{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,17]],"date-time":"2026-04-17T03:38:11Z","timestamp":1776397091299,"version":"3.51.2"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1009575","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2022,3,4]],"date-time":"2022-03-04T00:00:00Z","timestamp":1646352000000}}],"reference-count":100,"publisher":"Public Library of Science (PLoS)","issue":"2","license":[{"start":{"date-parts":[[2022,2,22]],"date-time":"2022-02-22T00:00:00Z","timestamp":1645488000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000053","name":"National Eye Institute","doi-asserted-by":"publisher","award":["1T32-EY021462"],"award-info":[{"award-number":["1T32-EY021462"]}],"id":[{"id":"10.13039\/100000053","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000053","name":"National Eye Institute","doi-asserted-by":"publisher","award":["R01-EY05729"],"award-info":[{"award-number":["R01-EY05729"]}],"id":[{"id":"10.13039\/100000053","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000053","name":"National Eye Institute","doi-asserted-by":"publisher","award":["K99\/R00 EY028229"],"award-info":[{"award-number":["K99\/R00 EY028229"]}],"id":[{"id":"10.13039\/100000053","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>We examine the structure of the visual motion projected on the retina during natural locomotion in real world environments. Bipedal gait generates a complex, rhythmic pattern of head translation and rotation in space, so without gaze stabilization mechanisms such as the vestibular-ocular-reflex (VOR) a walker\u2019s visually specified heading would vary dramatically throughout the gait cycle. The act of fixation on stable points in the environment nulls image motion at the fovea, resulting in stable patterns of outflow on the retinae centered on the point of fixation. These outflowing patterns retain a higher order structure that is informative about the stabilized trajectory of the eye through space. We measure this structure by applying the curl and divergence operations on the retinal flow velocity vector fields and found features that may be valuable for the control of locomotion. In particular, the sign and magnitude of foveal curl in retinal flow specifies the body\u2019s trajectory relative to the gaze point, while the point of maximum divergence in the retinal flow field specifies the walker\u2019s instantaneous overground velocity\/momentum vector in retinotopic coordinates. Assuming that walkers can determine the body position relative to gaze direction, these time-varying retinotopic cues for the body\u2019s momentum could provide a visual control signal for locomotion over complex terrain. In contrast, the temporal variation of the eye-movement-free, head-centered flow fields is large enough to be problematic for use in steering towards a goal. Consideration of optic flow in the context of real-world locomotion therefore suggests a re-evaluation of the role of optic flow in the control of action during natural behavior.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1009575","type":"journal-article","created":{"date-parts":[[2022,2,22]],"date-time":"2022-02-22T13:45:55Z","timestamp":1645537555000},"page":"e1009575","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":86,"title":["Retinal optic flow during natural locomotion"],"prefix":"10.1371","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-3683-646X","authenticated-orcid":true,"given":"Jonathan Samir","family":"Matthis","sequence":"first","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1319-9293","authenticated-orcid":true,"given":"Karl S.","family":"Muller","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9210-8275","authenticated-orcid":true,"given":"Kathryn L.","family":"Bonnen","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6671-5207","authenticated-orcid":true,"given":"Mary M.","family":"Hayhoe","sequence":"additional","affiliation":[]}],"member":"340","published-online":{"date-parts":[[2022,2,22]]},"reference":[{"key":"pcbi.1009575.ref001","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1007\/7854_2018_60","volume-title":"Processes of Visuospatial Attention and Working Memory","author":"MF Land","year":"2018"},{"issue":"3","key":"pcbi.1009575.ref002","doi-asserted-by":"crossref","first-page":"819","DOI":"10.1113\/jphysiol.2004.061523","article-title":"Contribution of cone photoreceptors and post-receptoral mechanisms to the human photopic electroretinogram: Human cone ERG","volume":"556","author":"C Friedburg","year":"2004","journal-title":"The Journal of Physiology"},{"key":"pcbi.1009575.ref003","article-title":"Freely-moving mice visually pursue prey using a retinal area with least optic flow","author":"CD Holmgren","year":"2021","journal-title":"Neuroscience"},{"issue":"1-4","key":"pcbi.1009575.ref004","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1016\/0042-6989(62)90064-0","article-title":"The evolutionary history of eye movements","volume":"2","author":"GL Walls","year":"1962","journal-title":"Vision Research"},{"key":"pcbi.1009575.ref005","volume-title":"The perception of the visual world","author":"JJ Gibson","year":"1950"},{"key":"pcbi.1009575.ref006","volume-title":"The ecological approach to visual perception","author":"JJ Gibson","year":"1979"},{"key":"pcbi.1009575.ref007","first-page":"339","article-title":"Action Modes and Laws of Control for the Visual Guidance Of Action","volume-title":"Advances in Psychology","author":"WH Warren","year":"1988"},{"issue":"9","key":"pcbi.1009575.ref008","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1016\/S1364-6613(99)01364-9","article-title":"Perception of self-motion from visual flow","volume":"3","author":"M Lappe","year":"1999","journal-title":"Trends in Cognitive Sciences"},{"key":"pcbi.1009575.ref009","first-page":"243","volume-title":"Closing the gap: The scientific writings of David N. 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I. A continuum of response selectivity to large-field stimuli","volume":"65","author":"CJ Duffy","year":"1991","journal-title":"Journal of Neurophysiology"},{"issue":"12","key":"pcbi.1009575.ref015","doi-asserted-by":"crossref","first-page":"1322","DOI":"10.1038\/81860","article-title":"A cortical area that responds specifically to optic flow, revealed by fMRI","volume":"3","author":"MC Morrone","year":"2000","journal-title":"Nature Neuroscience"},{"issue":"3","key":"pcbi.1009575.ref016","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1016\/j.cub.2007.12.053","article-title":"The Representation of Egomotion in the Human Brain","volume":"18","author":"MB Wall","year":"2008","journal-title":"Current Biology"},{"issue":"1","key":"pcbi.1009575.ref017","doi-asserted-by":"crossref","first-page":"920","DOI":"10.1038\/s41467-017-01021-5","article-title":"Heading representations in primates are compressed by saccades","volume":"8","author":"F Bremmer","year":"2017","journal-title":"Nature Communications"},{"issue":"10","key":"pcbi.1009575.ref018","doi-asserted-by":"crossref","first-page":"2470","DOI":"10.1152\/jn.00410.2014","article-title":"Visual selectivity for heading in the macaque ventral intraparietal area","volume":"112","author":"A Kaminiarz","year":"2014","journal-title":"Journal of Neurophysiology"},{"issue":"1","key":"pcbi.1009575.ref019","doi-asserted-by":"crossref","first-page":"160","DOI":"10.1364\/JOSAA.7.000160","article-title":"Eye movements and optical flow","volume":"7","author":"WH Warren","year":"1990","journal-title":"Journal of the Optical Society of America A"},{"key":"pcbi.1009575.ref020","first-page":"13","article-title":"The interpretation of a moving retinal image","volume":"208","author":"HC Longuet-Higgins","year":"1980","journal-title":"Proceedings of the Royal Society B: Biological 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Nakamura","year":"2016","journal-title":"i-Perception"},{"issue":"1","key":"pcbi.1009575.ref033","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1068\/p2990","article-title":"Global-Perspective Jitter Improves Vection in Central Vision","volume":"29","author":"S Palmisano","year":"2000","journal-title":"Perception"},{"issue":"8","key":"pcbi.1009575.ref034","doi-asserted-by":"crossref","first-page":"1224","DOI":"10.1016\/j.cub.2018.03.008","article-title":"Gaze and the Control of Foot Placement When Walking in Natural Terrain","volume":"28","author":"JS Matthis","year":"2018","journal-title":"Current Biology"},{"issue":"2","key":"pcbi.1009575.ref035","doi-asserted-by":"crossref","first-page":"233","DOI":"10.1007\/s00221-012-3254-x","article-title":"Mind the step: complementary effects of an implicit task on eye and head movements in real-life gaze allocation","volume":"223","author":"BM \u2019t Hart","year":"2012","journal-title":"Experimental Brain Research"},{"issue":"3","key":"pcbi.1009575.ref036","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1080\/09548980701671094","article-title":"Human eye-head co-ordination in natural exploration","volume":"18","author":"W Einh\u00e4user","year":"2007","journal-title":"Network: Computation in Neural Systems"},{"key":"pcbi.1009575.ref037","first-page":"32","article-title":"Wayfinding on Foot From Information in Retinal, Not Optical, Flow","author":"JE Cutting","year":"1992","journal-title":"Journal of Experimental Psychology: General"},{"issue":"5","key":"pcbi.1009575.ref038","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1007\/BF00216964","article-title":"On the sufficiency of the velocity field for perception of heading","volume":"65","author":"WH Warren","year":"1991","journal-title":"Biological Cybernetics"},{"issue":"52","key":"pcbi.1009575.ref039","doi-asserted-by":"crossref","first-page":"33161","DOI":"10.1073\/pnas.2022984117","article-title":"Heading 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Speed","volume":"408","author":"H Dietrich","year":"2019","journal-title":"Neuroscience"},{"key":"pcbi.1009575.ref043","doi-asserted-by":"crossref","first-page":"4","DOI":"10.3389\/fnsys.2018.00004","article-title":"The Interaction of Pre-programmed Eye Movements With the Vestibulo-Ocular Reflex","volume":"12","author":"SE Haggerty","year":"2018","journal-title":"Frontiers in Systems Neuroscience"},{"issue":"7","key":"pcbi.1009575.ref044","doi-asserted-by":"crossref","first-page":"e40264","DOI":"10.1371\/journal.pone.0040264","article-title":"Vestibular Facilitation of Optic Flow Parsing","volume":"7","author":"PR MacNeilage","year":"2012","journal-title":"PLoS ONE"},{"issue":"2","key":"pcbi.1009575.ref045","doi-asserted-by":"crossref","first-page":"730","DOI":"10.1152\/jn.2000.84.2.730","article-title":"Responses to Continuously Changing Optic Flow in Area MST","volume":"84","author":"M Paolini","year":"2000","journal-title":"Journal of Neurophysiology"},{"issue":"6404","key":"pcbi.1009575.ref046","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1038\/360583a0","article-title":"The perception of heading during eye movements","volume":"360","author":"CS Royden","year":"1992","journal-title":"Nature"},{"key":"pcbi.1009575.ref047","doi-asserted-by":"crossref","unstructured":"Weinzaepfel P, Revaud J, Harchaoui Z, Schmid C. DeepFlow: Large Displacement Optical Flow with Deep Matching. In: 2013 IEEE International Conference on Computer Vision. Sydney, Australia: IEEE; 2013. p. 1385\u20131392. Available from: http:\/\/ieeexplore.ieee.org\/document\/6751282\/.","DOI":"10.1109\/ICCV.2013.175"},{"issue":"1762","key":"pcbi.1009575.ref048","doi-asserted-by":"crossref","first-page":"20130700","DOI":"10.1098\/rspb.2013.0700","article-title":"Humans exploit the biomechanics of bipedal gait during visually guided walking over complex terrain","volume":"280","author":"JS Matthis","year":"2013","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"issue":"1480","key":"pcbi.1009575.ref049","doi-asserted-by":"crossref","first-page":"1985","DOI":"10.1098\/rspb.2001.1761","article-title":"Mechanical and metabolic determinants of the preferred step width in human walking","volume":"268","author":"M Donelan","year":"2001","journal-title":"Proceedings of the Royal Society of London Series B: Biological Sciences"},{"issue":"4","key":"pcbi.1009575.ref050","doi-asserted-by":"crossref","first-page":"617","DOI":"10.1016\/j.humov.2007.04.003","article-title":"The six 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Glennerster","year":"2001","journal-title":"Vision Research"},{"key":"pcbi.1009575.ref054","doi-asserted-by":"crossref","first-page":"1271","DOI":"10.3389\/fphys.2018.01271","article-title":"Neural Control of Balance During Walking","volume":"9","author":"H Reimann","year":"2018","journal-title":"Frontiers in Physiology"},{"issue":"2","key":"pcbi.1009575.ref055","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1038\/nn.4197","article-title":"Thalamic nuclei convey diverse contextual information to layer 1 of visual cortex","volume":"19","author":"MM Roth","year":"2016","journal-title":"Nature Neuroscience"},{"issue":"5","key":"pcbi.1009575.ref056","doi-asserted-by":"crossref","first-page":"969","DOI":"10.1152\/jn.00513.2013","article-title":"A novel optic flow pattern speeds split-belt locomotor adaptation","volume":"111","author":"JM Finley","year":"2014","journal-title":"Journal of 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Collecting and Analyzing Eye-tracking Data in Outdoor Environments. R A. 2012; p. 19.","DOI":"10.16910\/jemr.5.2.6"},{"key":"pcbi.1009575.ref097","volume-title":"Software that sees","author":"GR Bradski","year":"2011","edition":"1"},{"key":"pcbi.1009575.ref098","unstructured":"Legland D. geom3d; 2020. 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