{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T04:02:31Z","timestamp":1792036951288,"version":"4.4.0"},"posted":{"date-parts":[[2025,3,15]]},"group-title":"Genetics","reference-count":100,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2025,3,15]],"date-time":"2025-03-15T00:00:00Z","timestamp":1741996800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2025,3,16]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                <jats:p>\n                  Transposable elements are abundant in host genomes but are generally considered to be confined to the cell in which they are expressed, with the notable exception of endogenous retroviruses. Here, we identify a group of LTR retrotransposons that infect the germline from somatic cells within the\n                  <jats:italic>Drosophila<\/jats:italic>\n                  ovary, despite lacking the fusogenic Envelope protein typically required for retroviral entry. Instead, these elements encode a short transmembrane protein, sORF2, with structural features reminiscent of viral cell-cell fusogens. Through genetics, imaging, and electron microscopy, we show that sORF2 localizes to invasive somatic protrusions, enabling the direct transfer of retrotransposon capsids into the oocyte. Remarkably, sORF2-like proteins are widespread among insect retrotransposons and also occur in piscine nackednaviruses and avian picornaviruses. These findings reveal a noncanonical, Envelope-independent transmission mechanism shared by retrotransposons and non-enveloped viruses, offering important insights into host-pathogen evolution and soma-germline interactions.\n                <\/jats:p>","DOI":"10.1101\/2025.03.14.642691","type":"posted-content","created":{"date-parts":[[2025,3,18]],"date-time":"2025-03-18T18:20:18Z","timestamp":1742322018000},"source":"Crossref","is-referenced-by-count":10,"title":["Direct cell-to-cell transmission of retrotransposons"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0508-0682","authenticated-orcid":false,"given":"Maya","family":"Voichek","sequence":"first","affiliation":[{"name":"Institute of Molecular Biotechnology of the Austrian Academy of Sciences (IMBA), Vienna BioCenter (VBC); Dr. Bohr-Gasse 3, 1030 Vienna, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andreas","family":"Bernhard","sequence":"additional","affiliation":[{"name":"Institute of Molecular Biotechnology of the Austrian Academy of Sciences (IMBA), Vienna BioCenter (VBC); Dr. Bohr-Gasse 3, 1030 Vienna, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria","family":"Novatchkova","sequence":"additional","affiliation":[{"name":"Institute of Molecular Biotechnology of the Austrian Academy of Sciences (IMBA), Vienna BioCenter (VBC); Dr. Bohr-Gasse 3, 1030 Vienna, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dominik","family":"Handler","sequence":"additional","affiliation":[{"name":"Institute of Molecular Biotechnology of the Austrian Academy of Sciences (IMBA), Vienna BioCenter (VBC); Dr. Bohr-Gasse 3, 1030 Vienna, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paul","family":"M\u00f6seneder","sequence":"additional","affiliation":[{"name":"Institute of Molecular Biotechnology of the Austrian Academy of Sciences (IMBA), Vienna BioCenter (VBC); Dr. Bohr-Gasse 3, 1030 Vienna, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Baptiste","family":"Rafanel","sequence":"additional","affiliation":[{"name":"Institute of Molecular Biotechnology of the Austrian Academy of Sciences (IMBA), Vienna BioCenter (VBC); Dr. Bohr-Gasse 3, 1030 Vienna, Austria"},{"name":"Vienna BioCenter PhD Program, Doctoral School of the University of Vienna and Medical University of Vienna, Vienna, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peter","family":"Duchek","sequence":"additional","affiliation":[{"name":"Institute of Molecular Biotechnology of the Austrian Academy of Sciences (IMBA), Vienna BioCenter (VBC); Dr. Bohr-Gasse 3, 1030 Vienna, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kirsten-Andr\u00e9","family":"Senti","sequence":"additional","affiliation":[{"name":"Institute of Molecular Biotechnology of the Austrian Academy of Sciences (IMBA), Vienna BioCenter (VBC); Dr. Bohr-Gasse 3, 1030 Vienna, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5141-0814","authenticated-orcid":false,"given":"Julius","family":"Brennecke","sequence":"additional","affiliation":[{"name":"Institute of Molecular Biotechnology of the Austrian Academy of Sciences (IMBA), Vienna BioCenter (VBC); Dr. Bohr-Gasse 3, 1030 Vienna, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026041120471282000_2025.03.14.642691v2.1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-024-07487-w"},{"issue":"11","key":"2026041120471282000_2025.03.14.642691v2.2","first-page":"1905","article-title":"[Features of the structural organization of the MDG1 retrotransposon of Drosophila, revealed during its sequencing]","volume":"26","year":"1990","journal-title":"Genetika"},{"key":"2026041120471282000_2025.03.14.642691v2.3","doi-asserted-by":"publisher","DOI":"10.1186\/s12915-017-0404-4"},{"key":"2026041120471282000_2025.03.14.642691v2.4","doi-asserted-by":"publisher","DOI":"10.1186\/s13100-015-0041-9"},{"key":"2026041120471282000_2025.03.14.642691v2.5","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gky761"},{"key":"2026041120471282000_2025.03.14.642691v2.6","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.80067"},{"key":"2026041120471282000_2025.03.14.642691v2.7","doi-asserted-by":"publisher","DOI":"10.3390\/jdb8040035"},{"key":"2026041120471282000_2025.03.14.642691v2.8","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.03807-13"},{"key":"2026041120471282000_2025.03.14.642691v2.9","doi-asserted-by":"publisher","DOI":"10.1186\/s13059-018-1577-z"},{"key":"2026041120471282000_2025.03.14.642691v2.10","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2018.00260"},{"key":"2026041120471282000_2025.03.14.642691v2.11","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2007.01.043"},{"key":"2026041120471282000_2025.03.14.642691v2.12","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2024.07.023"},{"key":"2026041120471282000_2025.03.14.642691v2.13","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2007526118"},{"key":"2026041120471282000_2025.03.14.642691v2.14","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.51358"},{"key":"2026041120471282000_2025.03.14.642691v2.15","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/19.12.3213"},{"key":"2026041120471282000_2025.03.14.642691v2.16","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.biochem.72.121801.161504"},{"key":"2026041120471282000_2025.03.14.642691v2.17","doi-asserted-by":"publisher","DOI":"10.1016\/j.tim.2014.08.005"},{"key":"2026041120471282000_2025.03.14.642691v2.18","doi-asserted-by":"publisher","DOI":"10.1002\/mrd.23645"},{"key":"2026041120471282000_2025.03.14.642691v2.19","doi-asserted-by":"publisher","DOI":"10.1007\/s002390010206"},{"key":"2026041120471282000_2025.03.14.642691v2.20","doi-asserted-by":"publisher","DOI":"10.1101\/2024.02.07.579293"},{"key":"2026041120471282000_2025.03.14.642691v2.21","doi-asserted-by":"publisher","DOI":"10.1038\/nature05954"},{"key":"2026041120471282000_2025.03.14.642691v2.22","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.00813-19"},{"key":"2026041120471282000_2025.03.14.642691v2.23","doi-asserted-by":"publisher","DOI":"10.1021\/acs.jproteome.8b00113"},{"key":"2026041120471282000_2025.03.14.642691v2.24","doi-asserted-by":"publisher","DOI":"10.1021\/pr500202e"},{"key":"2026041120471282000_2025.03.14.642691v2.25","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-virology-092818-015523"},{"key":"2026041120471282000_2025.03.14.642691v2.26","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.1002195"},{"key":"2026041120471282000_2025.03.14.642691v2.27","doi-asserted-by":"publisher","DOI":"10.1091\/mbc.E19-01-0009"},{"key":"2026041120471282000_2025.03.14.642691v2.28","doi-asserted-by":"publisher","DOI":"10.1261\/rna.061184.117"},{"key":"2026041120471282000_2025.03.14.642691v2.29","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btn392"},{"key":"2026041120471282000_2025.03.14.642691v2.30","doi-asserted-by":"publisher","DOI":"10.1534\/g3.114.014126"},{"key":"2026041120471282000_2025.03.14.642691v2.31","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkaa472"},{"key":"2026041120471282000_2025.03.14.642691v2.32","doi-asserted-by":"publisher","DOI":"10.1038\/emboj.2011.308"},{"key":"2026041120471282000_2025.03.14.642691v2.33","doi-asserted-by":"publisher","DOI":"10.1637\/10663-091613-Reg.1"},{"key":"2026041120471282000_2025.03.14.642691v2.34","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1005883"},{"key":"2026041120471282000_2025.03.14.642691v2.35","doi-asserted-by":"publisher","DOI":"10.1007\/s00427-008-0233-0"},{"key":"2026041120471282000_2025.03.14.642691v2.36","doi-asserted-by":"publisher","DOI":"10.1038\/s41579-019-0189-2"},{"key":"2026041120471282000_2025.03.14.642691v2.37","doi-asserted-by":"publisher","DOI":"10.1038\/nmeth1113"},{"key":"2026041120471282000_2025.03.14.642691v2.38","doi-asserted-by":"publisher","DOI":"10.1016\/j.jmb.2004.03.016"},{"key":"2026041120471282000_2025.03.14.642691v2.39","doi-asserted-by":"publisher","DOI":"10.1186\/gb-2002-3-12-research0088"},{"key":"2026041120471282000_2025.03.14.642691v2.40","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0732024100"},{"key":"2026041120471282000_2025.03.14.642691v2.41","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-021-21366-2"},{"key":"2026041120471282000_2025.03.14.642691v2.42","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.91.4.1285"},{"key":"2026041120471282000_2025.03.14.642691v2.43","doi-asserted-by":"publisher","DOI":"10.1242\/jcs.213124"},{"key":"2026041120471282000_2025.03.14.642691v2.44","doi-asserted-by":"publisher","DOI":"10.1242\/jcs.s3-104.67.297"},{"key":"2026041120471282000_2025.03.14.642691v2.45","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2025.02.003"},{"key":"2026041120471282000_2025.03.14.642691v2.46","doi-asserted-by":"publisher","DOI":"10.1006\/jmbi.2000.4315"},{"key":"2026041120471282000_2025.03.14.642691v2.47","doi-asserted-by":"publisher","DOI":"10.1016\/j.chom.2017.07.019"},{"key":"2026041120471282000_2025.03.14.642691v2.48","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.74.22.10658-10669.2000"},{"key":"2026041120471282000_2025.03.14.642691v2.49","doi-asserted-by":"publisher","DOI":"10.1099\/0022-1317-78-9-2379"},{"key":"2026041120471282000_2025.03.14.642691v2.50","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkab301"},{"key":"2026041120471282000_2025.03.14.642691v2.51","doi-asserted-by":"publisher","DOI":"10.1038\/nrg3030"},{"key":"2026041120471282000_2025.03.14.642691v2.52","doi-asserted-by":"publisher","DOI":"10.1099\/jgv.0.001509"},{"key":"2026041120471282000_2025.03.14.642691v2.53","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-62703-646-7_10"},{"key":"2026041120471282000_2025.03.14.642691v2.54","doi-asserted-by":"publisher","DOI":"10.1016\/j.devcel.2022.05.016"},{"key":"2026041120471282000_2025.03.14.642691v2.55","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkae241"},{"key":"2026041120471282000_2025.03.14.642691v2.56","doi-asserted-by":"publisher","DOI":"10.1099\/jgv.0.001415"},{"issue":"7","key":"2026041120471282000_2025.03.14.642691v2.57","first-page":"1016","article-title":"[A small open reading frame of the Stalker retrotransposon reveals a high similarity to the second small frame of the MDG1 retrotransposon]","volume":"33","year":"1997","journal-title":"Genetika"},{"key":"2026041120471282000_2025.03.14.642691v2.58","doi-asserted-by":"publisher","DOI":"10.1101\/gr.145000"},{"key":"2026041120471282000_2025.03.14.642691v2.59","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-45963-z"},{"key":"2026041120471282000_2025.03.14.642691v2.60","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2009.03.040"},{"key":"2026041120471282000_2025.03.14.642691v2.61","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2009.01.045"},{"key":"2026041120471282000_2025.03.14.642691v2.62","doi-asserted-by":"publisher","DOI":"10.1016\/j.jsb.2005.07.007"},{"key":"2026041120471282000_2025.03.14.642691v2.63","doi-asserted-by":"publisher","DOI":"10.1016\/j.jsb.2016.07.011"},{"key":"2026041120471282000_2025.03.14.642691v2.64","doi-asserted-by":"publisher","DOI":"10.1093\/bib\/bbs012"},{"key":"2026041120471282000_2025.03.14.642691v2.65","doi-asserted-by":"publisher","DOI":"10.1099\/vir.0.79911-0"},{"key":"2026041120471282000_2025.03.14.642691v2.66","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msi060"},{"key":"2026041120471282000_2025.03.14.642691v2.67","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0607435103"},{"key":"2026041120471282000_2025.03.14.642691v2.68","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.201111116"},{"key":"2026041120471282000_2025.03.14.642691v2.69","doi-asserted-by":"publisher","DOI":"10.1038\/emboj.2010.212"},{"key":"2026041120471282000_2025.03.14.642691v2.70","doi-asserted-by":"publisher","DOI":"10.1186\/s12915-021-01158-2"},{"key":"2026041120471282000_2025.03.14.642691v2.71","doi-asserted-by":"publisher","DOI":"10.1002\/j.1460-2075.1994.tb06760.x"},{"key":"2026041120471282000_2025.03.14.642691v2.72","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkae1011"},{"key":"2026041120471282000_2025.03.14.642691v2.73","doi-asserted-by":"publisher","DOI":"10.1002\/pro.3943"},{"key":"2026041120471282000_2025.03.14.642691v2.74","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-37068-w"},{"key":"2026041120471282000_2025.03.14.642691v2.75","doi-asserted-by":"publisher","DOI":"10.1038\/nature08501"},{"key":"2026041120471282000_2025.03.14.642691v2.76","doi-asserted-by":"publisher","DOI":"10.1016\/j.devcel.2007.12.008"},{"key":"2026041120471282000_2025.03.14.642691v2.77","doi-asserted-by":"publisher","DOI":"10.1016\/j.virol.2024.110258"},{"key":"2026041120471282000_2025.03.14.642691v2.78","doi-asserted-by":"publisher","DOI":"10.1016\/j.semcdb.2022.03.031"},{"key":"2026041120471282000_2025.03.14.642691v2.79","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2313866121"},{"key":"2026041120471282000_2025.03.14.642691v2.80","doi-asserted-by":"publisher","DOI":"10.1242\/jcs.02831"},{"key":"2026041120471282000_2025.03.14.642691v2.81","doi-asserted-by":"publisher","DOI":"10.1038\/nature10098"},{"key":"2026041120471282000_2025.03.14.642691v2.82","doi-asserted-by":"publisher","DOI":"10.1101\/2023.08.03.551782"},{"key":"2026041120471282000_2025.03.14.642691v2.83","doi-asserted-by":"publisher","DOI":"10.1126\/science.1234781"},{"key":"2026041120471282000_2025.03.14.642691v2.84","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.77.18.9769-9779.2003"},{"key":"2026041120471282000_2025.03.14.642691v2.85","unstructured":"Smit, A.F.A. Hubley, R. , & Green, P. (2013). RepeatMasker Open-4.0 [Computer software]. http:\/\/www.repeatmasker.org"},{"key":"2026041120471282000_2025.03.14.642691v2.86","doi-asserted-by":"publisher","DOI":"10.1101\/gad.8.17.2046"},{"key":"2026041120471282000_2025.03.14.642691v2.87","doi-asserted-by":"publisher","DOI":"10.1186\/s13100-020-00230-y"},{"key":"2026041120471282000_2025.03.14.642691v2.88","doi-asserted-by":"publisher","DOI":"10.1021\/pr200611n"},{"key":"2026041120471282000_2025.03.14.642691v2.89","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.72.1.853-856.1998"},{"key":"2026041120471282000_2025.03.14.642691v2.90","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.57.2.551"},{"key":"2026041120471282000_2025.03.14.642691v2.91","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1009635107"},{"key":"2026041120471282000_2025.03.14.642691v2.92","doi-asserted-by":"publisher","DOI":"10.1101\/2023.04.07.536061"},{"key":"2026041120471282000_2025.03.14.642691v2.93","doi-asserted-by":"publisher","DOI":"10.1038\/s41423-021-00663-2"},{"key":"2026041120471282000_2025.03.14.642691v2.94","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btp033"},{"key":"2026041120471282000_2025.03.14.642691v2.95","doi-asserted-by":"publisher","DOI":"10.1126\/science.ade0705"},{"key":"2026041120471282000_2025.03.14.642691v2.96","doi-asserted-by":"publisher","DOI":"10.1038\/srep28249"},{"key":"2026041120471282000_2025.03.14.642691v2.97","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-41733-5"},{"key":"2026041120471282000_2025.03.14.642691v2.98","doi-asserted-by":"publisher","DOI":"10.1111\/j.1432-1033.1986.tb09767.x"},{"key":"2026041120471282000_2025.03.14.642691v2.99","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1313677110"},{"key":"2026041120471282000_2025.03.14.642691v2.100","doi-asserted-by":"publisher","DOI":"10.1186\/s12985-018-1008-5"}],"container-title":[],"original-title":[],"link":[{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.1101\/2025.03.14.642691","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,4,12]],"date-time":"2026-04-12T03:47:17Z","timestamp":1775965637000},"score":1,"resource":{"primary":{"URL":"http:\/\/biorxiv.org\/lookup\/doi\/10.1101\/2025.03.14.642691"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,3,15]]},"references-count":100,"URL":"https:\/\/doi.org\/10.1101\/2025.03.14.642691","relation":{},"subject":[],"published":{"date-parts":[[2025,3,15]]},"subtype":"preprint"}}