{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,1,15]],"date-time":"2026-01-15T19:57:13Z","timestamp":1768507033051,"version":"3.49.0"},"posted":{"date-parts":[[2023,5,30]]},"group-title":"Genetics","reference-count":77,"publisher":"openRxiv","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2023,11,3]]},"abstract":"<jats:title>Summary<\/jats:title>\n                <jats:p>\n                  Epigenetic modifications that arise during plant and animal development, such as DNA and histone modification, are mostly reset during gamete formation, but some are inherited from the germline including those marking imprinted genes\n                  <jats:sup>1<\/jats:sup>\n                  . Small RNAs guide these epigenetic modifications, and some are also inherited by the next generation\n                  <jats:sup>2,3<\/jats:sup>\n                  . In\n                  <jats:italic>C. elegans<\/jats:italic>\n                  , these inherited small RNAs have poly (UG) tails\n                  <jats:sup>4<\/jats:sup>\n                  , but how inherited small RNAs are distinguished in other animals and plants is unknown. Pseudouridine (\u03a8) is the most abundant RNA modification but has not been explored in small RNAs. Here, we develop novel assays to detect \u03a8 in short RNA sequences, demonstrating its presence in mouse and\n                  <jats:italic>Arabidopsis<\/jats:italic>\n                  microRNAs and their precursors. We also detect substantial enrichment in germline small RNAs, namely epigenetically activated siRNAs (easiRNAs) in\n                  <jats:italic>Arabidopsis<\/jats:italic>\n                  pollen, and piwi-interacting piRNAs in mouse testis. In pollen, pseudouridylated easiRNAs are localized to sperm cells, and we found that\n                  <jats:italic>PAUSED\/HEN5 (PSD)<\/jats:italic>\n                  , the plant homolog of Exportin-t, interacts genetically with \u03a8 and is required for transport of easiRNAs into sperm cells from the vegetative nucleus. We further show that Exportin-t is required for the triploid block: chromosome dosage-dependent seed lethality that is epigenetically inherited from pollen. Thus, \u03a8 has a conserved role in marking inherited small RNAs in the germline.\n                <\/jats:p>\n                <jats:sec>\n                  <jats:title>One-Sentence Summary<\/jats:title>\n                  <jats:p>Pseudouridine marks germline small RNAs in plants and mammals, impacting epigenetic inheritance via nuclear transport.<\/jats:p>\n                <\/jats:sec>","DOI":"10.1101\/2023.05.27.542553","type":"posted-content","created":{"date-parts":[[2023,5,30]],"date-time":"2023-05-30T04:40:15Z","timestamp":1685421615000},"source":"Crossref","is-referenced-by-count":2,"title":["Pseudouridine guides germline small RNA transport and epigenetic inheritance"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4264-9292","authenticated-orcid":false,"given":"Rowan P","family":"Herridge","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6141-8863","authenticated-orcid":false,"given":"Jakub","family":"Dolata","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4978-2245","authenticated-orcid":false,"given":"Valentina","family":"Migliori","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Cristiane","family":"de Santis Alves","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7388-2118","authenticated-orcid":false,"given":"Filipe","family":"Borges","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Andrea J","family":"Schorn","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Fr\u00e9d\u00e9ric","family":"Van Ex","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7859-7415","authenticated-orcid":false,"given":"Jean-Sebastien","family":"Parent","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ann","family":"Lin","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7043-2827","authenticated-orcid":false,"given":"Mateusz","family":"Bajczyk","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4449-1863","authenticated-orcid":false,"given":"Tommaso","family":"Leonardi","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8604-0462","authenticated-orcid":false,"given":"Alan","family":"Hendrick","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8918-4162","authenticated-orcid":false,"given":"Tony","family":"Kouzarides","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1285-9608","authenticated-orcid":false,"given":"Robert A","family":"Martienssen","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"54368","reference":[{"key":"2023110803201399000_2023.05.27.542553v4.1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-arplant-042811-105514"},{"key":"2023110803201399000_2023.05.27.542553v4.2","doi-asserted-by":"publisher","DOI":"10.1038\/s41556-018-0087-2"},{"key":"2023110803201399000_2023.05.27.542553v4.3","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-018-07429-x"},{"key":"2023110803201399000_2023.05.27.542553v4.4","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-020-2323-8"},{"key":"2023110803201399000_2023.05.27.542553v4.5","doi-asserted-by":"publisher","DOI":"10.1038\/nature04917"},{"key":"2023110803201399000_2023.05.27.542553v4.6","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2008.12.038"},{"key":"2023110803201399000_2023.05.27.542553v4.7","doi-asserted-by":"crossref","unstructured":"Long, J. , et al. Nurse cell\u2013derived small RNAs define paternal epigenetic inheritance in Arabidopsis. 373, eabh0556 (2021).","DOI":"10.1126\/science.abh0556"},{"key":"2023110803201399000_2023.05.27.542553v4.8","doi-asserted-by":"publisher","DOI":"10.1038\/nplants.2016.167"},{"key":"2023110803201399000_2023.05.27.542553v4.9","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-017-0032-5"},{"key":"2023110803201399000_2023.05.27.542553v4.10","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2019.0417"},{"key":"2023110803201399000_2023.05.27.542553v4.11","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-017-0033-4"},{"key":"2023110803201399000_2023.05.27.542553v4.12","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1418918112"},{"key":"2023110803201399000_2023.05.27.542553v4.13","doi-asserted-by":"publisher","DOI":"10.1016\/j.molcel.2021.09.012"},{"key":"2023110803201399000_2023.05.27.542553v4.14","doi-asserted-by":"publisher","DOI":"10.1038\/nature13802"},{"key":"2023110803201399000_2023.05.27.542553v4.15","doi-asserted-by":"publisher","DOI":"10.1038\/nchembio.1836"},{"key":"2023110803201399000_2023.05.27.542553v4.16","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2014.08.028"},{"key":"2023110803201399000_2023.05.27.542553v4.17","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0110799"},{"key":"2023110803201399000_2023.05.27.542553v4.18","doi-asserted-by":"publisher","DOI":"10.1261\/rna.078940.121"},{"key":"2023110803201399000_2023.05.27.542553v4.19","doi-asserted-by":"publisher","DOI":"10.1038\/s41587-022-01505-w"},{"key":"2023110803201399000_2023.05.27.542553v4.20","first-page":"e104708","volume":"39","year":"2020","journal-title":"The tRNA pseudouridine synthase TruB1 regulates the maturation of let-7 miRNA"},{"key":"2023110803201399000_2023.05.27.542553v4.21","doi-asserted-by":"publisher","DOI":"10.1126\/science.1154040"},{"key":"2023110803201399000_2023.05.27.542553v4.22","first-page":"1","article-title":"Meiosis-specific gene discovery in plants: RNA-Seq applied to isolated Arabidopsis male meiocytes","volume":"10","year":"2010","journal-title":"BMC plant biology"},{"key":"2023110803201399000_2023.05.27.542553v4.23","doi-asserted-by":"crossref","first-page":"101","DOI":"10.4161\/rna.4.2.5177","article-title":"Differential roles of archaeal box H\/ACA proteins in guide RNA-dependent and independent pseudouridine formation","volume":"4","year":"2007","journal-title":"RNA biology"},{"key":"2023110803201399000_2023.05.27.542553v4.24","doi-asserted-by":"publisher","DOI":"10.1101\/gad.12.4.527"},{"key":"2023110803201399000_2023.05.27.542553v4.25","doi-asserted-by":"publisher","DOI":"10.1017\/S1355838298980761"},{"key":"2023110803201399000_2023.05.27.542553v4.26","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2006.05.032"},{"key":"2023110803201399000_2023.05.27.542553v4.27","doi-asserted-by":"publisher","DOI":"10.1080\/15476286.2018.1462654"},{"key":"2023110803201399000_2023.05.27.542553v4.28","doi-asserted-by":"publisher","DOI":"10.1128\/MCB.01490-07"},{"key":"2023110803201399000_2023.05.27.542553v4.29","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erz273"},{"key":"2023110803201399000_2023.05.27.542553v4.30","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.109.072199"},{"key":"2023110803201399000_2023.05.27.542553v4.31","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2008.02.034"},{"key":"2023110803201399000_2023.05.27.542553v4.32","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2008.02.033"},{"key":"2023110803201399000_2023.05.27.542553v4.33","doi-asserted-by":"publisher","DOI":"10.1093\/plphys\/kiab474"},{"key":"2023110803201399000_2023.05.27.542553v4.34","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erq452"},{"key":"2023110803201399000_2023.05.27.542553v4.35","doi-asserted-by":"publisher","DOI":"10.1093\/plcell\/koab280"},{"key":"2023110803201399000_2023.05.27.542553v4.36","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M107141200"},{"key":"2023110803201399000_2023.05.27.542553v4.37","doi-asserted-by":"publisher","DOI":"10.1038\/nature08394"},{"key":"2023110803201399000_2023.05.27.542553v4.38","doi-asserted-by":"publisher","DOI":"10.1016\/j.tcb.2018.05.006"},{"key":"2023110803201399000_2023.05.27.542553v4.39","doi-asserted-by":"publisher","DOI":"10.1104\/pp.103.023309"},{"key":"2023110803201399000_2023.05.27.542553v4.40","doi-asserted-by":"publisher","DOI":"10.1104\/pp.103.023291"},{"key":"2023110803201399000_2023.05.27.542553v4.41","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2006.03.064"},{"key":"2023110803201399000_2023.05.27.542553v4.42","unstructured":"Brioudes, F. et al. HASTY, the Arabidopsis EXPORTIN5 ortholog, regulates cell-to-cell and vascular microRNA movement. The EMBO Journal, e107455 (2021)."},{"key":"2023110803201399000_2023.05.27.542553v4.43","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2012.09.001"},{"key":"2023110803201399000_2023.05.27.542553v4.44","doi-asserted-by":"publisher","DOI":"10.21203\/rs.3.rs-2947716\/v1"},{"key":"2023110803201399000_2023.05.27.542553v4.45","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0405570102"},{"key":"2023110803201399000_2023.05.27.542553v4.46","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.15.01056"},{"key":"2023110803201399000_2023.05.27.542553v4.47","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-20606-1"},{"key":"2023110803201399000_2023.05.27.542553v4.48","doi-asserted-by":"publisher","DOI":"10.1111\/nph.16871"},{"key":"2023110803201399000_2023.05.27.542553v4.49","doi-asserted-by":"publisher","DOI":"10.1038\/nature08828"},{"key":"2023110803201399000_2023.05.27.542553v4.50","doi-asserted-by":"publisher","DOI":"10.1101\/gad.343871.120"},{"key":"2023110803201399000_2023.05.27.542553v4.51","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0911341107"},{"key":"2023110803201399000_2023.05.27.542553v4.52","doi-asserted-by":"publisher","DOI":"10.1126\/science.aad6780"},{"key":"2023110803201399000_2023.05.27.542553v4.53","doi-asserted-by":"publisher","DOI":"10.1126\/science.aad7977"},{"key":"2023110803201399000_2023.05.27.542553v4.54","doi-asserted-by":"publisher","DOI":"10.3389\/fgene.2019.00096"},{"key":"2023110803201399000_2023.05.27.542553v4.55","doi-asserted-by":"publisher","DOI":"10.1002\/1873-3468.14188"},{"key":"2023110803201399000_2023.05.27.542553v4.56","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.1002326"},{"key":"2023110803201399000_2023.05.27.542553v4.57","doi-asserted-by":"publisher","DOI":"10.1242\/dev.00362"},{"key":"2023110803201399000_2023.05.27.542553v4.58","doi-asserted-by":"publisher","DOI":"10.2144\/000113941"},{"key":"2023110803201399000_2023.05.27.542553v4.59","doi-asserted-by":"crossref","unstructured":"Li, X . Infiltration of Nicotiana benthamiana protocol for transient expression via Agrobacterium. Bio-protocol, e95\u2013e95 (2011).","DOI":"10.21769\/BioProtoc.95"},{"key":"2023110803201399000_2023.05.27.542553v4.60","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btp616"},{"key":"2023110803201399000_2023.05.27.542553v4.61","doi-asserted-by":"publisher","DOI":"10.1186\/s13059-014-0550-8"},{"key":"2023110803201399000_2023.05.27.542553v4.62","doi-asserted-by":"publisher","DOI":"10.1002\/cbic.201500013"},{"key":"2023110803201399000_2023.05.27.542553v4.63","doi-asserted-by":"publisher","DOI":"10.1038\/nprot.2008.67"},{"key":"2023110803201399000_2023.05.27.542553v4.64","unstructured":"(!!! INVALID CITATION !!! {})."},{"key":"2023110803201399000_2023.05.27.542553v4.65","doi-asserted-by":"publisher","DOI":"10.1186\/1746-4811-8-1"},{"key":"2023110803201399000_2023.05.27.542553v4.66","doi-asserted-by":"publisher","DOI":"10.3791\/56441"},{"key":"2023110803201399000_2023.05.27.542553v4.67","doi-asserted-by":"publisher","DOI":"10.1002\/0471250953.bi1107s32"},{"key":"2023110803201399000_2023.05.27.542553v4.68","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btq033"},{"key":"2023110803201399000_2023.05.27.542553v4.69","unstructured":"Marker, C. et al. Experimental RNomics: identification of 140 candidates for small non-messenger RNAs in the plant Arabidopsis thaliana. Current Biology 12 (2002)."},{"key":"2023110803201399000_2023.05.27.542553v4.70","doi-asserted-by":"crossref","first-page":"D178","DOI":"10.1093\/nar\/gkm855","article-title":"The 3D rRNA modification maps database: with interactive tools for ribosome analysis","volume":"36","year":"2007","journal-title":"Nucleic acids research"},{"key":"2023110803201399000_2023.05.27.542553v4.71","doi-asserted-by":"publisher","DOI":"10.14806\/ej.17.1.200"},{"key":"2023110803201399000_2023.05.27.542553v4.72","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkw427"},{"key":"2023110803201399000_2023.05.27.542553v4.73","doi-asserted-by":"publisher","DOI":"10.7717\/peerj.1332"},{"key":"2023110803201399000_2023.05.27.542553v4.74","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btp324"},{"key":"2023110803201399000_2023.05.27.542553v4.75","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btp352"},{"key":"2023110803201399000_2023.05.27.542553v4.76","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gku302"},{"key":"2023110803201399000_2023.05.27.542553v4.77","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2017.06.013"}],"container-title":[],"original-title":[],"link":[{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.1101\/2023.05.27.542553","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,15]],"date-time":"2026-01-15T04:42:12Z","timestamp":1768452132000},"score":1,"resource":{"primary":{"URL":"http:\/\/biorxiv.org\/lookup\/doi\/10.1101\/2023.05.27.542553"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,5,30]]},"references-count":77,"URL":"https:\/\/doi.org\/10.1101\/2023.05.27.542553","relation":{"is-preprint-of":[{"id-type":"doi","id":"10.1038\/s41594-024-01392-6","asserted-by":"subject"}]},"subject":[],"published":{"date-parts":[[2023,5,30]]},"subtype":"preprint"}}