{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,6,4]],"date-time":"2026-06-04T07:19:37Z","timestamp":1780557577323,"version":"3.54.1"},"posted":{"date-parts":[[2025,4,7]]},"group-title":"Neuroscience","reference-count":133,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2025,4,7]],"date-time":"2025-04-07T00:00:00Z","timestamp":1743984000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000053","name":"National Eye Institute","doi-asserted-by":"crossref","award":["EY-028282"],"award-info":[{"award-number":["EY-028282"]}],"id":[{"id":"10.13039\/100000053","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["RF1-MH129260"],"award-info":[{"award-number":["RF1-MH129260"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["P51 OD010425"],"award-info":[{"award-number":["P51 OD010425"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2026,1,28]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                <jats:p>The fovea is a unique specialization of the primate retina and is a promising site for obtaining the first complete connectome of a human central nervous system (CNS) structure. Within the fovea, neural cells and circuits have been miniaturized and compressed during evolution to sample the visual image at highest spatial resolution and begin the neural processing that serves human form, color, and motion perception. Here we present a comprehensive analysis of a sample of human foveal retina using deep learning-based segmentation to reconstruct all cells and synaptic connections at nanoscale resolution. We classified \u223c3,000 cells into 51 distinct morphological types based on their structural features and connectivity patterns. Our observations reveal novel synaptic pathways absent in non-human primates, suggesting specialized circuits contribute uniquely to human trichromatic color vision. A biophysical model of the distinct connectomes made by gap junctions (electrical synapses) between short- (S) and medium-long- (ML) wavelength-sensitive cone photoreceptors, suggests chromatic interactions between S and ML cones prior to the first chemical synapse. Segmentation of retinal ganglion cells (RGCs) suggests the presence of only 11 visual pathways, with 5 high-density RGC pathways accounting for over 95% of foveal output to the brain: a dramatic contrast to the 40+ ganglion cell types recognized in mouse retina. Our connectomic analysis reveals distinctive features of human neural circuitry and demonstrates how AI-based computational approaches can advance understanding of human brain structure and function.<\/jats:p>\n                <jats:sec>\n                  <jats:title>Significant statement<\/jats:title>\n                  <jats:p>Vision begins in the retina, an accessible outpost of the brain located at the back of the eye. When we directly view an object, we use the fovea\u2014a tiny region containing over 50 distinct, densely packed neuron types. How these diverse cells interconnect through synapses to transmit visual information has been difficult to determine. Here, we used artificial intelligence-based computational methods to create a complete wiring diagram\u2014a connectome\u2014mapping every neuron and synapse in a sample of human fovea. This connectomic resource will be freely available to researchers worldwide. Discoveries so far include synaptic circuits linked to human color vision, demonstrating how connectomics can reveal what makes the human nervous system unique.<\/jats:p>\n                <\/jats:sec>","DOI":"10.1101\/2025.04.05.647403","type":"posted-content","created":{"date-parts":[[2025,4,8]],"date-time":"2025-04-08T00:06:22Z","timestamp":1744070782000},"source":"Crossref","is-referenced-by-count":3,"title":["Connectome of a human foveal retina"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6722-4063","authenticated-orcid":false,"given":"Yeon Jin","family":"Kim","sequence":"first","affiliation":[{"name":"Department of Neurobiology & Biophysics, University of Washington, Seattle, WA, 98195"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0885-0689","authenticated-orcid":false,"given":"Orin","family":"Packer","sequence":"additional","affiliation":[{"name":"Department of Neurobiology & Biophysics, University of Washington, Seattle, WA, 98195"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0622-9338","authenticated-orcid":false,"given":"Thomas","family":"Macrina","sequence":"additional","affiliation":[{"name":"Zetta AI LLC, Sherrill, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3902-239X","authenticated-orcid":false,"given":"Andreas","family":"Pollreisz","sequence":"additional","affiliation":[{"name":"Department of Ophthalmology, Medical University of Vienna, Vienna, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9769-1538","authenticated-orcid":false,"given":"Christine A.","family":"Curcio","sequence":"additional","affiliation":[{"name":"Department of Ophthalmology and Vision Sciences, University of Alabama at Birmingham, Alabama 35294"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5707-0155","authenticated-orcid":false,"given":"Kisuk","family":"Lee","sequence":"additional","affiliation":[{"name":"Zetta AI LLC, Sherrill, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0260-4532","authenticated-orcid":false,"given":"Nico","family":"Kemnitz","sequence":"additional","affiliation":[{"name":"Zetta AI LLC, Sherrill, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0295-8445","authenticated-orcid":false,"given":"Dodam","family":"Ih","sequence":"additional","affiliation":[{"name":"Zetta AI LLC, Sherrill, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9936-1794","authenticated-orcid":false,"given":"Tri","family":"Nguyen","sequence":"additional","affiliation":[{"name":"Zetta AI LLC, Sherrill, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2077-8165","authenticated-orcid":false,"given":"Ran","family":"Lu","sequence":"additional","affiliation":[{"name":"Zetta AI LLC, Sherrill, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sergiy","family":"Popovych","sequence":"additional","affiliation":[{"name":"Zetta AI LLC, Sherrill, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7367-4776","authenticated-orcid":false,"given":"Akhilesh","family":"Halageri","sequence":"additional","affiliation":[{"name":"Zetta AI LLC, Sherrill, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4681-6342","authenticated-orcid":false,"given":"J. Alexander","family":"Bae","sequence":"additional","affiliation":[{"name":"Zetta AI LLC, Sherrill, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4272-3576","authenticated-orcid":false,"given":"Joe","family":"Strout","sequence":"additional","affiliation":[{"name":"Zetta AI LLC, Sherrill, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4454-6171","authenticated-orcid":false,"given":"Stephan","family":"Gerhard","sequence":"additional","affiliation":[{"name":"Aware LLC, Rappenstrasse 19, 8307 Effretikon, Switzerland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Robert G.","family":"Smith","sequence":"additional","affiliation":[{"name":"Department of Neuroscience, University of Pennsylvania, Philadelphia, PA 19104"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5166-9658","authenticated-orcid":false,"given":"Paul R.","family":"Martin","sequence":"additional","affiliation":[{"name":"The University of Sydney, Save Sight Institute and Discipline of Ophthalmology, Faculty of Medicine and Health, Sydney, NSW, 2000, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9595-8152","authenticated-orcid":false,"given":"Ulrike","family":"Gr\u00fcnert","sequence":"additional","affiliation":[{"name":"The University of Sydney, Save Sight Institute and Discipline of Ophthalmology, Faculty of Medicine and Health, Sydney, NSW, 2000, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8476-5585","authenticated-orcid":false,"given":"Dennis M.","family":"Dacey","sequence":"additional","affiliation":[{"name":"Department of Neurobiology & Biophysics, University of Washington, Seattle, WA, 98195"},{"name":"Washington National Primate Research Center, Seattle, WA 98195"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026020510583500000_2025.04.05.647403v5.1","first-page":"119","article-title":"La r\u00e8tine des vert\u00e8br\u00e8s","volume":"9","year":"1892","journal-title":"La Cellule"},{"key":"2026020510583500000_2025.04.05.647403v5.2","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.neuro.23.1.249"},{"key":"2026020510583500000_2025.04.05.647403v5.3","unstructured":"J. E. Dowling , The Retina: An approachable part of the brain (Harvard University Press, Cambridge, MA, ed. 2nd, 2012), doi.org\/10.2307\/j.ctv31zqj2d, pp. 384."},{"key":"2026020510583500000_2025.04.05.647403v5.4","unstructured":"R. W. Rodieck , The First Steps in Seeing (Sinauer Associates, Inc., Sunderland MA, 1998), pp. 562 pp."},{"key":"2026020510583500000_2025.04.05.647403v5.5","doi-asserted-by":"publisher","DOI":"10.1038\/nature12346"},{"key":"2026020510583500000_2025.04.05.647403v5.6","doi-asserted-by":"publisher","DOI":"10.1038\/nature13240"},{"key":"2026020510583500000_2025.04.05.647403v5.7","doi-asserted-by":"publisher","DOI":"10.1038\/nature09818"},{"key":"2026020510583500000_2025.04.05.647403v5.8","doi-asserted-by":"publisher","DOI":"10.3390\/molecules17010355"},{"key":"2026020510583500000_2025.04.05.647403v5.9","doi-asserted-by":"publisher","DOI":"10.1038\/371070a0"},{"key":"2026020510583500000_2025.04.05.647403v5.10","doi-asserted-by":"publisher","DOI":"10.1038\/381613a0"},{"key":"2026020510583500000_2025.04.05.647403v5.11","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2016.07.054"},{"key":"2026020510583500000_2025.04.05.647403v5.12","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2019.01.004"},{"key":"2026020510583500000_2025.04.05.647403v5.13","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-020-66092-9"},{"key":"2026020510583500000_2025.04.05.647403v5.14","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-023-06638-9"},{"key":"2026020510583500000_2025.04.05.647403v5.15","doi-asserted-by":"publisher","DOI":"10.21203\/rs.3.rs-3471275\/v1"},{"key":"2026020510583500000_2025.04.05.647403v5.16","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-024-07558-y"},{"key":"2026020510583500000_2025.04.05.647403v5.17","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-024-07389-x"},{"key":"2026020510583500000_2025.04.05.647403v5.18","doi-asserted-by":"publisher","DOI":"10.1126\/science.adk4858"},{"key":"2026020510583500000_2025.04.05.647403v5.19","doi-asserted-by":"publisher","DOI":"10.1126\/science.abo0924"},{"key":"2026020510583500000_2025.04.05.647403v5.20","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2022.01.023"},{"key":"2026020510583500000_2025.04.05.647403v5.21","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-025-08840-3"},{"key":"2026020510583500000_2025.04.05.647403v5.22","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2020.08.010"},{"key":"2026020510583500000_2025.04.05.647403v5.23","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2008.08.010"},{"key":"2026020510583500000_2025.04.05.647403v5.24","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2007.10.012"},{"key":"2026020510583500000_2025.04.05.647403v5.25","doi-asserted-by":"crossref","first-page":"787737","DOI":"10.3389\/fnsys.2022.787737","article-title":"Topological Maps and Brain Computations From Low to High","volume":"16","year":"2022","journal-title":"Frontiers in systems neuroscience"},{"key":"2026020510583500000_2025.04.05.647403v5.26","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.2572-11.2011"},{"key":"2026020510583500000_2025.04.05.647403v5.27","doi-asserted-by":"publisher","DOI":"10.1126\/science.7754376"},{"key":"2026020510583500000_2025.04.05.647403v5.28","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1760-09.2009"},{"key":"2026020510583500000_2025.04.05.647403v5.29","doi-asserted-by":"publisher","DOI":"10.1038\/nature16468"},{"key":"2026020510583500000_2025.04.05.647403v5.30","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2018.04.040"},{"key":"2026020510583500000_2025.04.05.647403v5.31","doi-asserted-by":"crossref","first-page":"111040","DOI":"10.1016\/j.celrep.2022.111040","article-title":"Unified classification of mouse retinal ganglion cells using function, morphology, and gene expression","volume":"40","year":"2022","journal-title":"Cell reports"},{"key":"2026020510583500000_2025.04.05.647403v5.32","doi-asserted-by":"publisher","DOI":"10.1002\/cne.24157"},{"key":"2026020510583500000_2025.04.05.647403v5.33","doi-asserted-by":"crossref","first-page":"2862","DOI":"10.1038\/s41467-022-30405-5","article-title":"Origins of direction selectivity in the primate retina","volume":"13","year":"2022","journal-title":"Nature communications"},{"key":"2026020510583500000_2025.04.05.647403v5.34","doi-asserted-by":"publisher","DOI":"10.1002\/cne.902920402"},{"key":"2026020510583500000_2025.04.05.647403v5.35","doi-asserted-by":"crossref","unstructured":"C. A. Curcio , D. Kar , C. Owsley , K. R. Sloan , T. Ach , Age-Related Macular Degeneration, a Mathematically Tractable Disease. Investigative ophthalmology & visual science 65, 4 (2024).","DOI":"10.1167\/iovs.65.3.4"},{"key":"2026020510583500000_2025.04.05.647403v5.36","doi-asserted-by":"publisher","DOI":"10.1002\/cne.902690403"},{"key":"2026020510583500000_2025.04.05.647403v5.37","doi-asserted-by":"publisher","DOI":"10.1016\/j.visres.2007.01.007"},{"key":"2026020510583500000_2025.04.05.647403v5.38","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1671-20.2020"},{"key":"2026020510583500000_2025.04.05.647403v5.39","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2019.04.001"},{"key":"2026020510583500000_2025.04.05.647403v5.40","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1038\/s41592-021-01330-0","article-title":"FlyWire: online community for whole-brain connectomics","volume":"19","year":"2022","journal-title":"Nature methods"},{"key":"2026020510583500000_2025.04.05.647403v5.41","doi-asserted-by":"publisher","DOI":"10.1016\/j.preteyeres.2020.100844"},{"key":"2026020510583500000_2025.04.05.647403v5.42","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1146\/annurev-vision-100419-115801","article-title":"and Connections of Ganglion Cells in Primate Retina","volume":"7","year":"2021","journal-title":"Annu Rev Vis Sci"},{"key":"2026020510583500000_2025.04.05.647403v5.43","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2313820121"},{"key":"2026020510583500000_2025.04.05.647403v5.44","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2300545120"},{"key":"2026020510583500000_2025.04.05.647403v5.45","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.13-12-05334.1993"},{"key":"2026020510583500000_2025.04.05.647403v5.46","doi-asserted-by":"publisher","DOI":"10.1017\/S0952523800010191"},{"key":"2026020510583500000_2025.04.05.647403v5.47","doi-asserted-by":"publisher","DOI":"10.1038\/367731a0"},{"key":"2026020510583500000_2025.04.05.647403v5.48","doi-asserted-by":"publisher","DOI":"10.1038\/13189"},{"key":"2026020510583500000_2025.04.05.647403v5.49","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.18-09-03373.1998"},{"key":"2026020510583500000_2025.04.05.647403v5.50","doi-asserted-by":"crossref","unstructured":"50. R. A. Masri , U. Greferath , E. L. Fletcher , P. R. Martin , U. Gr\u00fcnert , Immunohistochemistry and spatial density of M\u00fcller cells in the human fovea. Investigative ophthalmology & visual science in press (2025).","DOI":"10.1167\/iovs.66.2.46"},{"key":"2026020510583500000_2025.04.05.647403v5.51","doi-asserted-by":"publisher","DOI":"10.1007\/s00429-017-1370-x"},{"key":"2026020510583500000_2025.04.05.647403v5.52","doi-asserted-by":"crossref","first-page":"633","DOI":"10.1038\/s41598-023-27453-2","article-title":"Trogocytosis of neurons and glial cells by microglia in a healthy adult macaque retina","volume":"13","year":"2023","journal-title":"Sci Rep"},{"key":"2026020510583500000_2025.04.05.647403v5.53","doi-asserted-by":"publisher","DOI":"10.1016\/S0896-6273(00)00011-8"},{"key":"2026020510583500000_2025.04.05.647403v5.54","doi-asserted-by":"crossref","first-page":"3486","DOI":"10.1038\/s41467-023-38943-2","article-title":"Mechanisms of simultaneous linear and nonlinear computations at the mammalian cone photoreceptor synapse","volume":"14","year":"2023","journal-title":"Nature communications"},{"key":"2026020510583500000_2025.04.05.647403v5.55","doi-asserted-by":"publisher","DOI":"10.1017\/S0952523810000465"},{"key":"2026020510583500000_2025.04.05.647403v5.56","doi-asserted-by":"publisher","DOI":"10.1002\/cne.903250203"},{"key":"2026020510583500000_2025.04.05.647403v5.57","doi-asserted-by":"crossref","unstructured":"E. Strettoi , R. A. Masri , U. Gr\u00fcnert , AII amacrine cells in the primate fovea contribute to photopic vision. Sci Rep 8, 16429 (2018).","DOI":"10.1038\/s41598-018-34621-2"},{"key":"2026020510583500000_2025.04.05.647403v5.58","doi-asserted-by":"publisher","DOI":"10.1002\/cne.903610315"},{"key":"2026020510583500000_2025.04.05.647403v5.59","doi-asserted-by":"publisher","DOI":"10.1111\/j.1460-9568.1997.tb01638.x"},{"key":"2026020510583500000_2025.04.05.647403v5.60","doi-asserted-by":"publisher","DOI":"10.1038\/nn1274"},{"key":"2026020510583500000_2025.04.05.647403v5.61","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.4749-11.2012"},{"key":"2026020510583500000_2025.04.05.647403v5.62","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0778-19.2019"},{"key":"2026020510583500000_2025.04.05.647403v5.63","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1976.sp011631"},{"key":"2026020510583500000_2025.04.05.647403v5.64","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.1988.0043"},{"key":"2026020510583500000_2025.04.05.647403v5.65","doi-asserted-by":"crossref","first-page":"1900","DOI":"10.1016\/S0960-9822(02)01261-7","article-title":"Electrical coupling between mammalian cones","volume":"12","year":"2002","journal-title":"Current biology : CB"},{"key":"2026020510583500000_2025.04.05.647403v5.66","doi-asserted-by":"publisher","DOI":"10.1364\/JOSAA.17.000635"},{"key":"2026020510583500000_2025.04.05.647403v5.67","doi-asserted-by":"publisher","DOI":"10.1038\/nn1275"},{"key":"2026020510583500000_2025.04.05.647403v5.68","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1096-9861(19970929)386:3<443::AID-CNE8>3.0.CO;2-1"},{"key":"2026020510583500000_2025.04.05.647403v5.69","doi-asserted-by":"publisher","DOI":"10.1016\/0042-6989(92)90042-H"},{"key":"2026020510583500000_2025.04.05.647403v5.70","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.2414-05.2005"},{"key":"2026020510583500000_2025.04.05.647403v5.71","doi-asserted-by":"publisher","DOI":"10.1016\/j.preteyeres.2012.04.003"},{"key":"2026020510583500000_2025.04.05.647403v5.72","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.4385-10.2011"},{"key":"2026020510583500000_2025.04.05.647403v5.73","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1688-17.2017"},{"key":"2026020510583500000_2025.04.05.647403v5.74","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.2010.194076"},{"key":"2026020510583500000_2025.04.05.647403v5.75","doi-asserted-by":"publisher","DOI":"10.1017\/S0952523800009093"},{"key":"2026020510583500000_2025.04.05.647403v5.76","doi-asserted-by":"publisher","DOI":"10.1126\/science.271.5249.656"},{"key":"2026020510583500000_2025.04.05.647403v5.77","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1096-9861(19980406)393:2<196::AID-CNE5>3.0.CO;2-Y"},{"key":"2026020510583500000_2025.04.05.647403v5.78","first-page":"105","article-title":"Some OFF bipolar cell types make contact with both rods and cones in macaque and mouse retinas","volume":"8","year":"2014","journal-title":"Frontiers in neuroanatomy"},{"key":"2026020510583500000_2025.04.05.647403v5.79","first-page":"122","article-title":"OFF bipolar cells in macaque retina: type-specific connectivity in the outer and inner synaptic layers","volume":"9","year":"2015","journal-title":"Frontiers in neuroanatomy"},{"key":"2026020510583500000_2025.04.05.647403v5.80","doi-asserted-by":"crossref","first-page":"2070","DOI":"10.1016\/j.cub.2016.06.016","article-title":"Glutamatergic Monopolar Interneurons Provide a Novel Pathway of Excitation in the Mouse Retina","volume":"26","year":"2016","journal-title":"Current biology : CB"},{"key":"2026020510583500000_2025.04.05.647403v5.81","doi-asserted-by":"publisher","DOI":"10.1002\/cne.23821"},{"key":"2026020510583500000_2025.04.05.647403v5.82","doi-asserted-by":"publisher","DOI":"10.1017\/S0952523811000368"},{"key":"2026020510583500000_2025.04.05.647403v5.83","first-page":"1","article-title":"Inhibitory Interneurons in the Retina: Types, Circuitry, and Function","volume":"3","year":"2017","journal-title":"Annu Rev Vis Sci"},{"key":"2026020510583500000_2025.04.05.647403v5.84","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.4274-07.2008"},{"key":"2026020510583500000_2025.04.05.647403v5.85","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.4442-08.2009"},{"key":"2026020510583500000_2025.04.05.647403v5.86","doi-asserted-by":"publisher","DOI":"10.1017\/S0952523813000230"},{"key":"2026020510583500000_2025.04.05.647403v5.87","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1017\/S0952523813000308","article-title":"Processing of S-cone signals in the inner plexiform layer of the mammalian retina","volume":"31","year":"2014","journal-title":"Visual neuroscience"},{"key":"2026020510583500000_2025.04.05.647403v5.88","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2020.08.013"},{"key":"2026020510583500000_2025.04.05.647403v5.89","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1249-13.2013"},{"key":"2026020510583500000_2025.04.05.647403v5.90","doi-asserted-by":"crossref","first-page":"109916","DOI":"10.1016\/j.isci.2024.109916","article-title":"Comprehensive single-cell atlas of the mouse retina","volume":"27","year":"2024","journal-title":"iScience"},{"key":"2026020510583500000_2025.04.05.647403v5.91","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0471-20.2020"},{"key":"2026020510583500000_2025.04.05.647403v5.92","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1146\/annurev-vision-102122-104700","article-title":"Behavior-Specific Computations in the Vertebrate Retina","volume":"11","year":"2025","journal-title":"Annu Rev Vis Sci"},{"key":"2026020510583500000_2025.04.05.647403v5.93","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1146\/annurev-vision-100419-112009","article-title":"Feature Detection by Retinal Ganglion Cells","volume":"8","year":"2022","journal-title":"Annu Rev Vis Sci"},{"key":"2026020510583500000_2025.04.05.647403v5.94","doi-asserted-by":"crossref","first-page":"1920","DOI":"10.1038\/s41467-024-46348-y","article-title":"Distributed feature representations of natural stimuli across parallel retinal pathways","volume":"15","year":"2024","journal-title":"Nature communications"},{"key":"2026020510583500000_2025.04.05.647403v5.95","doi-asserted-by":"publisher","DOI":"10.1364\/JOSAA.22.002013"},{"key":"2026020510583500000_2025.04.05.647403v5.96","unstructured":"J. D. Crook , O. S. Packer , J. B. Troy , D. M. Dacey , \u201cSynaptic mechanisms of color and luminance coding: rediscovering the X-Y dichotomy in primate retinal ganglion cells\u201d in The New Visual Neurosciences, L. M. Chalupa , J. S. Werner , Eds. (MIT press, Cambridge, MA, 2014), chap. 11, pp. 123\u2013144."},{"key":"2026020510583500000_2025.04.05.647403v5.97","doi-asserted-by":"publisher","DOI":"10.1016\/j.visres.2004.09.001"},{"key":"2026020510583500000_2025.04.05.647403v5.98","doi-asserted-by":"publisher","DOI":"10.1109\/JRPROC.1959.287207"},{"key":"2026020510583500000_2025.04.05.647403v5.99","doi-asserted-by":"crossref","first-page":"499","DOI":"10.1146\/annurev-vision-082114-035742","article-title":"Control and Functions of Fixational Eye Movements","volume":"1","year":"2015","journal-title":"Annu Rev Vis Sci"},{"key":"2026020510583500000_2025.04.05.647403v5.100","doi-asserted-by":"publisher","DOI":"10.1038\/nn.2663"},{"key":"2026020510583500000_2025.04.05.647403v5.101","doi-asserted-by":"publisher","DOI":"10.1016\/j.visres.2015.01.018"},{"key":"2026020510583500000_2025.04.05.647403v5.102","first-page":"1099","article-title":"Journal of the Optical Society of America","volume":"34","year":"2017","journal-title":"A, Optics, image science, and vision"},{"key":"2026020510583500000_2025.04.05.647403v5.103","unstructured":"A. Stockman , D. H. Brainard , \u201cColor Vision Mechanisms\u201d in OSA Handbook of Optics, M. Bass , Ed. (McGraw-Hill, New York, 2010), pp. 11.11\u201311.104."},{"key":"2026020510583500000_2025.04.05.647403v5.104","doi-asserted-by":"crossref","unstructured":"P. Lennie , P. W. Haake , D. R. Williams , \u201cThe design of chromatically opponent receptive fields\u201d in Computational models of visual processing, M. S. Landy , J. A. Movshon , Eds. (The MIT Press, Cambridge MA, 1991), pp. 71\u201382.","DOI":"10.7551\/mitpress\/2002.003.0010"},{"key":"2026020510583500000_2025.04.05.647403v5.105","doi-asserted-by":"publisher","DOI":"10.1038\/nature09424"},{"key":"2026020510583500000_2025.04.05.647403v5.106","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2405138121"},{"key":"2026020510583500000_2025.04.05.647403v5.107","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1038\/s41467-023-44354-0","article-title":"Petascale pipeline for precise alignment of images from serial section electron microscopy","volume":"15","year":"2024","journal-title":"Nature communications"},{"key":"2026020510583500000_2025.04.05.647403v5.108","doi-asserted-by":"crossref","unstructured":"D. G. Lowe (1999) Object recognition from local scale-invariant features. in Proceedings of the Seventh IEEE International Conference on Computer Vision (IEEE, Kerkyra, Greece), pp 1150\u20131157.","DOI":"10.1109\/ICCV.1999.790410"},{"key":"2026020510583500000_2025.04.05.647403v5.109","doi-asserted-by":"crossref","first-page":"717","DOI":"10.1038\/nmeth.2072","article-title":"Elastic volume reconstruction from series of ultra-thin microscopy sections","volume":"9","year":"2012","journal-title":"Nature methods"},{"key":"2026020510583500000_2025.04.05.647403v5.110","doi-asserted-by":"publisher","DOI":"10.1101\/2021.07.28.454025"},{"key":"2026020510583500000_2025.04.05.647403v5.111","unstructured":"111. K. Lee, J. Zung, P. H. Li, V. Jain, H. S. Seung, Superhuman accuracy on the SNEMI3D connectomics challenge. arXiv preprint arXiv:1706.00120 (2017)."},{"key":"2026020510583500000_2025.04.05.647403v5.112","doi-asserted-by":"publisher","DOI":"10.1109\/TMI.2021.3097826"},{"key":"2026020510583500000_2025.04.05.647403v5.113","doi-asserted-by":"publisher","DOI":"10.1101\/2021.08.04.455162"},{"key":"2026020510583500000_2025.04.05.647403v5.114","doi-asserted-by":"publisher","DOI":"10.1038\/s41592-024-02426-z"},{"key":"2026020510583500000_2025.04.05.647403v5.115","unstructured":"115. J. Maitin-Shepard, et al. (2021) google\/neuroglancer."},{"key":"2026020510583500000_2025.04.05.647403v5.116","doi-asserted-by":"publisher","DOI":"10.1111\/j.1460-9568.1991.tb00043.x"},{"key":"2026020510583500000_2025.04.05.647403v5.117","doi-asserted-by":"publisher","DOI":"10.1152\/physrev.1991.71.2.447"},{"key":"2026020510583500000_2025.04.05.647403v5.118","doi-asserted-by":"crossref","unstructured":"118. N. L. Turner , et al. (2020) Synaptic Partner Assignment Using Attentional Voxel Association Networks. in IEEE 17th International Symposium on Biomedical Imaging (ISBI), pp 1\u20135.","DOI":"10.1109\/ISBI45749.2020.9098489"},{"key":"2026020510583500000_2025.04.05.647403v5.119","unstructured":"119. R. Lu, A. Zlateski, H. S. Seung, Large-scale image segmentation based on distributed clustering algorithms. arXiv arXiv. http:\/\/arxiv.org\/abs\/2106.10795 [cs.CV], 1-8 (2021)."},{"key":"2026020510583500000_2025.04.05.647403v5.120","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2020.07.035"},{"key":"2026020510583500000_2025.04.05.647403v5.121","doi-asserted-by":"crossref","unstructured":"J. van Hateren , T. Lamb , The photocurrent response of human cones is fast and monophasic. BMC neuroscience 7, 34 (2006).","DOI":"10.1186\/1471-2202-7-34"},{"key":"2026020510583500000_2025.04.05.647403v5.122","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1992.sp019111"},{"key":"2026020510583500000_2025.04.05.647403v5.123","doi-asserted-by":"publisher","DOI":"10.1085\/jgp.111.1.7"},{"key":"2026020510583500000_2025.04.05.647403v5.124","doi-asserted-by":"publisher","DOI":"10.1085\/jgp.111.1.39"},{"key":"2026020510583500000_2025.04.05.647403v5.125","doi-asserted-by":"publisher","DOI":"10.1017\/S0952523805224045"},{"key":"2026020510583500000_2025.04.05.647403v5.126","doi-asserted-by":"publisher","DOI":"10.1167\/5.8.331"},{"key":"2026020510583500000_2025.04.05.647403v5.127","doi-asserted-by":"publisher","DOI":"10.1167\/7.1.1"},{"key":"2026020510583500000_2025.04.05.647403v5.128","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1987.sp016691"},{"key":"2026020510583500000_2025.04.05.647403v5.129","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.1983.0091"},{"key":"2026020510583500000_2025.04.05.647403v5.130","doi-asserted-by":"publisher","DOI":"10.1016\/S0042-6989(98)00225-9"},{"key":"2026020510583500000_2025.04.05.647403v5.131","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.19-22-09848.1999"},{"key":"2026020510583500000_2025.04.05.647403v5.132","doi-asserted-by":"publisher","DOI":"10.1007\/s002320001094"},{"key":"2026020510583500000_2025.04.05.647403v5.133","doi-asserted-by":"publisher","DOI":"10.1016\/0165-0270(92)90019-A"}],"container-title":[],"original-title":[],"link":[{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.1101\/2025.04.05.647403","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,5]],"date-time":"2026-02-05T18:58:49Z","timestamp":1770317929000},"score":1,"resource":{"primary":{"URL":"http:\/\/biorxiv.org\/lookup\/doi\/10.1101\/2025.04.05.647403"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,4,7]]},"references-count":133,"URL":"https:\/\/doi.org\/10.1101\/2025.04.05.647403","relation":{},"subject":[],"published":{"date-parts":[[2025,4,7]]},"subtype":"preprint"}}