{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,12]],"date-time":"2026-07-12T07:07:52Z","timestamp":1783840072420,"version":"3.55.0"},"reference-count":77,"publisher":"Wiley","issue":"5","license":[{"start":{"date-parts":[[2026,6,12]],"date-time":"2026-06-12T00:00:00Z","timestamp":1781222400000},"content-version":"vor","delay-in-days":11,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,6,1]],"date-time":"2026-06-01T00:00:00Z","timestamp":1780272000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["DGE 2139754"],"award-info":[{"award-number":["DGE 2139754"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100006510","name":"Duke University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100006510","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The Plant Journal"],"published-print":{"date-parts":[[2026,6]]},"abstract":"<jats:title>SUMMARY<\/jats:title>\n                  <jats:p>\n                    RNA editing is a post\u2010transcriptional pyrimidine exchange process that alters plastid and mitochondrial transcripts in nearly all land plants. Although confined to organelles, it is directed by nuclear\u2010encoded PLS\u2010type pentatricopeptide repeat (PPR) proteins, each typically recognizing a specific RNA target. While many editing sites are functionally neutral, edits at cryptic start and internal stop codons have been implicated in modulating organellar gene expression. Ferns\u2014and some lycophytes\u2014are unique among vascular plants in exhibiting both C\u2010to\u2010U and U\u2010to\u2010C editing, making them valuable for studying the evolution of both forms. Here, we examine chloroplast RNA editing in four Schizaeales species (\n                    <jats:italic>Schizaea dichotoma<\/jats:italic>\n                    ,\n                    <jats:italic>Actinostachys digitata<\/jats:italic>\n                    ,\n                    <jats:italic>Anemia phyllitidis<\/jats:italic>\n                    ,\n                    <jats:italic>Lygodium microphyllum<\/jats:italic>\n                    ).\n                    <jats:italic>Schizaea<\/jats:italic>\n                    and\n                    <jats:italic>Actinostachys<\/jats:italic>\n                    possess non\u2010photosynthetic gametophytes, providing a natural contrast with fully photosynthetic relatives. Despite extensive plastome reduction, including loss of the\n                    <jats:italic>ndh<\/jats:italic>\n                    suite and, in\n                    <jats:italic>Actinostachys<\/jats:italic>\n                    , all\n                    <jats:italic>chl<\/jats:italic>\n                    genes,\n                    <jats:italic>Schizaea<\/jats:italic>\n                    and\n                    <jats:italic>Actinostachys<\/jats:italic>\n                    exhibit dramatically elevated numbers of C\u2010to\u2010U edits. Genes evolving under relaxed selection accumulate more editing sites, and editing abundance per gene correlates with the magnitude of relaxed constraint, suggesting relaxed selection promotes edit proliferation.\n                    <jats:italic>Schizaea dichotoma<\/jats:italic>\n                    and\n                    <jats:italic>A.\u00a0digitata<\/jats:italic>\n                    also show expansion of the chloroplast inverted repeat (IR), and genes translocated into the IR exhibit reduced substitution rates and higher editing densities, indicating that IR expansion slows the loss of edits. Finally, annotation of PPR proteins revealed few full\u2010length editing factors, consistent with catalytic domains assembling\n                    <jats:italic>in trans<\/jats:italic>\n                    and highlighting the modular nature of the fern editosome.\n                  <\/jats:p>","DOI":"10.1111\/tpj.70919","type":"journal-article","created":{"date-parts":[[2026,6,12]],"date-time":"2026-06-12T22:10:41Z","timestamp":1781302241000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Losing genes, gaining edits: how relaxed selection and inverted repeat expansion shape\n                    <scp>RNA<\/scp>\n                    editing in Schizaeaceae plastomes"],"prefix":"10.1111","volume":"126","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5251-0959","authenticated-orcid":false,"given":"Blake D.","family":"Fauskee","sequence":"first","affiliation":[{"name":"Department of Biology Duke University  Durham 27708 North Carolina USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3388-3757","authenticated-orcid":false,"given":"Li\u2010Yaung","family":"Kuo","sequence":"additional","affiliation":[{"name":"Institute of Molecular and Cellular Biology National Tsing Hua University  Hsinchu Taiwan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8640-190X","authenticated-orcid":false,"given":"Farley","family":"Kwok van der Giezen","sequence":"additional","affiliation":[{"name":"ARC Centre of Excellence in Plant Energy Biology, School of Molecular Sciences The University of Western Australia  Perth Western Australia 6009 Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9776-6736","authenticated-orcid":false,"given":"Kathleen M.","family":"Pryer","sequence":"additional","affiliation":[{"name":"Department of Biology Duke University  Durham 27708 North Carolina USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,6,12]]},"reference":[{"key":"e_1_2_8_2_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1002910"},{"key":"e_1_2_8_3_1","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msu252"},{"key":"e_1_2_8_4_1","doi-asserted-by":"publisher","DOI":"10.1093\/gbe\/evv251"},{"key":"e_1_2_8_5_1","doi-asserted-by":"publisher","DOI":"10.1002\/j.1537-2197.1975.tb14055.x"},{"key":"e_1_2_8_6_1","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btu170"},{"key":"e_1_2_8_7_1","doi-asserted-by":"publisher","DOI":"10.7717\/peerj.1660"},{"key":"e_1_2_8_8_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.112.099507"},{"key":"e_1_2_8_9_1","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btx063"},{"key":"e_1_2_8_10_1","doi-asserted-by":"publisher","DOI":"10.1093\/gigascience\/giz100"},{"key":"e_1_2_8_11_1","unstructured":"Bushnell B.(2014)BBMap: a fast accurate splice\u2010aware aligner.9th Annual Genomics of Energy and Environment Meeting Walnut Creek CA USA."},{"key":"e_1_2_8_12_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-313X.2008.03634.x"},{"key":"e_1_2_8_13_1","doi-asserted-by":"publisher","DOI":"10.1002\/wrna.72"},{"key":"e_1_2_8_14_1","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btx364"},{"key":"e_1_2_8_15_1","doi-asserted-by":"publisher","DOI":"10.1016\/0168-9525(93)90011-6"},{"key":"e_1_2_8_16_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.107.054841"},{"key":"e_1_2_8_17_1","doi-asserted-by":"crossref","unstructured":"Dennis M. 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