key":"2026092220114752700_RSIF20251198C13","doi-asserted-by":"crossref","first-page":"jeb246106","DOI":"10.1242\/jeb.245432","article-title":"Bumblebees compensate for the adverse effects of sidewind during visually guided landings","volume":"227","author":"Goyal","year":"2024","journal-title":"J. Exp. Biol."},{"key":"2026092220114752700_RSIF20251198C14","doi-asserted-by":"crossref","first-page":"E1182","DOI":"10.1073\/pnas.1323529111","article-title":"Flying Drosophila stabilize their vision-based velocity controller by sensing wind with their antennae","volume":"111","author":"Fuller","year":"2014","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026092220114752700_RSIF20251198C15","first-page":"2301","article-title":"Control of moth flight posture is mediated by wing mechanosensory feedback","volume":"217","author":"Dickerson","year":"2014","journal-title":"J. Exp. Biol."},{"key":"2026092220114752700_RSIF20251198C16","first-page":"347","article-title":"The gyroscopic mechanism of the halteres of Diptera","volume":"233","author":"Pringle","year":"1948","journal-title":"Phil. Trans. R. Soc. B"},{"key":"2026092220114752700_RSIF20251198C17","doi-asserted-by":"crossref","first-page":"20141088","DOI":"10.1098\/rsif.2014.1088","article-title":"A new twist on gyroscopic sensing: body rotations lead to torsion in flapping, flexing insect wings","volume":"12","author":"Eberle","year":"2015","journal-title":"J. R. Soc. Interface"},{"key":"2026092220114752700_RSIF20251198C18","first-page":"385","article-title":"The interpretation of a moving retinal image","volume":"208","author":"Longuet-Higgins","year":"1980","journal-title":"Proc. R. Soc. B"},{"key":"2026092220114752700_RSIF20251198C19","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1098\/rsif.2010.0225","article-title":"Behavioural system identification of visual flight speed control in Drosophila melanogaster","volume":"8","author":"Rohrseitz","year":"2011","journal-title":"J. R. Soc. Interface"},{"key":"2026092220114752700_RSIF20251198C20","doi-asserted-by":"crossref","first-page":"e2121660119","DOI":"10.1073\/pnas.2121660119","article-title":"Complementary feedback control enables effective gaze stabilization in animals","volume":"119","author":"Cellini","year":"2022","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026092220114752700_RSIF20251198C21","doi-asserted-by":"crossref","first-page":"21-00113","DOI":"10.1299\/jbse.21-00113","article-title":"A six-degree-of-freedom proportional-derivative control strategy for bumblebee flight stabilization","volume":"16","author":"Cai","year":"2021","journal-title":"J. Biomech. Sci. Eng."},{"key":"2026092220114752700_RSIF20251198C22","doi-asserted-by":"crossref","first-page":"1060933","DOI":"10.3389\/frobt.2022.1060933","article-title":"Embodied airflow sensing for improved in-gust flight of flapping wing MAVs","volume":"9","author":"Wang","year":"2022","journal-title":"Front. Robot. AI"},{"key":"2026092220114752700_RSIF20251198C23","doi-asserted-by":"crossref","DOI":"10.2514\/6.2006-34","article-title":"An integrative model of insect flight control","author":"Dickson","year":"2006","journal-title":"44th AIAA Aerospace Sciences Meeting and Exhibit"},{"key":"2026092220114752700_RSIF20251198C24","doi-asserted-by":"crossref","first-page":"4820","DOI":"10.1073\/pnas.1000615107","article-title":"Discovering the flight autostabilizer of fruit flies by inducing aerial stumbles","volume":"107","author":"Ristroph","year":"2010","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026092220114752700_RSIF20251198C25","doi-asserted-by":"crossref","first-page":"849","DOI":"10.1109\/TRO.2011.2156170","article-title":"Translational and rotational damping of flapping flight and its dynamics and stability at hovering","volume":"27","author":"Cheng","year":"2011","journal-title":"IEEE Trans. Robot."},{"key":"2026092220114752700_RSIF20251198C26","first-page":"3508","article-title":"Pitch perfect: how fruit flies control their body pitch angle","volume":"218","author":"Whitehead","year":"2015","journal-title":"J. Exp. Biol."},{"key":"2026092220114752700_RSIF20251198C27","doi-asserted-by":"crossref","first-page":"20150075","DOI":"10.1098\/rsif.2015.0075","article-title":"Controlling roll perturbations in fruit flies","volume":"12","author":"Beatus","year":"2015","journal-title":"J. R. Soc. Interface"},{"key":"2026092220114752700_RSIF20251198C28","doi-asserted-by":"crossref","first-page":"1245","DOI":"10.1126\/science.aaa3042","article-title":"Luminance-dependent visual processing enables moth flight in low light","volume":"348","author":"Sponberg","year":"2015","journal-title":"Science"},{"key":"2026092220114752700_RSIF20251198C29","volume-title":"Feedback systems: an introduction for scientists and engineers","author":"\u00c5str\u00f6m","year":"2021","edition":"2nd"},{"key":"2026092220114752700_RSIF20251198C30","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1242\/jeb.01957","article-title":"Induced airflow in flying insects I. A theoretical model of the induced flow","volume":"209","author":"Sane","year":"2006","journal-title":"J. Exp. Biol."},{"key":"2026092220114752700_RSIF20251198C31","doi-asserted-by":"crossref","first-page":"1281","DOI":"10.1038\/s41593-018-0209-y","article-title":"DeepLabCut: markerless pose estimation of user-defined body parts with deep learning","volume":"21","author":"Mathis","year":"2018","journal-title":"Nat. Neurosci."},{"key":"2026092220114752700_RSIF20251198C32","first-page":"1843","article-title":"A protocol and calibration method for accurate multi-camera field videography","volume":"217","author":"Theriault","year":"2014","journal-title":"J. Exp. Biol."},{"key":"2026092220114752700_RSIF20251198C33","article-title":"A Python package to track the 3D flight kinematic of insects with DeepLabCut","author":"Cribellier","year":"2024","journal-title":"SEB Conf. Prague 2024, Prague, Czech Republic, 2\u20135 July 2024"},{"key":"2026092220114752700_RSIF20251198C34","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1016\/0042-6989(86)90078-7","article-title":"Optic flow","volume":"26","author":"Koenderink","year":"1986","journal-title":"Vision. Res."},{"key":"2026092220114752700_RSIF20251198C35","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1126\/science.1248955","article-title":"Flies evade looming targets by executing rapid visually directed banked turns","volume":"344","author":"Muijres","year":"2014","journal-title":"Science"},{"key":"2026092220114752700_RSIF20251198C36","doi-asserted-by":"crossref","first-page":"jeb230920","DOI":"10.1242\/jeb.230920","article-title":"Hovering flight in hummingbird hawkmoths: kinematics, wake dynamics and aerodynamic power","volume":"224","author":"Warfvinge","year":"2021","journal-title":"J. Exp. Biol."},{"key":"2026092220114752700_RSIF20251198C37","doi-asserted-by":"crossref","first-page":"1643","DOI":"10.1242\/jeb.041426","article-title":"The spatial frequency tuning of optic-flow-dependent behaviors in the bumblebee Bombus impatiens","volume":"213","author":"Dyhr","year":"2010","journal-title":"J. Exp. Biol."},{"key":"2026092220114752700_RSIF20251198C38","doi-asserted-by":"crossref","first-page":"1625","DOI":"10.1242\/jeb.038802","article-title":"Minimum viewing angle for visually guided ground speed control in bumblebees","volume":"213","author":"Baird","year":"2010","journal-title":"J. Exp. Biol."},{"key":"2026092220114752700_RSIF20251198C39","article-title":"Data and code from: Visual-inertial integration enables bumblebees to land on rapidly moving flowers [Dataset]","author":"Wang","year":"2026"},{"key":"2026092220114752700_RSIF20251198C40","article-title":"Supplementary material from: Visual-inertial integration enables bumblebees to land on rapidly moving flowers","author":"Wang","year":"2026"}],"container-title":["Journal of the Royal Society Interface"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/royalsocietypublishing.org\/rsif\/article-pdf\/doi\/10.1098\/rsif.2025.1198\/6167060\/rsif.2025.1198.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/royalsocietypublishing.org\/rsif\/article-pdf\/doi\/10.1098\/rsif.2025.1198\/6167060\/rsif.2025.1198.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:13:11Z","timestamp":1790122391000},"score":1,"resource":{"primary":{"URL":"https:\/\/royalsocietypublishing.org\/rsif\/article\/23\/242\/20251198\/483371\/Visual-inertial-integration-enables-bumblebees-to"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,23]]},"references-count":40,"journal-issue":{"issue":"242","published-print":{"date-parts":[[2026,9,23]]}},"URL":"https:\/\/doi.org\/10.1098\/rsif.2025.1198","relation":{},"ISSN":["1742-5689","1742-5662"],"issn-type":[{"value":"1742-5689","type":"print"},{"value":"1742-5662","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,23]]},"article-number":"20251198"}}