{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T00:45:18Z","timestamp":1785199518804,"version":"3.55.0"},"reference-count":97,"publisher":"Oxford University Press (OUP)","issue":"5","license":[{"start":{"date-parts":[[2025,5,1]],"date-time":"2025-05-01T00:00:00Z","timestamp":1746057600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"funder":[{"name":"Rosen Center for Advanced Computing at Purdue University"},{"DOI":"10.13039\/100006377","name":"Purdue University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100006377","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100022786","name":"Center for Plant Biology","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100022786","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100008982","name":"National Science Foundation","doi-asserted-by":"publisher","award":["DEB-1831493"],"award-info":[{"award-number":["DEB-1831493"]}],"id":[{"id":"10.13039\/501100008982","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100008982","name":"National Science Foundation","doi-asserted-by":"publisher","award":["IOS-1931114"],"award-info":[{"award-number":["IOS-1931114"]}],"id":[{"id":"10.13039\/501100008982","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2025,5,9]]},"abstract":"<jats:title>Abstract<\/jats:title>\n               <jats:p>Most ferns, unlike all seed plants, are homosporous and produce sexually undifferentiated spores. Sex ratio in many homosporous species is environmentally established by the secretion of antheridiogen from female\/hermaphrodite gametophytes. Nearby undetermined gametophytes perceive antheridiogen, which induces male development. In the fern Ceratopteris richardii (Ceratopteris), hermaphroditic (her) mutants develop as hermaphrodites even in the presence of antheridiogen. Modern sequencing and genomic tools make the molecular identification of mutants in the 11-Gbp genome of this fern possible. We mapped 2 linked mutants, her7-14 and her7-19, to the same 16-Mbp interval on chromosome 29 of the Ceratopteris genome. An ortholog of the receptor kinase gene BRASSINOSTEROID INSENSITIVE 1 (BRI1) within this interval encoded a deletion mutation in her7-14 and a missense mutation in her7-19. Three other linked her mutants encoded missense mutations in the same gene, which we name HER7. Consistent with a function as a receptor kinase, HER7-GFP fusion protein localized to the plasma membrane and cytoplasm. Analysis of gene expression showed that brassinosteroid biosynthesis was upregulated in hermaphrodites compared with male gametophytes. Our work demonstrates that HER7 is required for sex determination in Ceratopteris and opens avenues for studying the evolution of antheridiogen systems.<\/jats:p>","DOI":"10.1093\/plcell\/koaf058","type":"journal-article","created":{"date-parts":[[2025,5,9]],"date-time":"2025-05-09T12:01:43Z","timestamp":1746792103000},"source":"Crossref","is-referenced-by-count":7,"title":["A BRASSINOSTEROID INSENSISTIVE 1 receptor kinase ortholog is required for sex determination in <i>Ceratopteris richardii<\/i>"],"prefix":"10.1093","volume":"37","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9358-8146","authenticated-orcid":false,"given":"Katelin M","family":"Burow","sequence":"first","affiliation":[{"name":"Department of Biochemistry, Purdue University , West Lafayette, IN 47907 ,","place":["USA"]},{"name":"Purdue Center for Plant Biology, Purdue University , West Lafayette, IN 47907 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0648-1728","authenticated-orcid":false,"given":"Xi","family":"Yang","sequence":"additional","affiliation":[{"name":"Purdue Center for Plant Biology, Purdue University , West Lafayette, IN 47907 ,","place":["USA"]},{"name":"Department of Botany and Plant Pathology, Purdue University , West Lafayette, IN 47907 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4105-5174","authenticated-orcid":false,"given":"Yun","family":"Zhou","sequence":"additional","affiliation":[{"name":"Purdue Center for Plant Biology, Purdue University , West Lafayette, IN 47907 ,","place":["USA"]},{"name":"Department of Botany and Plant Pathology, Purdue University , West Lafayette, IN 47907 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2799-954X","authenticated-orcid":false,"given":"Brian P","family":"Dilkes","sequence":"additional","affiliation":[{"name":"Department of Biochemistry, Purdue University , West Lafayette, IN 47907 ,","place":["USA"]},{"name":"Purdue Center for Plant Biology, Purdue University , West Lafayette, IN 47907 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6843-5906","authenticated-orcid":false,"given":"Jennifer H","family":"Wisecaver","sequence":"additional","affiliation":[{"name":"Department of Biochemistry, Purdue University , West Lafayette, IN 47907 ,","place":["USA"]},{"name":"Purdue Center for Plant Biology, Purdue University , West Lafayette, IN 47907 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2025,5,9]]},"reference":[{"issue":"5757","key":"2025053005272104400_koaf058-B1","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1126\/science.1118642","article-title":"Integration of plant responses to environmentally activated phytohormonal signals","volume":"311","author":"Achard","year":"2006","journal-title":"Science"},{"issue":"7","key":"2025053005272104400_koaf058-B2","doi-asserted-by":"publisher","first-page":"2205","DOI":"10.1534\/g3.118.200292","article-title":"Sex determination in Ceratopteris richardii is accompanied by transcriptome changes that drive epigenetic reprogramming of the young gametophyte","volume":"8","author":"Atallah","year":"2018","journal-title":"G3 (Bethesda)"},{"issue":"7","key":"2025053005272104400_koaf058-B3","doi-asserted-by":"publisher","first-page":"1949","DOI":"10.1242\/dev.120.7.1949","article-title":"Sex-determining genes in the homosporous fern Ceratopteris","volume":"120","author":"Banks","year":"1994","journal-title":"Development"},{"issue":"4","key":"2025053005272104400_koaf058-B4","doi-asserted-by":"publisher","first-page":"1885","DOI":"10.1093\/genetics\/147.4.1885","article-title":"The TRANSFORMER genes of the fern Ceratopteris simultaneously promote meristem and archegonia development and repress antheridia development in the developing gametophyte","volume":"147","author":"Banks","year":"1997","journal-title":"Genetics"},{"issue":"4","key":"2025053005272104400_koaf058-B5","doi-asserted-by":"publisher","first-page":"522","DOI":"10.1086\/297135","article-title":"The programming of sexual phenotype in the homosporous fern Ceratopteris richardii","volume":"154","author":"Banks","year":"1993","journal-title":"Int J Plant Sci"},{"key":"2025053005272104400_koaf058-B6","doi-asserted-by":"publisher","first-page":"164","DOI":"10.1016\/j.compbiomed.2014.07.016","article-title":"LRRsearch: an asynchronous server-based application for the prediction of leucine-rich repeat motifs and an integrative database of NOD-like receptors","volume":"53","author":"Bej","year":"2014","journal-title":"Comput Biol Med"},{"issue":"7","key":"2025053005272104400_koaf058-B7","doi-asserted-by":"publisher","first-page":"e501","DOI":"10.1002\/pld3.501","article-title":"Genetic evidence that brassinosteroids suppress pistils in the maize tassel independent of the jasmonic acid pathway","volume":"7","author":"Best","year":"2023","journal-title":"Plant Direct"},{"issue":"4","key":"2025053005272104400_koaf058-B8","doi-asserted-by":"publisher","first-page":"2633","DOI":"10.1104\/pp.16.00399","article-title":"nana plant2 encodes a maize ortholog of the Arabidopsis brassinosteroid biosynthesis gene DWARF1, identifying developmental interactions between brassinosteroids and gibberellins","volume":"171","author":"Best","year":"2016","journal-title":"Plant Physiol"},{"issue":"2","key":"2025053005272104400_koaf058-B9","doi-asserted-by":"publisher","first-page":"10.1002\/pld3.9","DOI":"10.1002\/pld3.9","article-title":"Phytohormone inhibitor treatments phenocopy brassinosteroid\u2013gibberellin dwarf mutant interactions in maize","volume":"1","author":"Best","year":"2017","journal-title":"Plant Direct"},{"issue":"D1","key":"2025053005272104400_koaf058-B10","doi-asserted-by":"publisher","first-page":"D444","DOI":"10.1093\/nar\/gkae1082","article-title":"InterPro: the protein sequence classification resource in 2025","volume":"53","author":"Blum","year":"2025","journal-title":"Nucleic Acids Res"},{"issue":"15","key":"2025053005272104400_koaf058-B11","doi-asserted-by":"publisher","first-page":"2114","DOI":"10.1093\/bioinformatics\/btu170","article-title":"Trimmomatic: a flexible trimmer for illumina sequence data","volume":"30","author":"Bolger","year":"2014","journal-title":"Bioinformatics"},{"issue":"5","key":"2025053005272104400_koaf058-B12","doi-asserted-by":"publisher","first-page":"525","DOI":"10.1038\/nbt.3519","article-title":"Near-optimal probabilistic RNA-Seq quantification","volume":"34","author":"Bray","year":"2016","journal-title":"Nat Biotechnol"},{"issue":"1","key":"2025053005272104400_koaf058-B13","doi-asserted-by":"publisher","first-page":"44","DOI":"10.1186\/s40104-019-0359-0","article-title":"The GATK joint genotyping workflow is appropriate for calling variants in RNA-Seq experiments","volume":"10","author":"Brouard","year":"2019","journal-title":"J Anim Sci Biotechnol"},{"issue":"1","key":"2025053005272104400_koaf058-B14","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1038\/nmeth.3176","article-title":"Fast and sensitive protein alignment using DIAMOND","volume":"12","author":"Buchfink","year":"2015","journal-title":"Nat Methods"},{"issue":"26","key":"2025053005272104400_koaf058-B15","doi-asserted-by":"publisher","first-page":"9651","DOI":"10.1073\/pnas.1316717111","article-title":"GSK3-like kinases positively modulate abscisic acid signaling through phosphorylating subgroup III SnRK2s in Arabidopsis","volume":"111","author":"Cai","year":"2014","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"3","key":"2025053005272104400_koaf058-B16","doi-asserted-by":"publisher","first-page":"284","DOI":"10.1007\/s003440000032","article-title":"Ceratopteris richardii: a productive model for revealing secrets of signaling and development","volume":"19","author":"Chatterjee","year":"2000","journal-title":"J Plant Growth Regul"},{"issue":"17","key":"2025053005272104400_koaf058-B17","doi-asserted-by":"publisher","first-page":"i884","DOI":"10.1093\/bioinformatics\/bty560","article-title":"Fastp: an ultra-fast all-in-one FASTQ preprocessor","volume":"34","author":"Chen","year":"2018","journal-title":"Bioinformatics"},{"issue":"12","key":"2025053005272104400_koaf058-B18","doi-asserted-by":"publisher","first-page":"e81938","DOI":"10.1371\/journal.pone.0081938","article-title":"Brassinosteroids regulate plant growth through distinct signaling pathways in Selaginella and arabidopsis","volume":"8","author":"Cheon","year":"2013","journal-title":"PLoS One"},{"issue":"1","key":"2025053005272104400_koaf058-B19","doi-asserted-by":"publisher","first-page":"270","DOI":"10.1186\/1471-2229-10-270","article-title":"Arabidopsis brassinosteroid biosynthetic mutant dwarf7-1 exhibits slower rates of cell division and shoot induction","volume":"10","author":"Cheon","year":"2010","journal-title":"BMC Plant Biol"},{"issue":"6","key":"2025053005272104400_koaf058-B20","doi-asserted-by":"publisher","first-page":"396","DOI":"10.1080\/07352689.2013.797856","article-title":"The regulation of brassinosteroid biosynthesis in Arabidopsis","volume":"32","author":"Chung","year":"2013","journal-title":"CRC Crit Rev Plant Sci."},{"issue":"2","key":"2025053005272104400_koaf058-B21","doi-asserted-by":"publisher","first-page":"80","DOI":"10.4161\/fly.19695","article-title":"A program for annotating and predicting the effects of single nucleotide polymorphisms, SnpEff: SNPs in the genome of Drosophila melanogaster strain w1118; iso-2; iso-3","volume":"6","author":"Cingolani","year":"2012","journal-title":"Fly (Austin)"},{"issue":"42","key":"2025053005272104400_koaf058-B22","doi-asserted-by":"publisher","first-page":"5083","DOI":"10.1016\/S0040-4039(00)85138-2","article-title":"Constitution of antheridium-inducing factor of anemia phyllitidis","volume":"27","author":"Corey","year":"1986","journal-title":"Tetrahedron Lett"},{"issue":"2","key":"2025053005272104400_koaf058-B23","doi-asserted-by":"publisher","first-page":"462","DOI":"10.1104\/pp.103.027979","article-title":"A gateway cloning vector set for high-throughput functional analysis of genes in planta","volume":"133","author":"Curtis","year":"2003","journal-title":"Plant Physiol"},{"issue":"2","key":"2025053005272104400_koaf058-B24","doi-asserted-by":"publisher","first-page":"giab008","DOI":"10.1093\/gigascience\/giab008","article-title":"Twelve years of SAMtools and BCFtools","volume":"10","author":"Danecek","year":"2021","journal-title":"Gigascience"},{"issue":"1","key":"2025053005272104400_koaf058-B25","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1093\/bioinformatics\/bts635","article-title":"STAR: ultrafast universal RNA-seq aligner","volume":"29","author":"Dobin","year":"2013","journal-title":"Bioinformatics"},{"issue":"5","key":"2025053005272104400_koaf058-B26","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1111\/j.1438-8677.1951.tb01498.x","article-title":"Eine die antheridienbildung bei farnen f\u00f6rdernde substanz in den prothallien von Pteridium aquilinum (L.) Kuhn","volume":"63","author":"D\u00f6pp","year":"1950","journal-title":"Ber Deutsch Bot Ges"},{"issue":"3","key":"2025053005272104400_koaf058-B27","doi-asserted-by":"publisher","first-page":"359","DOI":"10.1086\/297257","article-title":"Ceratopteris: a model system for studying sex-determining mechanisms in plants","volume":"156","author":"Eberle","year":"1995","journal-title":"Int J Plant Sci"},{"issue":"3","key":"2025053005272104400_koaf058-B28","doi-asserted-by":"publisher","first-page":"973","DOI":"10.1093\/genetics\/142.3.973","article-title":"Genetic interactions among sex-determining genes in the fern Ceratopteris richardii","volume":"142","author":"Eberle","year":"1996","journal-title":"Genetics"},{"key":"2025053005272104400_koaf058-B29","doi-asserted-by":"publisher","first-page":"238","DOI":"10.1186\/s13059-019-1832-y","article-title":"OrthoFinder: phylogenetic orthology inference for comparative genomics","volume":"20","author":"Emms","year":"2019","journal-title":"Genome Biol"},{"issue":"19","key":"2025053005272104400_koaf058-B30","doi-asserted-by":"publisher","first-page":"3047","DOI":"10.1093\/bioinformatics\/btw354","article-title":"MultiQC: summarize analysis results for multiple tools and samples in a single report","volume":"32","author":"Ewels","year":"2016","journal-title":"Bioinformatics"},{"key":"2025053005272104400_koaf058-B31","doi-asserted-by":"publisher","first-page":"718932","DOI":"10.3389\/fpls.2021.718932","article-title":"Sexual and apogamous species of woodferns show different protein and phytohormone profiles","volume":"12","author":"Fern\u00e1ndez","year":"2021","journal-title":"Front Plant Sci"},{"issue":"3","key":"2025053005272104400_koaf058-B32","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1111\/j.1469-8137.2006.01787.x","article-title":"Seed dormancy and the control of germination","volume":"171","author":"Finch-Savage","year":"2006","journal-title":"New Phytol"},{"issue":"11","key":"2025053005272104400_koaf058-B33","doi-asserted-by":"publisher","first-page":"1951","DOI":"10.1105\/tpc.9.11.1951","article-title":"The Arabidopsis deetiolated2 mutant is blocked early in brassinosteroid biosynthesis","volume":"9","author":"Fujioka","year":"1997","journal-title":"Plant Cell"},{"issue":"10","key":"2025053005272104400_koaf058-B34","doi-asserted-by":"publisher","first-page":"1601","DOI":"10.1093\/pcp\/pcae037","article-title":"Multiple roles of brassinosteroid signaling in vascular development","volume":"65","author":"Furuya","year":"2024","journal-title":"Plant Cell Physiol"},{"issue":"7","key":"2025053005272104400_koaf058-B35","doi-asserted-by":"publisher","first-page":"dev162537","DOI":"10.1242\/dev.162537","article-title":"Evolutionary divergence of the sex-determining gene MID uncoupled from the transition to anisogamy in volvocine algae","volume":"145","author":"Geng","year":"2018","journal-title":"Development"},{"issue":"3","key":"2025053005272104400_koaf058-B36","doi-asserted-by":"publisher","first-page":"evab042","DOI":"10.1093\/gbe\/evab042","article-title":"A de novo transcriptome assembly of Ceratopteris richardii provides insights into the evolutionary dynamics of complex gene families in land plants","volume":"13","author":"Geng","year":"2021","journal-title":"Genome Biol Evol"},{"issue":"1","key":"2025053005272104400_koaf058-B37","doi-asserted-by":"publisher","first-page":"1214","DOI":"10.1038\/s41467-020-14989-4","article-title":"A signal cascade originated from epidermis defines apical-basal patterning of Arabidopsis shoot apical meristems","volume":"11","author":"Han","year":"2020","journal-title":"Nat Commun"},{"issue":"49","key":"2025053005272104400_koaf058-B38","doi-asserted-by":"publisher","first-page":"19814","DOI":"10.1073\/pnas.1108359108","article-title":"Brassinosteroid control of sex determination in maize","volume":"108","author":"Hartwig","year":"2011","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"11","key":"2025053005272104400_koaf058-B39","doi-asserted-by":"publisher","first-page":"928","DOI":"10.1093\/biosci\/biw108","article-title":"Sex and the single gametophyte: revising the homosporous vascular plant life cycle in light of contemporary research","volume":"66","author":"Haufler","year":"2016","journal-title":"BioScience"},{"issue":"5475","key":"2025053005272104400_koaf058-B40","doi-asserted-by":"publisher","first-page":"2360","DOI":"10.1126\/science.288.5475.2360","article-title":"Perception of brassinosteroids by the extracellular domain of the receptor kinase BRI1","volume":"288","author":"He","year":"2000","journal-title":"Science"},{"issue":"12","key":"2025053005272104400_koaf058-B41","doi-asserted-by":"publisher","first-page":"523","DOI":"10.1016\/S1360-1385(00)01790-8","article-title":"Gibberellin metabolism: new insights revealed by the genes","volume":"5","author":"Hedden","year":"2000","journal-title":"Trends Plant Sci"},{"issue":"4","key":"2025053005272104400_koaf058-B42","doi-asserted-by":"publisher","first-page":"2065","DOI":"10.1073\/pnas.98.4.2065","article-title":"The CYP88A cytochrome P450, ent-kaurenoic acid oxidase, catalyzes three steps of the gibberellin biosynthesis pathway","volume":"98","author":"Helliwell","year":"2001","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"15","key":"2025053005272104400_koaf058-B43","doi-asserted-by":"publisher","first-page":"9019","DOI":"10.1073\/pnas.95.15.9019","article-title":"Cloning of the Arabidopsis ent-kaurene oxidase gene GA3","volume":"95","author":"Helliwell","year":"1998","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"3","key":"2025053005272104400_koaf058-B44","doi-asserted-by":"publisher","first-page":"888","DOI":"10.1139\/b83-098","article-title":"Abscisic acid blocks antheridiogen-induced antheridium formation in gametophytes of the fern Ceratopteris","volume":"61","author":"Hickok","year":"1983","journal-title":"Can J Bot"},{"issue":"3","key":"2025053005272104400_koaf058-B45","doi-asserted-by":"publisher","first-page":"332","DOI":"10.1086\/297255","article-title":"The biology of the fern ceratopteris and its use as a model system","volume":"156","author":"Hickok","year":"1995","journal-title":"Int J Plant Sci"},{"issue":"8","key":"2025053005272104400_koaf058-B46","doi-asserted-by":"publisher","first-page":"1304","DOI":"10.1002\/j.1537-2197.1987.tb08743.x","article-title":"Ceratopteris richardii: applications for experimental plant biology","volume":"74","author":"Hickok","year":"1987","journal-title":"Am J Bot"},{"issue":"13","key":"2025053005272104400_koaf058-B47","doi-asserted-by":"publisher","first-page":"3488","DOI":"10.1073\/pnas.1714972115","article-title":"Mechanistic basis for the activation of plant membrane receptor kinases by SERK-family coreceptors","volume":"115","author":"Hohmann","year":"2018","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"1","key":"2025053005272104400_koaf058-B48","doi-asserted-by":"publisher","first-page":"607","DOI":"10.1111\/nph.16836","article-title":"Insights into the evolutionary history and widespread occurrence of antheridiogen systems in ferns","volume":"229","author":"Hornych","year":"2021","journal-title":"New Phytol"},{"issue":"11","key":"2025053005272104400_koaf058-B49","doi-asserted-by":"publisher","first-page":"4394","DOI":"10.1105\/tpc.114.130849","article-title":"BRASSINOSTEROID INSENSITIVE2 interacts with ABSCISIC ACID INSENSITIVE5 to mediate the antagonism of brassinosteroids to abscisic acid during seed germination in Arabidopsis","volume":"26","author":"Hu","year":"2014","journal-title":"Plant Cell"},{"issue":"6","key":"2025053005272104400_koaf058-B50","doi-asserted-by":"publisher","first-page":"1635","DOI":"10.1093\/molbev\/msw046","article-title":"ETE 3: reconstruction, analysis, and visualization of phylogenomic data","volume":"33","author":"Huerta-Cepas","year":"2016","journal-title":"Mol Biol Evol"},{"issue":"2","key":"2025053005272104400_koaf058-B51","doi-asserted-by":"publisher","first-page":"134","DOI":"10.1007\/s00344-003-0028-5","article-title":"DELLA proteins and GA signalling in Arabidopsis","volume":"22","author":"Hussain","year":"2003","journal-title":"J Plant Growth Regul"},{"issue":"5","key":"2025053005272104400_koaf058-B52","doi-asserted-by":"publisher","first-page":"1027","DOI":"10.1016\/j.jmb.2004.03.016","article-title":"A combined transmembrane topology and signal peptide prediction method","volume":"338","author":"K\u00e4ll","year":"2004","journal-title":"J Mol Biol"},{"issue":"6","key":"2025053005272104400_koaf058-B53","doi-asserted-by":"publisher","first-page":"587","DOI":"10.1038\/nmeth.4285","article-title":"ModelFinder: fast model selection for accurate phylogenetic estimates","volume":"14","author":"Kalyaanamoorthy","year":"2017","journal-title":"Nat Methods"},{"issue":"4","key":"2025053005272104400_koaf058-B54","doi-asserted-by":"publisher","first-page":"772","DOI":"10.1093\/molbev\/mst010","article-title":"MAFFT multiple sequence alignment software version 7: improvements in performance and usability","volume":"30","author":"Katoh","year":"2013","journal-title":"Mol Biol Evol"},{"issue":"4","key":"2025053005272104400_koaf058-B55","doi-asserted-by":"publisher","first-page":"3072","DOI":"10.1093\/plphys\/kiae147","article-title":"A maize semi-dwarf mutant reveals a GRAS transcription factor involved in brassinosteroid signaling","volume":"195","author":"Kaur","year":"2024","journal-title":"Plant Physiol"},{"issue":"4","key":"2025053005272104400_koaf058-B56","doi-asserted-by":"publisher","first-page":"614","DOI":"10.1016\/j.bbrc.2008.07.073","article-title":"Castasterone is a likely end product of brassinosteroid biosynthetic pathway in rice","volume":"374","author":"Kim","year":"2008","journal-title":"Biochem Biophys Res Commun"},{"issue":"4","key":"2025053005272104400_koaf058-B57","doi-asserted-by":"publisher","first-page":"561","DOI":"10.1016\/j.molcel.2011.05.037","article-title":"The CDG1 kinase mediates brassinosteroid signal transduction from bri1 receptor kinase to BSU1 phosphatase and GSK3-like kinase BIN2","volume":"43","author":"Kim","year":"2011","journal-title":"Mol Cell"},{"issue":"7385","key":"2025053005272104400_koaf058-B58","doi-asserted-by":"publisher","first-page":"419","DOI":"10.1038\/nature10794","article-title":"Brassinosteroid regulates stomatal development by GSK3-mediated inhibition of a MAPK pathway","volume":"482","author":"Kim","year":"2012","journal-title":"Nature"},{"issue":"10","key":"2025053005272104400_koaf058-B59","doi-asserted-by":"publisher","first-page":"2311","DOI":"10.1271\/bbb.65.2311","article-title":"Isolation and identification of antheridiogens in the ferns, Lygodium microphyllum and Lygodium reticulatum","volume":"65","author":"Kurumatani","year":"2001","journal-title":"Biosci Biotechnol Biochem"},{"issue":"14","key":"2025053005272104400_koaf058-B60","doi-asserted-by":"publisher","first-page":"1754","DOI":"10.1093\/bioinformatics\/btp324","article-title":"Fast and accurate short read alignment with Burrows\u2013Wheeler transform","volume":"25","author":"Li","year":"2009","journal-title":"Bioinformatics"},{"issue":"2","key":"2025053005272104400_koaf058-B61","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1016\/s0092-8674(02)00812-7","article-title":"BAK1, an Arabidopsis LRR receptor-like protein kinase, interacts with BRI1 and modulates brassinosteroid signaling","volume":"110","author":"Li","year":"2002","journal-title":"Cell"},{"issue":"12","key":"2025053005272104400_koaf058-B62","doi-asserted-by":"publisher","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol"},{"issue":"1","key":"2025053005272104400_koaf058-B63","doi-asserted-by":"publisher","first-page":"e30798","DOI":"10.1371\/journal.pone.0030798","article-title":"Brd1 gene in maize encodes a brassinosteroid C-6 oxidase","volume":"7","author":"Makarevitch","year":"2012","journal-title":"PLoS One"},{"issue":"9","key":"2025053005272104400_koaf058-B64","doi-asserted-by":"publisher","first-page":"1038","DOI":"10.1038\/s41477-022-01226-7","article-title":"Dynamic genome evolution in a model fern","volume":"8","author":"Marchant","year":"2022","journal-title":"Nat Plants"},{"issue":"1","key":"2025053005272104400_koaf058-B65","doi-asserted-by":"publisher","first-page":"122","DOI":"10.1186\/s13059-016-0974-4","article-title":"The ensembl variant effect predictor","volume":"17","author":"McLaren","year":"2016","journal-title":"Genome Biol"},{"issue":"4","key":"2025053005272104400_koaf058-B66","doi-asserted-by":"publisher","first-page":"448","DOI":"10.1101\/gad.1174204","article-title":"Nuclear protein phosphatases with Kelch-repeat domains modulate the response to brassinosteroids in Arabidopsis","volume":"18","author":"Mora-Garc\u00eda","year":"2004","journal-title":"Genes Dev"},{"issue":"9","key":"2025053005272104400_koaf058-B67","doi-asserted-by":"publisher","first-page":"2763","DOI":"10.1093\/molbev\/msaa141","article-title":"GeneRax: a tool for species-tree-aware maximum likelihood-based gene family tree inference under gene duplication, transfer, and loss","volume":"37","author":"Morel","year":"2020","journal-title":"Mol Biol Evol"},{"issue":"1","key":"2025053005272104400_koaf058-B68","doi-asserted-by":"publisher","first-page":"268","DOI":"10.1093\/molbev\/msu300","article-title":"IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies","volume":"32","author":"Nguyen","year":"2015","journal-title":"Mol Biol Evol"},{"issue":"3","key":"2025053005272104400_koaf058-B69","doi-asserted-by":"publisher","first-page":"743","DOI":"10.1104\/pp.121.3.743","article-title":"Brassinosteroid-insensitive dwarf mutants of arabidopsis accumulate brassinosteroids","volume":"121","author":"Noguchi","year":"1999","journal-title":"Plant Physiol"},{"issue":"6","key":"2025053005272104400_koaf058-B70","doi-asserted-by":"publisher","first-page":"563","DOI":"10.1111\/jse.12229","article-title":"A community-derived classification for extant lycophytes and ferns","volume":"54","author":"PPG","year":"2016","journal-title":"J Syst Evol"},{"issue":"1","key":"2025053005272104400_koaf058-B71","doi-asserted-by":"publisher","first-page":"24","DOI":"10.1038\/nbt.1754","article-title":"Integrative genomics viewer","volume":"29","author":"Robinson","year":"2011","journal-title":"Nat Biotechnol"},{"issue":"6982","key":"2025053005272104400_koaf058-B72","doi-asserted-by":"publisher","first-page":"553","DOI":"10.1038\/nature02361","article-title":"Ferns diversified in the shadow of angiosperms","volume":"428","author":"Schneider","year":"2004","journal-title":"Nature"},{"key":"2025053005272104400_koaf058-B73","doi-asserted-by":"publisher","first-page":"134","DOI":"10.1017\/CBO9780511541827.006","volume-title":"Biology and evolution of ferns and lycophytes","author":"Schneller","year":"2008"},{"issue":"7352","key":"2025053005272104400_koaf058-B74","doi-asserted-by":"publisher","first-page":"472","DOI":"10.1038\/nature10178","article-title":"Structural insight into brassinosteroid perception by BRI1","volume":"474","author":"She","year":"2011","journal-title":"Nature"},{"key":"2025053005272104400_koaf058-B75","doi-asserted-by":"publisher","first-page":"1521","DOI":"10.12688\/f1000research.7563.2","article-title":"Differential analyses for RNA-Seq: transcript-level estimates improve gene-level inferences","volume":"4","author":"Soneson","year":"2015","journal-title":"F1000Res"},{"issue":"2","key":"2025053005272104400_koaf058-B100","doi-asserted-by":"publisher","first-page":"275","DOI":"10.1093\/biostatistics\/kxw041","article-title":"False discovery rates: a new deal","volume":"18","author":"Stephens","year":"2017","journal-title":"Biostatistics"},{"issue":"3","key":"2025053005272104400_koaf058-B76","doi-asserted-by":"publisher","first-page":"1271","DOI":"10.1093\/genetics\/159.3.1271","article-title":"Characterization of mutations that feminize gametophytes of the fern ceratopteris","volume":"159","author":"Strain","year":"2001","journal-title":"Genetics"},{"issue":"3","key":"2025053005272104400_koaf058-B77","doi-asserted-by":"publisher","first-page":"1881","DOI":"10.1104\/pp.17.00118","article-title":"Scanning for new BRI1 mutations via TILLING analysis","volume":"174","author":"Sun","year":"2017","journal-title":"Plant Physiol"},{"issue":"10","key":"2025053005272104400_koaf058-B78","doi-asserted-by":"publisher","first-page":"1509","DOI":"10.1105\/tpc.6.10.1509","article-title":"The Arabidopsis GA1 locus encodes the cyclase ent-kaurene synthetase A of gibberellin biosynthesis","volume":"6","author":"Sun","year":"1994","journal-title":"Plant Cell"},{"issue":"2","key":"2025053005272104400_koaf058-B79","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1093\/oxfordjournals.pcp.a077730","article-title":"Biological activities of the methyl ester of gibberellin A73, a novel and principal antheridiogen in Lygodium japonicum","volume":"30","author":"Takeno","year":"1989","journal-title":"Plant Cell Physiol"},{"issue":"6208","key":"2025053005272104400_koaf058-B80","doi-asserted-by":"publisher","first-page":"469","DOI":"10.1126\/science.1259923","article-title":"Antheridiogen determines sex in ferns via a spatiotemporally split gibberellin synthesis pathway","volume":"346","author":"Tanaka","year":"2014","journal-title":"Science"},{"issue":"5888","key":"2025053005272104400_koaf058-B81","doi-asserted-by":"publisher","first-page":"557","DOI":"10.1126\/science.1156973","article-title":"BSKs mediate signal transduction from the receptor kinase BRI1 in Arabidopsis","volume":"321","author":"Tang","year":"2008","journal-title":"Science"},{"issue":"8","key":"2025053005272104400_koaf058-B82","doi-asserted-by":"publisher","first-page":"4698","DOI":"10.1073\/pnas.96.8.4698","article-title":"Molecular cloning and functional expression of gibberellin 2- oxidases, multifunctional enzymes involved in gibberellin deactivation","volume":"96","author":"Thomas","year":"1999","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"7059","key":"2025053005272104400_koaf058-B83","doi-asserted-by":"publisher","first-page":"693","DOI":"10.1038\/nature04028","article-title":"GIBBERELLIN INSENSITIVE DWARF1 encodes a soluble receptor for gibberellin","volume":"437","author":"Ueguchi-Tanaka","year":"2005","journal-title":"Nature"},{"issue":"6","key":"2025053005272104400_koaf058-B84","doi-asserted-by":"publisher","first-page":"1738","DOI":"10.1093\/mp\/sst096","article-title":"From squalene to brassinolide: the steroid metabolic and signaling pathways across the plant kingdom","volume":"6","author":"Vriet","year":"2013","journal-title":"Mol Plant"},{"issue":"2","key":"2025053005272104400_koaf058-B85","doi-asserted-by":"publisher","first-page":"315","DOI":"10.1016\/j.molp.2017.12.013","article-title":"Abscisic acid signaling inhibits brassinosteroid signaling through dampening the dephosphorylation of BIN2 by ABI1 and ABI2","volume":"11","author":"Wang","year":"2018","journal-title":"Mol Plant"},{"issue":"1","key":"2025053005272104400_koaf058-B86","doi-asserted-by":"publisher","first-page":"701","DOI":"10.1146\/annurev-genet-102209-163450","article-title":"Brassinosteroid signaling network and regulation of photomorphogenesis","volume":"46","author":"Wang","year":"2012","journal-title":"Annu Rev Genet"},{"issue":"6826","key":"2025053005272104400_koaf058-B87","doi-asserted-by":"publisher","first-page":"380","DOI":"10.1038\/35066597","article-title":"BRI1 is a critical component of a plasma-membrane receptor for plant steroids","volume":"410","author":"Wang","year":"2001","journal-title":"Nature"},{"issue":"2","key":"2025053005272104400_koaf058-B88","doi-asserted-by":"publisher","first-page":"535","DOI":"10.1104\/pp.89.2.535","article-title":"Evidence for a gibberellin biosynthetic origin of ceratopteris antheridiogen","volume":"89","author":"Warne","year":"1989","journal-title":"Plant Physiol"},{"issue":"4","key":"2025053005272104400_koaf058-B89","doi-asserted-by":"publisher","first-page":"371","DOI":"10.1111\/j.1095-8339.1988.tb00692.x","article-title":"Characterization and genetic analysis of antheridiogen-insensitive mutants in the fern Ceratopteris","volume":"96","author":"Warne","year":"1988","journal-title":"Bot J Linn Soc"},{"issue":"4","key":"2025053005272104400_koaf058-B90","doi-asserted-by":"publisher","first-page":"1271","DOI":"10.1104\/pp.116.4.1271","article-title":"The GA2 locus of Arabidopsis thaliana encodes ent-kaurene synthase of gibberellin biosynthesis","volume":"116","author":"Yamaguchi","year":"1998","journal-title":"Plant Physiol"},{"key":"2025053005272104400_koaf058-B91","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/S0074-7696(08)62177-4","volume-title":"International review of cytology","author":"Yamane","year":"1998"},{"issue":"3","key":"2025053005272104400_koaf058-B92","doi-asserted-by":"publisher","first-page":"741","DOI":"10.1104\/pp.111.3.741","article-title":"Identification of antheridiogens in Lygodium circinnatum and Lygodium flexuosum","volume":"111","author":"Yamauchi","year":"1996","journal-title":"Plant Physiol"},{"issue":"1","key":"2025053005272104400_koaf058-B93","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1105\/tpc.17.00816","article-title":"Brassinosteroids modulate meristem fate and differentiation of unique inflorescence morphology in Setaria viridis","volume":"30","author":"Yang","year":"2018","journal-title":"Plant Cell"},{"key":"2025053005272104400_koaf058-B94","doi-asserted-by":"publisher","first-page":"46","DOI":"10.1016\/j.phytochem.2016.12.020","article-title":"Occurrence of brassinosteroids in non-flowering land plants, liverwort, moss, lycophyte and fern","volume":"136","author":"Yokota","year":"2017","journal-title":"Phytochemistry"},{"issue":"11","key":"2025053005272104400_koaf058-B95","doi-asserted-by":"publisher","first-page":"4543","DOI":"10.1073\/pnas.0900349106","article-title":"The primary signaling outputs of brassinosteroids are regulated by abscisic acid signaling","volume":"106","author":"Zhang","year":"2009","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"12","key":"2025053005272104400_koaf058-B96","doi-asserted-by":"publisher","first-page":"6795","DOI":"10.3390\/ijms23126795","article-title":"Evolutionary analysis and functional identification of ancient brassinosteroid receptors in Ceratopteris richardii","volume":"23","author":"Zheng","year":"2022","journal-title":"Int J Mol Sci"}],"container-title":["The Plant Cell"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/plcell\/article-pdf\/37\/5\/koaf058\/63135133\/koaf058.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/plcell\/article-pdf\/37\/5\/koaf058\/63135133\/koaf058.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,5,30]],"date-time":"2025-05-30T09:27:29Z","timestamp":1748597249000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/plcell\/article\/doi\/10.1093\/plcell\/koaf058\/8127778"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,5]]},"references-count":97,"journal-issue":{"issue":"5","published-print":{"date-parts":[[2025,5,9]]}},"URL":"https:\/\/doi.org\/10.1093\/plcell\/koaf058","relation":{},"ISSN":["1040-4651","1532-298X"],"issn-type":[{"value":"1040-4651","type":"print"},{"value":"1532-298X","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2025,5]]},"published":{"date-parts":[[2025,5]]},"article-number":"koaf058"}}