{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T17:45:15Z","timestamp":1785433515045,"version":"3.56.0"},"posted":{"date-parts":[[2025,7,30]]},"group-title":"Plant Biology","reference-count":86,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2025,7,30]],"date-time":"2025-07-30T00:00:00Z","timestamp":1753833600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["T32GM007276"],"award-info":[{"award-number":["T32GM007276"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"id":[{"id":"https:\/\/ror.org\/006w34k90","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2025,10,21]]},"abstract":"<jats:title>Summary<\/jats:title>\n                <jats:p>\n                  Plants regulate stomatal development and function to acquire atmospheric carbon dioxide for photosynthesis while minimizing water loss. The ancestral basic helix-loop-helix transcription factor (TF) gene that drove stomata production in early land plants diversified to become paralogs\n                  <jats:italic>SPEECHLESS<\/jats:italic>\n                  (\n                  <jats:italic>SPCH<\/jats:italic>\n                  ),\n                  <jats:italic>MUTE<\/jats:italic>\n                  , and\n                  <jats:italic>FAMA<\/jats:italic>\n                  . Extant grasses exhibit a particularly interesting set of duplications and losses of\n                  <jats:italic>SPCH<\/jats:italic>\n                  .\n                <\/jats:p>\n                <jats:p>\n                  Using phylogenetic methods, we tracked the history of SPCH duplications.\n                  <jats:italic>Brachypodium distachyon<\/jats:italic>\n                  and\n                  <jats:italic>Oryza sativa<\/jats:italic>\n                  plants bearing mutations in either\n                  <jats:italic>SPCH1<\/jats:italic>\n                  or\n                  <jats:italic>SPCH2, and B. distachyon<\/jats:italic>\n                  plants with SPCH1 or SPCH2 translational reporters were assayed under different environmental conditions for their effects on stomatal development.\n                <\/jats:p>\n                <jats:p>\n                  We identified the Poaceae-specific\n                  <jats:italic>rho<\/jats:italic>\n                  whole genome duplication as the origin of\n                  <jats:italic>SPCH1<\/jats:italic>\n                  and\n                  <jats:italic>SPCH2<\/jats:italic>\n                  and demonstrated that both paralogs remain under selection. We found paralog-specific divergence in response to two environmental perturbations in both\n                  <jats:italic>B. distachyon and O. sativa<\/jats:italic>\n                  . Plausible molecular mechanisms underpinning paralog divergence, and cellular mechanisms driving the stomatal phenotypes are supported by analyses of\n                  <jats:italic>BdSPCH1<\/jats:italic>\n                  and\n                  <jats:italic>BdSPCH2<\/jats:italic>\n                  RNA and protein expression and by sequence variation among grasses.\n                <\/jats:p>\n                <jats:p>These studies suggest ways in which a duplication of a key stomatal regulator enables adaptation and could inform genetic strategies to mitigate anticipated stressors in agronomically important plants.<\/jats:p>","DOI":"10.1101\/2025.07.29.667563","type":"posted-content","created":{"date-parts":[[2025,7,30]],"date-time":"2025-07-30T23:40:14Z","timestamp":1753918814000},"source":"Crossref","is-referenced-by-count":0,"title":["SPEECHLESS duplication in grasses expands potential for environmental regulation of stomatal development"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3844-9386","authenticated-orcid":false,"given":"Joel M.","family":"Erberich","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Britney","family":"Bennett","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0873-3543","authenticated-orcid":false,"given":"Dominique C.","family":"Bergmann","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2025102318000755000_2025.07.29.667563v2.1","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1093\/nar\/gkq291","article-title":"TranslatorX: multiple alignment of nucleotide sequences guided by amino acid translations","volume":"38","year":"2010","journal-title":"Nucleic Acids Research"},{"key":"2025102318000755000_2025.07.29.667563v2.2","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msad141"},{"key":"2025102318000755000_2025.07.29.667563v2.3","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkv416"},{"key":"2025102318000755000_2025.07.29.667563v2.4","doi-asserted-by":"publisher","DOI":"10.1126\/science.1096014"},{"key":"2025102318000755000_2025.07.29.667563v2.5","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.arplant.58.032806.104023"},{"key":"2025102318000755000_2025.07.29.667563v2.6","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-8137.2009.03087.x"},{"key":"2025102318000755000_2025.07.29.667563v2.7","doi-asserted-by":"publisher","DOI":"10.1093\/plcell\/koac076"},{"key":"2025102318000755000_2025.07.29.667563v2.8","doi-asserted-by":"crossref","first-page":"371","DOI":"10.1111\/nph.15344","article-title":"Rice with reduced stomatal density conserves water and has improved drought tolerance under future climate conditions","volume":"221","year":"2019","journal-title":"The New Phytologist"},{"key":"2025102318000755000_2025.07.29.667563v2.9","doi-asserted-by":"crossref","first-page":"16179","DOI":"10.1038\/nplants.2016.179","article-title":"Origin and function of stomata in the moss Physcomitrella patens","volume":"2","year":"2016","journal-title":"Nature Plants"},{"key":"2025102318000755000_2025.07.29.667563v2.10","doi-asserted-by":"publisher","DOI":"10.1016\/j.tplants.2016.09.005"},{"key":"2025102318000755000_2025.07.29.667563v2.11","doi-asserted-by":"publisher","DOI":"10.1093\/sysbio\/syt072"},{"key":"2025102318000755000_2025.07.29.667563v2.12","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2022.04.040"},{"key":"2025102318000755000_2025.07.29.667563v2.13","doi-asserted-by":"publisher","DOI":"10.1104\/pp.15.00432"},{"key":"2025102318000755000_2025.07.29.667563v2.14","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1411766111"},{"key":"2025102318000755000_2025.07.29.667563v2.15","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msx069"},{"key":"2025102318000755000_2025.07.29.667563v2.16","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1300127110"},{"key":"2025102318000755000_2025.07.29.667563v2.17","doi-asserted-by":"crossref","unstructured":"Devi MJ , Reddy VR . 2018. 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