{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T12:27:13Z","timestamp":1785414433112,"version":"3.56.0"},"posted":{"date-parts":[[2026,6,10]]},"group-title":"Neuroscience","reference-count":95,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2026,6,10]],"date-time":"2026-06-10T00:00:00Z","timestamp":1781049600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"award":["NIH R01 EY026555"],"award-info":[{"award-number":["NIH R01 EY026555"]}],"id":[{"id":"https:\/\/ror.org\/001d55x84","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2026,6,10]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                <jats:p>Detecting approaching objects is essential for survival, and studies across animals have identified neurons and circuits selective for objects that grow in size over time, a hallmark of visual approach. However, approach as a physical process generates a rich ensemble of correlated cues governed by geometry and physics, of which size expansion is just one, and how the brain incorporates this broader cue structure to detect approach remains unknown. Here, we show that a visual stimulus without size expansion but with modulated luminance, a cue intrinsic to approach, elicits compelling percepts of approach and retreat in both humans and the fruit fly Drosophila, revealing an unsuspected, cross-species percept of approach based on luminance alone. Using targeted genetic silencing and two-photon calcium imaging in flies, we identify neurons that mediate evasive responses to both expansion-based and luminance-based approach, establishing them as general approach detectors. Guided by the geometry of approach, we show that luminance cues should precede detectable expansion signals during naturalistic approach. Consistent with this, these neurons combine luminance and expansion cues synergistically, but only when luminance changes precede expansion, matching their natural temporal order, thus performing sequenced cue integration. Together, these findings anchor a cross-species perceptual phenomenon to a defined circuit computation and reveal how neural circuits are tuned to the dynamic cue structure of natural events.<\/jats:p>","DOI":"10.64898\/2026.06.05.730527","type":"posted-content","created":{"date-parts":[[2026,6,10]],"date-time":"2026-06-10T18:55:12Z","timestamp":1781117712000},"source":"Crossref","is-referenced-by-count":0,"title":["Dynamic trajectory cues drive sequenced integration in approach detectors"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9411-5137","authenticated-orcid":false,"given":"Harsh","family":"Vashistha","sequence":"first","affiliation":[{"name":"Department of Molecular, Cellular and Developmental Biology, Yale University"},{"name":"Quantitative Biology Institute, Yale University"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Natalia CB","family":"matos","sequence":"additional","affiliation":[{"name":"Quantitative Biology Institute, Yale University"},{"name":"Interdepartmental Neuroscience Program, Yale University"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Heng","family":"Wu","sequence":"additional","affiliation":[{"name":"Department of Molecular, Cellular and Developmental Biology, Yale University"},{"name":"Quantitative Biology Institute, Yale University"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8487-700X","authenticated-orcid":false,"given":"Damon","family":"Clark","sequence":"additional","affiliation":[{"name":"Department of Molecular, Cellular and Developmental Biology, Yale University"},{"name":"Quantitative Biology Institute, Yale University"},{"name":"Interdepartmental Neuroscience Program, Yale University"},{"name":"Department of Physics, Yale University"},{"name":"Department of Neuroscience, Yale University"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026061320001234000_2026.06.05.730527v1.1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2019.01.079"},{"key":"2026061320001234000_2026.06.05.730527v1.2","doi-asserted-by":"publisher","DOI":"10.1364\/JOSAA.2.000284"},{"key":"2026061320001234000_2026.06.05.730527v1.3","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2002937117"},{"key":"2026061320001234000_2026.06.05.730527v1.4","doi-asserted-by":"publisher","DOI":"10.1037\/h0068517"},{"key":"2026061320001234000_2026.06.05.730527v1.5","doi-asserted-by":"publisher","DOI":"10.1126\/science.171.3973.818"},{"key":"2026061320001234000_2026.06.05.730527v1.6","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1965.sp007638"},{"key":"2026061320001234000_2026.06.05.730527v1.7","doi-asserted-by":"publisher","DOI":"10.1016\/0166-2236(89)90010-6"},{"key":"2026061320001234000_2026.06.05.730527v1.8","doi-asserted-by":"publisher","DOI":"10.1038\/373609a0"},{"key":"2026061320001234000_2026.06.05.730527v1.9","doi-asserted-by":"publisher","DOI":"10.1163\/156856897X00357"},{"key":"2026061320001234000_2026.06.05.730527v1.10","doi-asserted-by":"publisher","DOI":"10.1007\/s00359-018-1257-1"},{"key":"2026061320001234000_2026.06.05.730527v1.11","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2008.07.094"},{"key":"2026061320001234000_2026.06.05.730527v1.12","doi-asserted-by":"crossref","unstructured":"J. Chen , H. Mandel , J. Fitzgerald , and D. Clark . Asymmetric on-off processing of visual motion cancels variability induced by the structure of natural scenes, 2019.","DOI":"10.7554\/eLife.47579.sa2"},{"key":"2026061320001234000_2026.06.05.730527v1.13","doi-asserted-by":"publisher","DOI":"10.1038\/nature12354"},{"key":"2026061320001234000_2026.06.05.730527v1.14","doi-asserted-by":"publisher","DOI":"10.1038\/nn.3600"},{"key":"2026061320001234000_2026.06.05.730527v1.15","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1306428110"},{"key":"2026061320001234000_2026.06.05.730527v1.16","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1336-14.2014"},{"key":"2026061320001234000_2026.06.05.730527v1.17","doi-asserted-by":"publisher","DOI":"10.1037\/h0044343"},{"key":"2026061320001234000_2026.06.05.730527v1.18","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-024-07451-8"},{"key":"2026061320001234000_2026.06.05.730527v1.19","doi-asserted-by":"publisher","DOI":"10.1016\/j.jneumeth.2019.05.005"},{"key":"2026061320001234000_2026.06.05.730527v1.20","doi-asserted-by":"crossref","unstructured":"J. E. Cutting and P. M. Vishton . Perceiving Layout and Knowing Distances: The Integration, Relative Potency, and Contextual Use of Different Information about Depth. Perception of Space and Motion, pages 69\u2013117, 1 1995.","DOI":"10.1016\/B978-012240530-3\/50005-5"},{"key":"2026061320001234000_2026.06.05.730527v1.21","doi-asserted-by":"crossref","unstructured":"R. Dewell , Y. Zhu , M. Eisenbrandt , R. Morse , and F. Gabbiani . Contrast-polarity specific mapping improves efficiency of neuronal computation for collision detection, 2022.","DOI":"10.7554\/eLife.79772.sa2"},{"key":"2026061320001234000_2026.06.05.730527v1.22","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-022-05562-8"},{"key":"2026061320001234000_2026.06.05.730527v1.23","doi-asserted-by":"publisher","DOI":"10.1016\/0042-6989(86)90154-9"},{"key":"2026061320001234000_2026.06.05.730527v1.24","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2015.12.021"},{"key":"2026061320001234000_2026.06.05.730527v1.25","doi-asserted-by":"publisher","DOI":"10.1068\/p120167"},{"key":"2026061320001234000_2026.06.05.730527v1.26","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-018-0244-6"},{"key":"2026061320001234000_2026.06.05.730527v1.27","doi-asserted-by":"publisher","DOI":"10.1068\/p060287"},{"issue":"1","key":"2026061320001234000_2026.06.05.730527v1.28","first-page":"1","article-title":"Behavioral and neuronal underpinnings of safety in numbers in fruit flies","volume":"11","year":"2020","journal-title":"Nature Communications"},{"key":"2026061320001234000_2026.06.05.730527v1.29","doi-asserted-by":"crossref","unstructured":"J. Fitzgerald and D. Clark . Nonlinear circuits for naturalistic visual motion estimation. eLife, page 4, 2015.","DOI":"10.7554\/eLife.09123"},{"key":"2026061320001234000_2026.06.05.730527v1.30","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1015680108"},{"key":"2026061320001234000_2026.06.05.730527v1.31","doi-asserted-by":"crossref","unstructured":"H. Fotowat and F. Engert . Neural circuits underlying habituation of visually evoked escape behaviors in larval zebrafish. ELife, 12, 2023.","DOI":"10.7554\/eLife.82916"},{"key":"2026061320001234000_2026.06.05.730527v1.32","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-neuro-061010-113632"},{"key":"2026061320001234000_2026.06.05.730527v1.33","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2016.06.006"},{"key":"2026061320001234000_2026.06.05.730527v1.34","doi-asserted-by":"publisher","DOI":"10.1038\/281565a0"},{"key":"2026061320001234000_2026.06.05.730527v1.35","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.52760"},{"key":"2026061320001234000_2026.06.05.730527v1.36","doi-asserted-by":"publisher","DOI":"10.1038\/nature01190"},{"issue":"3","key":"2026061320001234000_2026.06.05.730527v1.37","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1038\/nmeth.1561","article-title":"A versatile in vivo system for directed dissection of gene expression patterns","volume":"8","year":"2011","journal-title":"Nature Methods"},{"key":"2026061320001234000_2026.06.05.730527v1.38","doi-asserted-by":"crossref","unstructured":"S. Hampel , R. Franconville , J. Simpson , and A. Seeds . A neural command circuit for grooming movement control, 2015.","DOI":"10.7554\/eLife.08758.036"},{"key":"2026061320001234000_2026.06.05.730527v1.39","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1515\/znb-1956-9-1004","article-title":"Systemtheoretische analyse der zeit-, reihenfolgen-und vorzeichenauswertung bei der bewegungsperzeption des r\u00fcsselk\u00e4fers chlorophanus","volume":"11","year":"1956","journal-title":"Zeitschrift fur Naturforschung - Section B Journal of Chemical Sciences"},{"issue":"1","key":"2026061320001234000_2026.06.05.730527v1.40","doi-asserted-by":"crossref","first-page":"21578","DOI":"10.1038\/s41598-024-70256-2","article-title":"Looming stimuli reliably drive innate defensive responses in male rats, but not learned defensive responses","volume":"14","year":"2024","journal-title":"Scientific Reports"},{"key":"2026061320001234000_2026.06.05.730527v1.41","doi-asserted-by":"crossref","unstructured":"P. Hibbard , R. Goutcher , R. Hornsey , D. Hunter , and P. Scarfe . Luminance contrast provides metric depth information, volume 10. Royal Society Open Science, 2023.","DOI":"10.1098\/rsos.220567"},{"key":"2026061320001234000_2026.06.05.730527v1.42","doi-asserted-by":"publisher","DOI":"10.1167\/10.1.9"},{"key":"2026061320001234000_2026.06.05.730527v1.43","doi-asserted-by":"publisher","DOI":"10.1038\/nn1497"},{"key":"2026061320001234000_2026.06.05.730527v1.44","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2010.10.025"},{"key":"2026061320001234000_2026.06.05.730527v1.45","doi-asserted-by":"publisher","DOI":"10.1093\/cercor\/bhm187"},{"key":"2026061320001234000_2026.06.05.730527v1.46","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2017.01.012"},{"key":"2026061320001234000_2026.06.05.730527v1.47","doi-asserted-by":"crossref","first-page":"2115","DOI":"10.1016\/j.celrep.2020.01.061","article-title":"Inhibitory interactions and columnar inputs to an object motion detector in drosophila","volume":"30","year":"2020","journal-title":"Cell Reports"},{"key":"2026061320001234000_2026.06.05.730527v1.48","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2022.12.014"},{"key":"2026061320001234000_2026.06.05.730527v1.49","doi-asserted-by":"crossref","first-page":"9920","DOI":"10.1126\/sciadv.abc9920","article-title":"Dendritic and parallel processing of visual threats in the retina control defensive responses","volume":"6","year":"2020","journal-title":"Science Advances"},{"key":"2026061320001234000_2026.06.05.730527v1.50","doi-asserted-by":"publisher","DOI":"10.1002\/neu.1018"},{"key":"2026061320001234000_2026.06.05.730527v1.51","doi-asserted-by":"publisher","DOI":"10.1038\/nature24626"},{"key":"2026061320001234000_2026.06.05.730527v1.52","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2022.02.013"},{"key":"2026061320001234000_2026.06.05.730527v1.53","unstructured":"M. Kleiner , D. Brainard , D. Pelli , A. Ingling , R. Murray , and C. Broussard , 2007. What\u2019s new in Psychtoolbox-3? ."},{"key":"2026061320001234000_2026.06.05.730527v1.54","doi-asserted-by":"publisher","DOI":"10.1068\/p050437"},{"key":"2026061320001234000_2026.06.05.730527v1.55","doi-asserted-by":"crossref","unstructured":"K. Lee , A. Tran , Z. Turan , and M. Meister . The sifting of visual information in the superior colliculus. ELife, 9, 2020.","DOI":"10.7554\/eLife.50678"},{"issue":"7","key":"2026061320001234000_2026.06.05.730527v1.56","doi-asserted-by":"crossref","first-page":"1292","DOI":"10.1364\/JOSAA.20.001292","article-title":"Statistical correlations between two-dimensional images and three-dimensional structures in natural scenes","volume":"20","year":"2003","journal-title":"JOSA A"},{"key":"2026061320001234000_2026.06.05.730527v1.57","doi-asserted-by":"publisher","DOI":"10.1038\/nn.4262"},{"key":"2026061320001234000_2026.06.05.730527v1.58","doi-asserted-by":"publisher","DOI":"10.1159\/000327045"},{"key":"2026061320001234000_2026.06.05.730527v1.59","doi-asserted-by":"publisher","DOI":"10.1126\/science.1248955"},{"key":"2026061320001234000_2026.06.05.730527v1.60","doi-asserted-by":"publisher","DOI":"10.1038\/nn.2389"},{"key":"2026061320001234000_2026.06.05.730527v1.61","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.070755"},{"key":"2026061320001234000_2026.06.05.730527v1.62","doi-asserted-by":"publisher","DOI":"10.1016\/0042-6989(94)90116-3"},{"key":"2026061320001234000_2026.06.05.730527v1.63","doi-asserted-by":"publisher","DOI":"10.1163\/156856897X00366"},{"key":"2026061320001234000_2026.06.05.730527v1.64","doi-asserted-by":"publisher","DOI":"10.1038\/nn.2259"},{"key":"2026061320001234000_2026.06.05.730527v1.65","doi-asserted-by":"crossref","unstructured":"T. Pologruto , B. Sabatini , and K. Svoboda . Scanimage: flexible software for operating laser scanning microscopes. springerta pologruto, bl sabatini. K SvobodaBiomedical engineering online, 2003.","DOI":"10.1186\/1475-925X-2-13"},{"key":"2026061320001234000_2026.06.05.730527v1.66","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2017.05.036"},{"key":"2026061320001234000_2026.06.05.730527v1.67","doi-asserted-by":"crossref","first-page":"477","DOI":"10.1007\/BF00603973","article-title":"Non-directional, movement sensitive neurones of the locust optic lobe","volume":"161","year":"1987","journal-title":"Journal of Comparative Physiology A"},{"key":"2026061320001234000_2026.06.05.730527v1.68","doi-asserted-by":"publisher","DOI":"10.1098\/rsbl.2003.0096"},{"key":"2026061320001234000_2026.06.05.730527v1.69","doi-asserted-by":"publisher","DOI":"10.1152\/jn.1992.68.5.1654"},{"key":"2026061320001234000_2026.06.05.730527v1.70","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1200125109"},{"key":"2026061320001234000_2026.06.05.730527v1.71","doi-asserted-by":"publisher","DOI":"10.1167\/18.8.5"},{"key":"2026061320001234000_2026.06.05.730527v1.72","doi-asserted-by":"publisher","DOI":"10.1126\/science.136.3520.982"},{"key":"2026061320001234000_2026.06.05.730527v1.73","doi-asserted-by":"crossref","first-page":"456","DOI":"10.3758\/BF03330007","article-title":"Luminance controls the perceived 3-d structure of dynamic 2-d displays","volume":"21","year":"1983","journal-title":"Bulletin of the Psychonomic Society"},{"key":"2026061320001234000_2026.06.05.730527v1.74","doi-asserted-by":"crossref","unstructured":"K. Shinomiya , G. Huang , Z. Lu , T. Parag , C. Xu , and R. Aniceto . Comparisons between the on-and off-edge motion pathways in the drosophila brain, 2019.","DOI":"10.7554\/eLife.40025.015"},{"key":"2026061320001234000_2026.06.05.730527v1.75","doi-asserted-by":"publisher","DOI":"10.1007\/s00359-003-0390-6"},{"key":"2026061320001234000_2026.06.05.730527v1.76","doi-asserted-by":"publisher","DOI":"10.1038\/1110"},{"key":"2026061320001234000_2026.06.05.730527v1.77","first-page":"49","volume":"33","year":"2016","journal-title":"The contributions of static visual cues, nonvisual cues, and optic flow in distance estimation"},{"key":"2026061320001234000_2026.06.05.730527v1.78","doi-asserted-by":"crossref","unstructured":"S. Takemura , A. Nern , D. Chklovskii , L. Scheffer , G. Rubin , and I. Meinertzhagen . The comprehensive connectome of a neural substrate for \u2018on, 2017. motion detection in Drosophila. ELife 6.","DOI":"10.7554\/eLife.24394"},{"key":"2026061320001234000_2026.06.05.730527v1.79","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.205.18.2785"},{"key":"2026061320001234000_2026.06.05.730527v1.80","first-page":"642","volume":"62","year":"1989","journal-title":"Underlying mechanisms of the response specificity of expansion\/contraction and rotation cells in the dorsal part of the medial superior temporal area of the macaque monkey"},{"key":"2026061320001234000_2026.06.05.730527v1.81","doi-asserted-by":"publisher","DOI":"10.1016\/J.CUB.2020.04.068"},{"key":"2026061320001234000_2026.06.05.730527v1.82","doi-asserted-by":"crossref","unstructured":"R. Tanaka and D. Clark . Neural mechanisms to exploit positional geometry for collision avoidance. Current Biology, 2022.","DOI":"10.1101\/2021.12.11.472218"},{"key":"2026061320001234000_2026.06.05.730527v1.83","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2015.06.002"},{"key":"2026061320001234000_2026.06.05.730527v1.84","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2012.01.007"},{"key":"2026061320001234000_2026.06.05.730527v1.85","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2021.03.025"},{"key":"2026061320001234000_2026.06.05.730527v1.86","doi-asserted-by":"publisher","DOI":"10.1038\/356236a0"},{"key":"2026061320001234000_2026.06.05.730527v1.87","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1111\/j.1475-1313.2004.00188.x","article-title":"Stimulating human accommodation without changes in focus","volume":"24","year":"2004","journal-title":"Ophthalmic and Physiological Optics"},{"key":"2026061320001234000_2026.06.05.730527v1.88","doi-asserted-by":"crossref","unstructured":"M. Wu , A. Nern , W. Ryan Williamson , M. Morimoto , M. Reiser , and G. Card . Visual projection neurons in the drosophila lobula link feature detection to distinct behavioral programs. ELife, 5, 2016.","DOI":"10.7554\/eLife.21022"},{"key":"2026061320001234000_2026.06.05.730527v1.89","first-page":"2410768122","article-title":"Broken time-reversal symmetry in visual motion detection","volume":"122","year":"2025","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"key":"2026061320001234000_2026.06.05.730527v1.90","doi-asserted-by":"publisher","DOI":"10.1007\/s00359-008-0403-6"},{"key":"2026061320001234000_2026.06.05.730527v1.91","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2013.08.015"},{"issue":"1","key":"2026061320001234000_2026.06.05.730527v1.92","doi-asserted-by":"crossref","first-page":"3697","DOI":"10.1038\/s41467-018-05875-1","article-title":"Speed dependent descending control of freezing behavior in drosophila melanogaster","volume":"9","year":"2018","journal-title":"Nature Communications"},{"key":"2026061320001234000_2026.06.05.730527v1.93","doi-asserted-by":"publisher","DOI":"10.1167\/jov.20.2.2"},{"key":"2026061320001234000_2026.06.05.730527v1.94","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2014.08.037"},{"key":"2026061320001234000_2026.06.05.730527v1.95","doi-asserted-by":"crossref","unstructured":"B. Zhou , Z. Li , S. Kim , J. Lafferty , and D. Clark . Shallow neural networks trained to detect collisions recover features of visual loom-selective neurons. ELife, 11, 2022.","DOI":"10.7554\/eLife.72067"}],"container-title":[],"original-title":[],"link":[{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.64898\/2026.06.05.730527","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,14]],"date-time":"2026-06-14T03:00:22Z","timestamp":1781406022000},"score":1,"resource":{"primary":{"URL":"http:\/\/biorxiv.org\/lookup\/doi\/10.64898\/2026.06.05.730527"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,10]]},"references-count":95,"URL":"https:\/\/doi.org\/10.64898\/2026.06.05.730527","relation":{},"subject":[],"published":{"date-parts":[[2026,6,10]]},"subtype":"preprint"}}