IsoNet2 determines cellular structures at submolecular resolution without averaging

IsoNet2 determines cellular structures at submolecular resolution without averaging | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article IsoNet2 determines cellular structures at submolecular resolution without averaging Z. Hong Zhou, Yun-Tao Liu, Hongcheng Fan, Jonathan Jih, Liam Tran, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8363763/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted You are reading this latest preprint version Abstract We introduce IsoNet2 , an end-to-end self-supervised deep-learning method that directly reconstructs high-quality 3D densities from cryogenic electron tomograms. A unified network simultaneously performs denoising, contrast transfer function correction, and missing-wedge restoration, achieving ~ 20 Å resolution without averaging. A feature-rich GUI enables rapid, dataset-specific fine-tuning for end-users. IsoNet2 resolves domain organization in HIV capsid proteins, tRNA occupancy in individual ribosomes, and in situ architectures of mitochondrial respiration-related complexes, enabling atomic-level interpretation of cellular environments. Biological sciences/Biophysics/Computational biophysics Biological sciences/Structural biology/Electron microscopy/Cryoelectron tomography Full Text Additional Declarations There is NO Competing Interest. Supplementary Files SupplementaryVideo1.mp4 Supplementary Video 1 SupplementaryVideo2.mp4 Supplementary Video 2 SupplementaryVideo3.mp4 Supplementary Video 3 Cite Share Download PDF Status: Under Review Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-8363763","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":577451661,"identity":"4fbb7dc8-643a-4b9b-9121-75dbb9de5ec0","order_by":0,"name":"Z. Hong Zhou","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA1ElEQVRIiWNgGAWjYBACgwNAgvEPkGTvMfzMYADkHSBGCwMPkOQ5YyzNUEGSFokcM26GM8RoOd57+AWDxB05g5tvzJgZ2xjk+G4k4Ndif+ZcmgWDwTNjg9s5YC3GkoS0GNzIMTNgSDicuAGqJXEDcVoOALXcPAPU8o+hnhgtxg8YG4BabvCYcQNtSTAgqOXMGTOGxIbDxpJn0pKlGdskDGeeeUBAy/Ee4w8f/xyW4zt++ODnn2028nzHCdgCBGwSSGokCCoHAeYPRCkbBaNgFIyCkQsAaDBNvK0sJeAAAAAASUVORK5CYII=","orcid":"https://orcid.org/0000-0002-8373-4717","institution":"University of California, Los Angeles","correspondingAuthor":true,"prefix":"","firstName":"Z.","middleName":"Hong","lastName":"Zhou","suffix":""},{"id":577451662,"identity":"78920e4f-29a9-40e7-94e8-709dd3eb7b0d","order_by":1,"name":"Yun-Tao Liu","email":"","orcid":"https://orcid.org/0000-0003-2577-1859","institution":"University of California, Los Angeles","correspondingAuthor":false,"prefix":"","firstName":"Yun-Tao","middleName":"","lastName":"Liu","suffix":""},{"id":577451663,"identity":"46bfc761-38d3-4857-ba72-1fe8863f2376","order_by":2,"name":"Hongcheng Fan","email":"","orcid":"https://orcid.org/0000-0001-9038-7998","institution":"The University of California, Los Angeles","correspondingAuthor":false,"prefix":"","firstName":"Hongcheng","middleName":"","lastName":"Fan","suffix":""},{"id":577451664,"identity":"b76ca1b8-160d-45b0-9332-ac9144571bb2","order_by":3,"name":"Jonathan Jih","email":"","orcid":"","institution":"University of California, Los Angeles","correspondingAuthor":false,"prefix":"","firstName":"Jonathan","middleName":"","lastName":"Jih","suffix":""},{"id":577451665,"identity":"afd3a95c-8a66-4778-b921-8aa31ef485ea","order_by":4,"name":"Liam Tran","email":"","orcid":"","institution":"University of California, Los Angeles","correspondingAuthor":false,"prefix":"","firstName":"Liam","middleName":"","lastName":"Tran","suffix":""},{"id":577451666,"identity":"9e8a5359-3277-4aa5-a643-5795cd9544ae","order_by":5,"name":"Xiaoying Zhang","email":"","orcid":"","institution":"University of California, Los Angeles","correspondingAuthor":false,"prefix":"","firstName":"Xiaoying","middleName":"","lastName":"Zhang","suffix":""}],"badges":[],"createdAt":"2025-12-15 08:45:56","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8363763/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8363763/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":100858732,"identity":"14f30a9d-6ead-4ba6-8a39-5710406c3675","added_by":"auto","created_at":"2026-01-22 07:24:42","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":22000750,"visible":true,"origin":"","legend":"","description":"","filename":"ManuscriptDec122025ForSubmissionv3yuntao.docx","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/53b3447d89fb08331ceda047.docx"},{"id":100818901,"identity":"0ce7901b-2e67-4e5f-a8ce-6963f3c126ce","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"json","order_by":1,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":6589,"visible":true,"origin":"","legend":"","description":"","filename":"NCOMMS26004037T.json","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/9b305fd45fd1058bb1ba7dcc.json"},{"id":100818927,"identity":"784fb3f7-965c-4541-95a8-13ea2e005f8c","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"xml","order_by":2,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":104235,"visible":true,"origin":"","legend":"","description":"","filename":"NCOMMS26004037T0enriched.xml","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/fa27c5a9bd483435669a8c07.xml"},{"id":100818906,"identity":"c764135d-bfd2-4609-bb38-457fb14ed2bd","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"png","order_by":3,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":3212853,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/1fcffa7214e295eb3b29c01e.png"},{"id":100818902,"identity":"c799b44e-5abf-43cd-bc67-dcafb1a8ce17","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"png","order_by":4,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1600468,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage10.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/49efd8e5cc51cbafedaaf3d2.png"},{"id":100857934,"identity":"9cce2811-dfb7-4b1b-8da2-144fdade7494","added_by":"auto","created_at":"2026-01-22 07:23:27","extension":"png","order_by":5,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1454847,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage11.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/bcb652d5c8675d3475b0284e.png"},{"id":102294776,"identity":"20e10f33-5751-4793-8968-4fea25a7ee3f","added_by":"auto","created_at":"2026-02-10 09:56:26","extension":"png","order_by":6,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":2644034,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/b5a9ff43078d880b134b87ce.png"},{"id":100818910,"identity":"557713e4-bd2e-4a6e-9bb0-0de45b6051ea","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"jpeg","order_by":7,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1826956,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage3.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/c6c11c254b3699937bb14ef0.jpeg"},{"id":100818904,"identity":"4c90ffe0-7062-43a3-b5b1-62efcb13575a","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"png","order_by":8,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":736490,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/b46d3cccd0691be7e05f2782.png"},{"id":100857855,"identity":"65d6cabc-6ec3-426a-8529-ce228ce75be6","added_by":"auto","created_at":"2026-01-22 07:23:14","extension":"png","order_by":9,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":457447,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/64092b8f7496418d0d2d5a92.png"},{"id":100858618,"identity":"713b98eb-91ce-418d-ad03-ab4e2d49ef15","added_by":"auto","created_at":"2026-01-22 07:24:30","extension":"png","order_by":10,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":2746264,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage6.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/9e31876f083c56675f5a34aa.png"},{"id":100949353,"identity":"14d233de-dfdc-4004-8f96-c9c733712013","added_by":"auto","created_at":"2026-01-23 07:01:04","extension":"png","order_by":11,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":760643,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage7.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/b86ebca547843d63744b6202.png"},{"id":100858090,"identity":"29e68aff-64b5-4696-8431-dd79d8791de5","added_by":"auto","created_at":"2026-01-22 07:23:50","extension":"png","order_by":12,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":603298,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage8.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/cc72b1cb72a15f36c78e070b.png"},{"id":100818911,"identity":"3b6e330a-acf6-442c-aea9-dbc664d8ee28","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"png","order_by":13,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":606121,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage9.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/f433a9c1646a210cd13342d3.png"},{"id":100818908,"identity":"1a32df4c-e8ab-4023-8b47-2d364d694522","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"png","order_by":14,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":655911,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/870dfcf0b74a0e62cf6abb0f.png"},{"id":100818924,"identity":"3b5f18cb-0bc5-43cd-9034-d4ee735af3dc","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"png","order_by":15,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":233280,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage10.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/cf38f05c5354816f4f36cac0.png"},{"id":100857858,"identity":"307a4489-9240-49b4-b01d-e217ec803db0","added_by":"auto","created_at":"2026-01-22 07:23:16","extension":"png","order_by":16,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":233478,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage11.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/4768b1c8c8020c0da78b1212.png"},{"id":100858132,"identity":"80b87de4-46fa-413a-9520-ff3719b28f98","added_by":"auto","created_at":"2026-01-22 07:23:54","extension":"png","order_by":17,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":523261,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/19f1e1f98439ade81bb6f4f7.png"},{"id":100858033,"identity":"96610f3e-9a05-4492-9807-cddf9e1ec3d6","added_by":"auto","created_at":"2026-01-22 07:23:44","extension":"png","order_by":18,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":915442,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/319970a64f9075e962f0aace.png"},{"id":100818923,"identity":"d1056608-c645-4140-81f9-cff01cbf064d","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"png","order_by":19,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":137779,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/6fe2722dc3d8074725456ec8.png"},{"id":100818912,"identity":"ec7f72b4-7066-4254-af73-3012e74e2618","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"png","order_by":20,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":71056,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/8c29f6a55629688d12d03383.png"},{"id":100818925,"identity":"70d4be1e-83e1-45ae-85ed-65d0e43b840e","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"png","order_by":21,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":474871,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage6.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/6cf56446f716eb6f952328ae.png"},{"id":100818929,"identity":"3fab46e1-2bd4-4710-a036-a8b70db77a44","added_by":"auto","created_at":"2026-01-21 17:21:59","extension":"png","order_by":22,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":156395,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage7.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/d1ada464dc024e3c5b473693.png"},{"id":100818928,"identity":"12ad253d-fac0-438a-af5b-90cdcb93625f","added_by":"auto","created_at":"2026-01-21 17:21:59","extension":"png","order_by":23,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":118665,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage8.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/0c1e63a46d9768c94985ed93.png"},{"id":100818919,"identity":"55fe6523-1a6c-4159-822c-44fc638542e5","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"png","order_by":24,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":99773,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage9.png","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/1fd7979f26da46ca16069acd.png"},{"id":100818931,"identity":"0be614e6-ad5d-470b-b8c8-2a8f3dde62d9","added_by":"auto","created_at":"2026-01-21 17:21:59","extension":"xml","order_by":25,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":102353,"visible":true,"origin":"","legend":"","description":"","filename":"NCOMMS26004037T0structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/2c01b0a5a8296bbf73fc4fac.xml"},{"id":100818922,"identity":"f4d49e05-2acf-4ff1-b19e-52600b9a2578","added_by":"auto","created_at":"2026-01-21 17:21:58","extension":"html","order_by":26,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":120738,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/8e306ca2a34537436257c1b9.html"},{"id":102298257,"identity":"19225b24-e4fe-4b5b-a5b2-e3569508eeb7","added_by":"auto","created_at":"2026-02-10 10:36:27","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":4165244,"visible":true,"origin":"","legend":"","description":"","filename":"ManuscriptDec122025ForSubmissionv3yuntao.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1_covered_43ad38f2-3e67-41e4-abad-78771c8ca55c.pdf"},{"id":100818930,"identity":"216730b0-d666-4666-94fb-bd456bb34307","added_by":"auto","created_at":"2026-01-21 17:21:59","extension":"mp4","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":57271125,"visible":true,"origin":"","legend":"Supplementary Video 1","description":"","filename":"SupplementaryVideo1.mp4","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/7cc6424ddcceef1f4278a954.mp4"},{"id":101397597,"identity":"15691239-07e6-4a04-89db-51c1dcf3b174","added_by":"auto","created_at":"2026-01-29 09:31:36","extension":"mp4","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":9627593,"visible":true,"origin":"","legend":"Supplementary Video 2","description":"","filename":"SupplementaryVideo2.mp4","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/522410d1598713b6386c70e0.mp4"},{"id":100818932,"identity":"0e62a433-9717-414f-af1f-77f2764bcee3","added_by":"auto","created_at":"2026-01-21 17:21:59","extension":"mp4","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":77621040,"visible":true,"origin":"","legend":"Supplementary Video 3","description":"","filename":"SupplementaryVideo3.mp4","url":"https://assets-eu.researchsquare.com/files/rs-8363763/v1/9b457e4f741995894c13e2d8.mp4"}],"financialInterests":"There is \u003cb\u003eNO\u003c/b\u003e Competing Interest.","formattedTitle":"IsoNet2 determines cellular structures at submolecular resolution without averaging","fulltext":[],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":false,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":true,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":true,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"nature-portfolio","isNatureJournal":true,"hasQc":false,"allowDirectSubmit":false,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"","title":"Nature Portfolio","twitterHandle":"","acdcEnabled":false,"dfaEnabled":false,"editorialSystem":"ejp","reportingPortfolio":"","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"","lastPublishedDoi":"10.21203/rs.3.rs-8363763/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8363763/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eWe introduce \u003cem\u003eIsoNet2\u003c/em\u003e, an end-to-end self-supervised deep-learning method that directly reconstructs high-quality 3D densities from cryogenic electron tomograms. A unified network simultaneously performs denoising, contrast transfer function correction, and missing-wedge restoration, achieving\u0026thinsp;~\u0026thinsp;20 \u0026Aring; resolution without averaging. A feature-rich GUI enables rapid, dataset-specific fine-tuning for end-users. \u003cem\u003eIsoNet2\u003c/em\u003e resolves domain organization in HIV capsid proteins, tRNA occupancy in individual ribosomes, and \u003cem\u003ein situ\u003c/em\u003e architectures of mitochondrial respiration-related complexes, enabling atomic-level interpretation of cellular environments.\u003c/p\u003e","manuscriptTitle":"IsoNet2 determines cellular structures at submolecular resolution without averaging","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-01-21 17:21:49","doi":"10.21203/rs.3.rs-8363763/v1","editorialEvents":[],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"nature-communications","isNatureJournal":true,"hasQc":false,"allowDirectSubmit":false,"externalIdentity":"NCOMMS","sideBox":"Learn more about [Nature Communications](http://www.nature.com/ncomms/)","snPcode":"","submissionUrl":"https://mts-ncomms.nature.com/","title":"Nature Communications","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"ejp","reportingPortfolio":"Nature Communications","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"f41e6bcd-65b1-4535-95af-ec3e160e587f","owner":[],"postedDate":"January 21st, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[{"id":61436135,"name":"Biological sciences/Biophysics/Computational biophysics"},{"id":61436136,"name":"Biological sciences/Structural biology/Electron microscopy/Cryoelectron tomography"}],"tags":[],"updatedAt":"2026-02-06T16:21:07+00:00","versionOfRecord":[],"versionCreatedAt":"2026-01-21 17:21:49","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8363763","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8363763","identity":"rs-8363763","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}