{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T08:55:46Z","timestamp":1791276946090,"version":"4.1.0"},"reference-count":113,"publisher":"Oxford University Press (OUP)","issue":"9","license":[{"start":{"date-parts":[[2026,9,7]],"date-time":"2026-09-07T00:00:00Z","timestamp":1788739200000},"content-version":"vor","delay-in-days":6,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R35GM131760"],"award-info":[{"award-number":["R35GM131760"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["1R35GM158110"],"award-info":[{"award-number":["1R35GM158110"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["#2310356"],"award-info":[{"award-number":["#2310356"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["#2531822"],"award-info":[{"award-number":["#2531822"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Transcription factors (TFs) integrate a diverse array of inputs to achieve the exquisite control of gene expression necessary for life. In plants, this is exemplified by the deeply conserved CLASS III HOMEODOMAIN LEUCINE ZIPPER (HD-Zip III) family of TFs. HD-Zip III activity is controlled by inputs at transcriptional, post-transcriptional, and post-translational levels. As part of their multidomain architecture, HD-Zip III TFs contain a StAR-related lipid transfer (START) domain, a ubiquitously distributed evolutionary module that binds various types of lipophilic ligands. Here, we show that HD-Zip III and HD-Zip IV proteins contain an additional cryptic, deeply conserved START domain which we term the disorder-containing START domain (dSTART). The dSTART domain is required for HD-Zip III developmental function, promoting homodimerization and controlling subcellular localization and DNA-binding competence. The dSTART domain also helps discriminate responsive from nonresponsive binding sites across the HD-Zip III shared genetic network. Finally, we identify candidate ligands of the dSTART domain including several species of phosphatidylglycerol. The identification and functional characterization of a cryptic START domain provides new mechanistic insights into a deeply conserved family of TFs with roles in nearly all aspects of plant development.<\/jats:p>","DOI":"10.1093\/plcell\/koag267","type":"journal-article","created":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T11:31:11Z","timestamp":1788521471000},"source":"Crossref","is-referenced-by-count":0,"title":["A cryptic START domain regulates deeply conserved transcription factors"],"prefix":"10.1093","volume":"38","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-8815-7226","authenticated-orcid":false,"given":"Courtney E","family":"Dresden","sequence":"first","affiliation":[{"name":"Department of Biology, University of Pennsylvania , 415 S. University Ave, Philadelphia, PA 19104 ,","place":["United States"]},{"name":"Molecular, Cellular, and Developmental Biology, The Ohio State University , 111 Biological Sciences Building, 484 W. 12th Avenue, Columbus, OH 43215 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3719-0293","authenticated-orcid":false,"given":"Ekaterina P","family":"Andrianova","sequence":"additional","affiliation":[{"name":"Department of Microbiology, The Ohio State University , 105 Biological Sciences Building, 484 W. 12th Avenue, Columbus, OH 43215 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-5418-7008","authenticated-orcid":false,"given":"Brian J","family":"Smith","sequence":"additional","affiliation":[{"name":"Department of Molecular Genetics, 105 Biological Sciences Building, 484 W. 12th Avenue, The Ohio State University , Columbus, OH 43215 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-2605-7055","authenticated-orcid":false,"given":"Nicole I","family":"Callery","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Pennsylvania , 415 S. University Ave, Philadelphia, PA 19104 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5165-8262","authenticated-orcid":false,"given":"Dominic","family":"Kolonay","sequence":"additional","affiliation":[{"name":"Molecular, Cellular, and Developmental Biology, The Ohio State University , 111 Biological Sciences Building, 484 W. 12th Avenue, Columbus, OH 43215 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9443-8358","authenticated-orcid":false,"given":"Ashton S","family":"Holub","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Pennsylvania , 415 S. University Ave, Philadelphia, PA 19104 ,","place":["United States"]},{"name":"Department of Molecular Genetics, 105 Biological Sciences Building, 484 W. 12th Avenue, The Ohio State University , Columbus, OH 43215 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5526-3938","authenticated-orcid":false,"given":"Ricardo A","family":"Urquidi Camacho","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Pennsylvania , 415 S. University Ave, Philadelphia, PA 19104 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2564-434X","authenticated-orcid":false,"given":"Sarah G","family":"Choudury","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Pennsylvania , 415 S. University Ave, Philadelphia, PA 19104 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-5770-8739","authenticated-orcid":false,"given":"Isabella J","family":"Higgins","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Pennsylvania , 415 S. University Ave, Philadelphia, PA 19104 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6708-5323","authenticated-orcid":false,"given":"Igor B","family":"Zhulin","sequence":"additional","affiliation":[{"name":"Department of Microbiology, The Ohio State University , 105 Biological Sciences Building, 484 W. 12th Avenue, Columbus, OH 43215 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4580-1779","authenticated-orcid":false,"given":"Aman Y","family":"Husbands","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Pennsylvania , 415 S. University Ave, Philadelphia, PA 19104 ,","place":["United States"]},{"name":"Molecular, Cellular, and Developmental Biology, The Ohio State University , 111 Biological Sciences Building, 484 W. 12th Avenue, Columbus, OH 43215 ,","place":["United States"]},{"name":"Department of Molecular Genetics, 105 Biological Sciences Building, 484 W. 12th Avenue, The Ohio State University , Columbus, OH 43215 ,","place":["United States"]},{"name":"Epigenetics Institute, University of Pennsylvania , 3400 Civic Center Boulevard, Philadelphia, PA 19104 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,9,7]]},"reference":[{"key":"2026100604463824800_koag267-B1","doi-asserted-by":"publisher","first-page":"2791","DOI":"10.1242\/jcs.02485","article-title":"Give lipids a START: the StAR-related lipid transfer (START) domain in mammals","volume":"118","author":"Alpy","year":"2005","journal-title":"J Cell Sci"},{"key":"2026100604463824800_koag267-B2","doi-asserted-by":"publisher","first-page":"17945","DOI":"10.1074\/jbc.M500723200","article-title":"Functional characterization of the MENTAL domain","volume":"280","author":"Alpy","year":"2005","journal-title":"J Biol Chem"},{"key":"2026100604463824800_koag267-B3","author":"Apprill","year":"2026"},{"key":"2026100604463824800_koag267-B4","doi-asserted-by":"publisher","first-page":"10223","DOI":"10.1074\/jbc.M611221200","article-title":"Cholesterol binding does not predict activity of the steroidogenic acute regulatory protein, StAR","volume":"282","author":"Baker","year":"2007","journal-title":"J Biol Chem"},{"key":"2026100604463824800_koag267-B5","doi-asserted-by":"publisher","first-page":"281","DOI":"10.1016\/S0092-8674(04)00045-5","article-title":"MicroRNAs: genomics, biogenesis, mechanism, and function","volume":"116","author":"Bartel","year":"2004","journal-title":"Cell"},{"key":"2026100604463824800_koag267-B6","doi-asserted-by":"publisher","first-page":"W335","DOI":"10.1093\/nar\/gkl217","article-title":"The MPI bioinformatics toolkit for protein sequence analysis","volume":"34","author":"Biegert","year":"2006","journal-title":"Nuc Acids Res"},{"key":"2026100604463824800_koag267-B7","doi-asserted-by":"publisher","first-page":"911","DOI":"10.1139\/y59-099","article-title":"A rapid method of total lipid extraction and purification","volume":"37","author":"Bligh","year":"1959","journal-title":"Can J Biochem Physiol"},{"key":"2026100604463824800_koag267-B8","doi-asserted-by":"publisher","first-page":"329","DOI":"10.1042\/bj3170329","article-title":"Multiple steps in the regulation of transcription-factor level and activity","volume":"317","author":"Calkhoven","year":"1996","journal-title":"Biochem J"},{"key":"2026100604463824800_koag267-B9","doi-asserted-by":"publisher","DOI":"10.1016\/j.molmet.2020.101015","article-title":"The type 2 diabetes gene product STARD10 is a phosphoinositide-binding protein that controls insulin secretory granule biogenesis","volume":"40","author":"Carrat","year":"2020","journal-title":"Mol Metab"},{"key":"2026100604463824800_koag267-B10","doi-asserted-by":"publisher","first-page":"711","DOI":"10.1016\/j.jtbi.2010.03.046","article-title":"The rules of variation: amino acid exchange according to the rotating circular genetic code","volume":"264","author":"Castro-Chavez","year":"2010","journal-title":"J Theor Biol"},{"key":"2026100604463824800_koag267-B11","doi-asserted-by":"publisher","first-page":"1653","DOI":"10.1242\/dev.001016","article-title":"The AP2 transcription factors of DORNR\u04e6SCHEN and DORNR\u04e6SCHEN-LIKE redundantly control Arabidopsis embryo patterning via interaction with PHAVOLUTA","volume":"134","author":"Chandler","year":"2007","journal-title":"Development"},{"key":"2026100604463824800_koag267-B12","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1093\/mp\/sst026","article-title":"Protein-protein interactions in the regulation of WRKY transcription factors","volume":"6","author":"Chi","year":"2013","journal-title":"Mol Plant"},{"key":"2026100604463824800_koag267-B13","doi-asserted-by":"publisher","DOI":"10.1186\/1752-0509-6-90","article-title":"Multiple independent analyses reveal only transcription factors as an enriched functional class associated with microRNAs","volume":"6","author":"Croft","year":"2012","journal-title":"BMC Syst Biol"},{"key":"2026100604463824800_koag267-B14","doi-asserted-by":"publisher","first-page":"109","DOI":"10.1016\/S0009-3084(01)00171-2","article-title":"The binding of phosphatidylcholine to the phosphatidylcholine transfer protein: affinity and role in folding","volume":"112","author":"de Brouwer","year":"2001","journal-title":"Chem Phys Lipids"},{"key":"2026100604463824800_koag267-B15","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"},{"key":"2026100604463824800_koag267-B16","doi-asserted-by":"publisher","DOI":"10.1016\/j.pbi.2021.102148","article-title":"Diverse regulatory mechanisms of StARkin domains in land plants and mammals","volume":"64","author":"Dresden","year":"2021","journal-title":"Current Opin Plant Biol"},{"key":"2026100604463824800_koag267-B17","doi-asserted-by":"publisher","first-page":"4405","DOI":"10.1158\/0008-5472.CAN-12-0777","article-title":"Functional interaction of tumor suppressor DLC1 and caveolin-1 in cancer cells","volume":"72","author":"Du","year":"2012","journal-title":"Cancer Res"},{"key":"2026100604463824800_koag267-B18","doi-asserted-by":"publisher","first-page":"D427","DOI":"10.1093\/nar\/gky995","article-title":"The pfam protein families database in 2019","volume":"47","author":"El-Gebali","year":"2019","journal-title":"Nucleic Acids Res."},{"key":"2026100604463824800_koag267-B19","doi-asserted-by":"publisher","first-page":"76","DOI":"10.1016\/j.tips.2013.11.005","article-title":"Regulation of transcription factor activity by interconnected, post-translational modifications","volume":"35","author":"Filtz","year":"2014","journal-title":"Trends Pharmacol Sci"},{"key":"2026100604463824800_koag267-B20","doi-asserted-by":"publisher","first-page":"187","DOI":"10.1007\/978-1-61779-585-5_8","article-title":"Evolution of protein domain architectures","volume":"856","author":"Forslund","year":"2012","journal-title":"Methods in Molecular Biology (Clifton, N.J.)"},{"key":"2026100604463824800_koag267-B21","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-30770-1","article-title":"ChIP-hub provides an integrative platform for exploring plant regulome","volume":"13","author":"Fu","year":"2022","journal-title":"Nat Commun."},{"key":"2026100604463824800_koag267-B22","doi-asserted-by":"publisher","first-page":"681","DOI":"10.1083\/jcb.200803129","article-title":"Cardiolipin provides an essential activating platform for caspase-8 on mitochondria","volume":"183","author":"Gonzalvez","year":"2008","journal-title":"J Cell Biol."},{"key":"2026100604463824800_koag267-B23","doi-asserted-by":"publisher","first-page":"W72","DOI":"10.1093\/nar\/gkaa243","article-title":"TREND: a platform for exploring protein function in prokaryotes based on phylogenetic, domain architecture and gene neighborhood analyses","volume":"48","author":"Gumerov","year":"2020","journal-title":"Nucleic Acids Res."},{"key":"2026100604463824800_koag267-B24","doi-asserted-by":"publisher","first-page":"2022","DOI":"10.1111\/nph.70309","article-title":"Mechanisms driving functional divergence of transcription factor paralogs","volume":"247","author":"Higgins","year":"2025","journal-title":"New Phytol."},{"key":"2026100604463824800_koag267-B25","doi-asserted-by":"publisher","first-page":"518","DOI":"10.1093\/molbev\/msx281","article-title":"UFBoot2: improving the ultrafast bootstrap approximation","volume":"35","author":"Hoang","year":"2018","journal-title":"Mol Biol Evol."},{"key":"2026100604463824800_koag267-B26","doi-asserted-by":"publisher","first-page":"1785","DOI":"10.1126\/science.1151651","article-title":"Gene regulation by transcription factors and microRNAs","volume":"319","author":"Hobert","year":"2008","journal-title":"Science (New York, N.Y.)"},{"key":"2026100604463824800_koag267-B27","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-54269-z","article-title":"START domains generate paralog-specific regulons from a single network architecture","volume":"15","author":"Holub","year":"2024","journal-title":"Nat Commun."},{"key":"2026100604463824800_koag267-B28","doi-asserted-by":"publisher","DOI":"10.1038\/srep42853","article-title":"Dual mechanisms regulate nucleocytoplasmic localization of human DDX6","volume":"7","author":"Huang","year":"2017","journal-title":"Sci Reports"},{"key":"2026100604463824800_koag267-B29","doi-asserted-by":"publisher","first-page":"1783","DOI":"10.1105\/tpc.16.00289","article-title":"In planta single-molecule pull-down reveals tetrameric stoichiometry of HD-ZIPIII:LITTLE ZIPPER complexes","volume":"28","author":"Husbands","year":"2016","journal-title":"Plant Cell."},{"key":"2026100604463824800_koag267-B30","doi-asserted-by":"publisher","first-page":"2332","DOI":"10.1093\/plcell\/koad058","article-title":"The START domain potentiates HD-ZIPIII transcriptional activity","volume":"35","author":"Husbands","year":"2023","journal-title":"Plant Cell"},{"key":"2026100604463824800_koag267-B31","doi-asserted-by":"publisher","DOI":"10.1242\/dev.169300","article-title":"ATML1 activity is restricted to the outermost cells of the embryo through post-transcriptional repressions","volume":"146","author":"Iida","year":"2019","journal-title":"Development (Cambridge, England)"},{"key":"2026100604463824800_koag267-B32","doi-asserted-by":"publisher","first-page":"134","DOI":"10.1002\/1097-0134(20010501)43:2&lt;134::AID-PROT1025&gt;3.0.CO;2-I","article-title":"Adaptations of the helix-grip fold for ligand binding and catalysis in the START domain superfamily","volume":"43","author":"Iyer","year":"2001","journal-title":"Proteins: Structure, Function, and Genetics"},{"key":"2026100604463824800_koag267-B33","doi-asserted-by":"publisher","first-page":"30728","DOI":"10.1074\/jbc.M703745200","article-title":"Interacting proteins dictate function of the minimal START domain phosphatidylcholine transfer protein\/StarD2","volume":"282","author":"Kanno","year":"2007","journal-title":"J Biol Chem."},{"key":"2026100604463824800_koag267-B34","doi-asserted-by":"publisher","first-page":"996","DOI":"10.1016\/0955-0674(90)90148-8","article-title":"Transcriptional control and the integration of cell-autonomous and environmental cues during development","volume":"2","author":"Karin","year":"1990","journal-title":"Curr Opin Cell Biol."},{"key":"2026100604463824800_koag267-B35","doi-asserted-by":"publisher","first-page":"3059","DOI":"10.1093\/nar\/gkf436","article-title":"MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform","volume":"30","author":"Katoh","year":"2002","journal-title":"Nucleic Acids Res."},{"key":"2026100604463824800_koag267-B36","doi-asserted-by":"publisher","first-page":"84","DOI":"10.1111\/j.1365-313X.2005.02354.x","article-title":"MicroRNA-directed cleavage of ATHB15 mRNA regulates vascular development in Arabidopsis inflorescence stems","volume":"42","author":"Kim","year":"2005","journal-title":"Plant J."},{"key":"2026100604463824800_koag267-B37","doi-asserted-by":"publisher","first-page":"920","DOI":"10.1105\/tpc.107.057448","article-title":"HD-ZIPIII activity is modulated by competitive inhibitors via a feedback loop in Arabidopsis shoot apical meristem development","volume":"20","author":"Kim","year":"2008","journal-title":"Plant Cell."},{"key":"2026100604463824800_koag267-B38","doi-asserted-by":"publisher","DOI":"10.1016\/j.cdev.2024.203936","article-title":"Self-organization underlies developmental robustness in plants","volume":"184","author":"Kong","year":"2025","journal-title":"Cells Dev."},{"key":"2026100604463824800_koag267-B39","doi-asserted-by":"publisher","DOI":"10.1128\/JB.00687-19","article-title":"The intrinsically disordered region of ExbD is required for signal transduction","volume":"202","author":"Kopp","year":"2020","journal-title":"J Bacteriol."},{"key":"2026100604463824800_koag267-B40","doi-asserted-by":"publisher","first-page":"M46","DOI":"10.1016\/S0962-8924(99)01679-7","article-title":"Eukaryotic transcriptional control","volume":"9","author":"Kornberg","year":"1999","journal-title":"Trends Cell Biol."},{"key":"2026100604463824800_koag267-B41","doi-asserted-by":"publisher","first-page":"3856","DOI":"10.1093\/nar\/gkab185","article-title":"Androgen and glucocorticoid receptor direct distinct transcriptional programs by receptor-specific and shared DNA binding sites","volume":"49","author":"Kulik","year":"2021","journal-title":"Nucleic Acids Res."},{"key":"2026100604463824800_koag267-B42","doi-asserted-by":"publisher","first-page":"1082","DOI":"10.1105\/tpc.114.135160","article-title":"VIH2 regulates the synthesis of inositol pyrophosphate InsP8 and jasmonate-dependent defenses in Arabidopsis","volume":"27","author":"Laha","year":"2015","journal-title":"Plant Cell."},{"key":"2026100604463824800_koag267-B43","doi-asserted-by":"publisher","first-page":"1305","DOI":"10.1016\/S1357-2725(97)00085-X","article-title":"Transcription factors: an overview","volume":"29","author":"Latchman","year":"1997","journal-title":"Int J Biochem Cell Biol."},{"key":"2026100604463824800_koag267-B44","doi-asserted-by":"publisher","first-page":"W78","DOI":"10.1093\/nar\/gkae268","article-title":"Interactive tree of life (iTOL) v6: recent updates to the phylogenetic tree display and annotation tool","volume":"52","author":"Letunic","year":"2024","journal-title":"Nucleic Acids Res."},{"key":"2026100604463824800_koag267-B45","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1007\/s004250050110","article-title":"Intracellular distribution and identification of the nuclear localization signals of two plant heat-stress transcription factors","volume":"202","author":"Lyck","year":"1997","journal-title":"Planta"},{"key":"2026100604463824800_koag267-B46","doi-asserted-by":"publisher","first-page":"1064","DOI":"10.1126\/science.1172408","article-title":"Regulators of PP2C phosphatase activity function as abscisic acid sensors","volume":"324","author":"Ma","year":"2009","journal-title":"Science (New York, N.Y.)"},{"key":"2026100604463824800_koag267-B47","doi-asserted-by":"publisher","first-page":"281","DOI":"10.1093\/nar\/30.1.281","article-title":"CDD: a database of conserved domain alignments with links to domain three-dimensional structure","volume":"30","author":"Marchler-Bauer","year":"2002","journal-title":"Nucleic Acids Res."},{"key":"2026100604463824800_koag267-B48","doi-asserted-by":"publisher","first-page":"1754","DOI":"10.1093\/emboj\/21.7.1754","article-title":"Regulated nuclear import of the STAT1 transcription factor by direct binding of importin-alpha","volume":"21","author":"McBride","year":"2002","journal-title":"EMBO J."},{"key":"2026100604463824800_koag267-B49","doi-asserted-by":"publisher","first-page":"2935","DOI":"10.1242\/dev.125.15.2935","article-title":"Leaf polarity and meristem formation in Arabidopsis","volume":"125","author":"McConnel","year":"1998","journal-title":"Development (Cambridge, England)"},{"key":"2026100604463824800_koag267-B50","doi-asserted-by":"publisher","first-page":"709","DOI":"10.1038\/35079635","article-title":"Role of PHABULOSA and PHAVOLUTA in determining radial patterning in shoots","volume":"411","author":"McConnell","year":"2001","journal-title":"Nature"},{"key":"2026100604463824800_koag267-B51","doi-asserted-by":"publisher","first-page":"1357","DOI":"10.1105\/tpc.16.00640","article-title":"Type-B ARABIDOPSIS RESPONSE REGULATORs specify the shoot stem cell niche by dual regulation of WUSCHEL","volume":"29","author":"Meng","year":"2017","journal-title":"Plant Cell."},{"key":"2026100604463824800_koag267-B52","doi-asserted-by":"publisher","first-page":"11973","DOI":"10.1073\/pnas.1516110113","article-title":"Regulation of MIR165\/166 by class II and class III homeodomain leucine zipper proteins establishes leaf polarity","volume":"113","author":"Merelo","year":"2016","journal-title":"Proc Natl Acad Sci U S A."},{"key":"2026100604463824800_koag267-B53","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1016\/j.tplants.2012.11.002","article-title":"Structure and function of abscisic acid receptors","volume":"18","author":"Miyakawa","year":"2013","journal-title":"Trends Plant Sci."},{"key":"2026100604463824800_koag267-B54","doi-asserted-by":"publisher","first-page":"E2274","DOI":"10.1073\/pnas.1719588115","article-title":"The timescale of early land plant evolution","volume":"115","author":"Morris","year":"2018","journal-title":"Proc Natl Acad Sci U S A."},{"key":"2026100604463824800_koag267-B55","doi-asserted-by":"publisher","first-page":"1142","DOI":"10.1104\/pp.105.073833","article-title":"MEKHLA, a novel domain with similarity to PAS domains, is fused to plant homeodomain-leucine zipper III proteins","volume":"140","author":"Mukherjee","year":"2006","journal-title":"Plant Physiol."},{"key":"2026100604463824800_koag267-B56","doi-asserted-by":"publisher","first-page":"2315","DOI":"10.1093\/plphys\/kiac383","article-title":"The START domain mediates Arabidopsis GLABRA2 dimerization and turnover independently of homeodomain DNA binding","volume":"190","author":"Mukherjee","year":"2022","journal-title":"Plant Physiol."},{"key":"2026100604463824800_koag267-B57","doi-asserted-by":"publisher","first-page":"448","DOI":"10.1111\/dgd.12753","article-title":"The lipid-binding START domain regulates the dimerization of ATML1 via modulating the ZIP motif activity in Arabidopsis thaliana","volume":"63","author":"Nagata","year":"2021","journal-title":"Dev Growth Differ."},{"key":"2026100604463824800_koag267-B58","doi-asserted-by":"publisher","DOI":"10.1242\/dev.194969","article-title":"Ceramides mediate positional signals in Arabidopsis thaliana protoderm differentiation","volume":"148","author":"Nagata","year":"2021","journal-title":"Development (Cambridge, England)"},{"key":"2026100604463824800_koag267-B59","doi-asserted-by":"publisher","first-page":"4766","DOI":"10.1128\/MCB.00647-13","article-title":"Androgen induces a switch from cytoplasmic retention to nuclear import of the androgen receptor","volume":"33","author":"Ni","year":"2013","journal-title":"Mol Cell Biol."},{"key":"2026100604463824800_koag267-B60","doi-asserted-by":"publisher","first-page":"1068","DOI":"10.1126\/science.1173041","article-title":"Abscisic acid inhibits type 2C protein phosphatases via the PYR\/PYL family of START proteins","volume":"324","author":"Park","year":"2009","journal-title":"Science (New York, N.Y.)"},{"key":"2026100604463824800_koag267-B61","doi-asserted-by":"publisher","first-page":"517","DOI":"10.1038\/nsmb.1598","article-title":"Active nuclear import and cytoplasmic retention of activation-induced deaminase","volume":"16","author":"Patenaude","year":"2009","journal-title":"Nat Struct Mol Biol."},{"key":"2026100604463824800_koag267-B62","doi-asserted-by":"publisher","first-page":"3333","DOI":"10.1002\/j.1460-2075.1987.tb02654.x","article-title":"Two signals mediate hormone-dependent nuclear localization of the glucocorticoid receptor","volume":"6","author":"Picard","year":"1987","journal-title":"EMBO J."},{"key":"2026100604463824800_koag267-B63","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1146\/annurev-cellbio-100617-062459","article-title":"Regulation of division and differentiation of plant stem cells","volume":"34","author":"Pierre-Jerome","year":"2018","journal-title":"Annu Rev Cell Dev Biol."},{"key":"2026100604463824800_koag267-B64","doi-asserted-by":"publisher","first-page":"130","DOI":"10.1016\/S0968-0004(99)01362-6","article-title":"START: a lipid-binding domain in StAR, HD-ZIP and signalling proteins","volume":"24","author":"Ponting","year":"1999","journal-title":"Trends Biochem Sci."},{"key":"2026100604463824800_koag267-B65","doi-asserted-by":"publisher","first-page":"14217","DOI":"10.1074\/jbc.M117.780007","article-title":"Interaction between the PH and START domains of ceramide transfer protein competes with phosphatidylinositol 4-phosphate binding by the PH domain","volume":"292","author":"Prashek","year":"2017","journal-title":"J Biol Chem."},{"key":"2026100604463824800_koag267-B66","doi-asserted-by":"publisher","first-page":"36441","DOI":"10.1074\/jbc.M004751200","article-title":"IkappaBalpha and IkappaBalpha \/NF-kappa B complexes are retained in the cytoplasm through interaction with a novel partner, RasGAP SH3-binding protein 2","volume":"275","author":"Prigent","year":"2000","journal-title":"J Biol Chem."},{"key":"2026100604463824800_koag267-B67","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1105\/tpc.104.026161","article-title":"Class III homeodomain-leucine zipper gene family members have overlapping, antagonistic, and distinct roles in Arabidopsis development","volume":"17","author":"Prigge","year":"2005","journal-title":"Plant Cell."},{"key":"2026100604463824800_koag267-B68","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2148-8-286","article-title":"The Bet v 1 fold: an ancient, versatile scaffold for binding of large, hydrophobic ligands","volume":"8","author":"Radauer","year":"2008","journal-title":"BMC Evol Biol."},{"key":"2026100604463824800_koag267-B69","doi-asserted-by":"publisher","first-page":"513","DOI":"10.1016\/S0092-8674(02)00863-2","article-title":"Prediction of plant microRNA targets","volume":"110","author":"Rhoades","year":"2002","journal-title":"Cell"},{"key":"2026100604463824800_koag267-B70","doi-asserted-by":"publisher","first-page":"507","DOI":"10.1038\/nsb812","article-title":"Structure of human phosphatidylcholine transfer protein in complex with its ligand","volume":"9","author":"Roderick","year":"2002","journal-title":"Nat Struct Biol."},{"key":"2026100604463824800_koag267-B71","doi-asserted-by":"publisher","first-page":"408","DOI":"10.1111\/nph.15133","article-title":"Evolutionary history of HOMEODOMAIN LECUINE ZIPPER transcription factors during plant transition to land","volume":"219","author":"Romani","year":"2018","journal-title":"New Phytol."},{"key":"2026100604463824800_koag267-B72","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1111\/nph.14132","article-title":"Regulatory function of homeodomain-leucine zipper (HD-ZIP) family proteins during embryogenesis","volume":"213","author":"Roodbarkelari","year":"2017","journal-title":"New Phytol."},{"key":"2026100604463824800_koag267-B73","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1042\/BSR20080111","article-title":"The mechanism of specific binding of free cholesterol by the steroidogenic acute regulatory protein: evidence for a role of the C-terminal \u03b1-helix in the gating of the binding site","volume":"29","author":"Roostaee","year":"2009","journal-title":"Biosci Rep."},{"key":"2026100604463824800_koag267-B74","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1210\/me.2007-0508","article-title":"Structure of SF-1 bound by different phospholipids: evidence for regulatory ligands","volume":"23","author":"Sablin","year":"2009","journal-title":"Molecular Endocrinology (Baltimore, Md.)"},{"key":"2026100604463824800_koag267-B75","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1111\/j.1440-169X.2011.01318.x","article-title":"Mir165\/166 and the development of land plants","volume":"54","author":"Sakaguchi","year":"2012","journal-title":"Dev Growth Differ."},{"key":"2026100604463824800_koag267-B76","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2017.00565","article-title":"MicroRNA and transcription factor: key players in plant regulatory network","volume":"8","author":"Samad","year":"2017","journal-title":"Front Plant Sci."},{"key":"2026100604463824800_koag267-B77","doi-asserted-by":"publisher","first-page":"2240","DOI":"10.1128\/MCB.18.4.2240","article-title":"The tomato Hsf system: HsfA2 needs interaction with HsfA1 for efficient nuclear import and may be localized in cytoplasmic heat stress granules","volume":"18","author":"Scharf","year":"1998","journal-title":"Mol Cell Biol."},{"key":"2026100604463824800_koag267-B78","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/nmeth.2019","article-title":"Fiji: an open-source platform for biological-image analysis","volume":"9","author":"Schindelin","year":"2012","journal-title":"Nat Methods."},{"key":"2026100604463824800_koag267-B79","doi-asserted-by":"publisher","DOI":"10.1186\/gb-2004-5-6-r41","article-title":"START lipid\/sterol-binding domains are amplified in plants and are predominantly associated with homeodomain transcription factors","volume":"5","author":"Schrick","year":"2004","journal-title":"Genome Biol."},{"key":"2026100604463824800_koag267-B80","doi-asserted-by":"publisher","DOI":"10.1016\/j.pbi.2023.102417","article-title":"HD-Zip IV transcription factors: drivers of epidermal cell fate integrate metabolic signals","volume":"75","author":"Schrick","year":"2023","journal-title":"Curr Opin Plant Biol."},{"key":"2026100604463824800_koag267-B81","doi-asserted-by":"publisher","first-page":"706","DOI":"10.1038\/s41586-019-1923-7","article-title":"Improved protein structure prediction using potentials from deep learning","volume":"577","author":"Senior","year":"2020","journal-title":"Nature"},{"key":"2026100604463824800_koag267-B82","doi-asserted-by":"publisher","DOI":"10.3390\/ijms19124047","article-title":"Multiple links between HD-Zip proteins and hormone networks","volume":"19","author":"Sessa","year":"2018","journal-title":"Int J Mol Sci."},{"key":"2026100604463824800_koag267-B83","doi-asserted-by":"publisher","first-page":"3408","DOI":"10.1038\/s41598-017-03632-w","article-title":"Phytohormonal crosstalk modulates the expression of miR166\/165s, target Class III HD-ZIPs, and KANADI genes during root growth in Arabidopsis thaliana","volume":"7","author":"Singh","year":"2017","journal-title":"Sci Rep."},{"key":"2026100604463824800_koag267-B84","doi-asserted-by":"publisher","first-page":"126","DOI":"10.1016\/j.pbi.2016.07.004","article-title":"Transcriptional regulation by complex interplay between post-translational modifications","volume":"33","author":"Skelly","year":"2016","journal-title":"Curr Opin Plant Biol."},{"key":"2026100604463824800_koag267-B85","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1016\/j.devcel.2017.10.003","article-title":"Boundary formation through a direct threshold-based readout of mobile small RNA gradients","volume":"43","author":"Skopelitis","year":"2017","journal-title":"Dev Cell."},{"key":"2026100604463824800_koag267-B86","doi-asserted-by":"publisher","first-page":"W244","DOI":"10.1093\/nar\/gki408","article-title":"The HHpred interactive server for protein homology detection and structure prediction","volume":"33","author":"Soding","year":"2005","journal-title":"Nucleic Acids Res."},{"key":"2026100604463824800_koag267-B87","author":"Stark"},{"key":"2026100604463824800_koag267-B88","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 (Oxford, England)"},{"key":"2026100604463824800_koag267-B89","doi-asserted-by":"publisher","first-page":"1137","DOI":"10.1126\/science.abh4054","article-title":"Florigen sequestration in cellular membranes modulates temperature-responsive flowering","volume":"373","author":"Susila","year":"2021","journal-title":"Science (New York, N.Y.)"},{"key":"2026100604463824800_koag267-B90","doi-asserted-by":"publisher","first-page":"e19521","DOI":"10.1371\/journal.pone.0019521","article-title":"Comparative structural analysis of lipid binding START domains","volume":"6","author":"Thorsell","year":"2011","journal-title":"PLoS One"},{"key":"2026100604463824800_koag267-B91","doi-asserted-by":"publisher","first-page":"408","DOI":"10.1038\/75192","article-title":"Structure and lipid transport mechanism of a StAR-related domain","volume":"7","author":"Tsujishita","year":"2000","journal-title":"Nat Struct Biol."},{"key":"2026100604463824800_koag267-B92","doi-asserted-by":"publisher","first-page":"3944","DOI":"10.1111\/febs.15474","article-title":"Molecular basis for the recognition of steroidogenic acute regulatory protein by the 14-3-3 protein family","volume":"287","author":"Tugaeva","year":"2020","journal-title":"FEBS J."},{"key":"2026100604463824800_koag267-B93","author":"Vijaya","year":"2024"},{"key":"2026100604463824800_koag267-B94","doi-asserted-by":"publisher","first-page":"110","DOI":"10.1002\/j.1460-2075.1996.tb00339.x","article-title":"Differential ligand-dependent interactions between the AF-2 activing domain of nuclear receptors and the putative transcriptional intermediary factors mSUG1 and TIF1","volume":"15","author":"vom Baur","year":"1996","journal-title":"EMBO J."},{"key":"2026100604463824800_koag267-B95","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1038\/s41477-019-0560-3","article-title":"Genomes of early-diverging streptophyte algae shed light on plant terrestrialization","volume":"6","author":"Wang","year":"2020","journal-title":"Nat Plants."},{"key":"2026100604463824800_koag267-B96","doi-asserted-by":"publisher","first-page":"1189","DOI":"10.1093\/bioinformatics\/btp033","article-title":"Jalview Version2\u2014a multiple sequence alignment editor and analysis workbench","volume":"25","author":"Waterhouse","year":"2009","journal-title":"Bioinformatics (Oxford, England)"},{"key":"2026100604463824800_koag267-B97","doi-asserted-by":"publisher","first-page":"3379","DOI":"10.1105\/tpc.107.055772","article-title":"A feedback regulatory module formed by LITTLE ZIPPER and HD-ZIPIII genes","volume":"19","author":"Wenkel","year":"2007","journal-title":"Plant Cell."},{"key":"2026100604463824800_koag267-B98","doi-asserted-by":"publisher","first-page":"1498","DOI":"10.1111\/nph.19917","article-title":"Arabidopsis PROTODERMAL FACTOR2 binds lysophosphatidylcholines and transcriptionally regulates phospholipid metabolism","volume":"244","author":"Wojciechowska","year":"2024","journal-title":"New Phytol."},{"key":"2026100604463824800_koag267-B99","doi-asserted-by":"publisher","first-page":"517","DOI":"10.1042\/BST20150275","article-title":"Lipid transfer proteins do their thing anchored at membrane contact sites\u2026but what is their thing?","volume":"44","author":"Wong","year":"2016","journal-title":"Biochem Soc Trans."},{"key":"2026100604463824800_koag267-B100","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msag117","article-title":"IQ-TREE3: phylogenomic inference software using complex evolutionary models","volume":"43","author":"Wong","year":"2026","journal-title":"Mol Biol Evol."},{"key":"2026100604463824800_koag267-B101","doi-asserted-by":"publisher","first-page":"3677","DOI":"10.1091\/mbc.e12-01-0046","article-title":"Sequence and structural analyses of nuclear export signals in the NESdb database","volume":"23","author":"Xu","year":"2012","journal-title":"Mol Biol Cell."},{"key":"2026100604463824800_koag267-B102","doi-asserted-by":"publisher","first-page":"559","DOI":"10.1038\/35019649","article-title":"The nuclear import functions of Smad2 is masked by SARA and unmasked by TGFb-dependent phosphorylation","volume":"2","author":"Xu","year":"2000","journal-title":"Nat Cell Biol."},{"key":"2026100604463824800_koag267-B103","doi-asserted-by":"publisher","first-page":"W174","DOI":"10.1093\/nar\/gkv342","article-title":"I-TASSER server: new development for protein structure and function predictions","volume":"43","author":"Yang","year":"2015","journal-title":"Nucleic Acids Res."},{"key":"2026100604463824800_koag267-B104","doi-asserted-by":"publisher","first-page":"2131","DOI":"10.1101\/gad.12.14.2131","article-title":"Control of cyclin B1 localization through regulated binding of the nuclear export factor CRM1","volume":"12","author":"Yang","year":"1998","journal-title":"Genes Dev."},{"key":"2026100604463824800_koag267-B105","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1038\/nmeth.3213","article-title":"The I-TASSER suite: protein structure and function prediction","volume":"12","author":"Yang","year":"2015","journal-title":"Nat Methods."},{"key":"2026100604463824800_koag267-B106","doi-asserted-by":"publisher","first-page":"5030","DOI":"10.1105\/tpc.113.120162","article-title":"Phosphatidic acid interacts with a MYB transcription factor and regulates its nuclear localization and function in Arabidopsis","volume":"25","author":"Yao","year":"2013","journal-title":"Plant Cell."},{"key":"2026100604463824800_koag267-B107","doi-asserted-by":"publisher","first-page":"307","DOI":"10.1016\/j.jgg.2017.03.006","article-title":"LRH-1 senses signaling from phosphatidylcholine to regulate the expansion growth of digestive organs via synergy with Wnt\/\u03b2-catenin signaling in zebrafish","volume":"44","author":"Zhai","year":"2017","journal-title":"J Genet Genomics."},{"key":"2026100604463824800_koag267-B108","doi-asserted-by":"publisher","first-page":"815","DOI":"10.1016\/j.tplants.2017.08.007","article-title":"Type-B ARABIDOPSIS RESPONSE REGULATORs directly activate WUSCHEL","volume":"22","author":"Zhang","year":"2017","journal-title":"Trends Plant Sci."},{"key":"2026100604463824800_koag267-B109","doi-asserted-by":"publisher","DOI":"10.1186\/gb-2008-9-9-r137","article-title":"Model-based analysis of ChIP-seq (MACS)","volume":"9","author":"Zhang","year":"2008","journal-title":"Genome Biol."},{"key":"2026100604463824800_koag267-B110","doi-asserted-by":"publisher","DOI":"10.1016\/j.crmeth.2021.100014","article-title":"Folding non-homology proteins by coupling deep-learning contact maps with I-TASSER assembly simulations","volume":"1","author":"Zheng","year":"2021","journal-title":"Cell Rep Methods."},{"key":"2026100604463824800_koag267-B111","doi-asserted-by":"publisher","first-page":"242","DOI":"10.1016\/j.cell.2011.03.024","article-title":"Arabidopsis Argonaute10 specifically sequesters miR166\/165 to regulate shoot apical meristem development","volume":"145","author":"Zhu","year":"2011","journal-title":"Cell"},{"key":"2026100604463824800_koag267-B112","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.79116","article-title":"Overcoming the cytoplasmic retention of GDOWN1 modulates global transcription and facilitates stress adaptation","volume":"11","author":"Zhu","year":"2022","journal-title":"eLife"},{"key":"2026100604463824800_koag267-B113","doi-asserted-by":"publisher","first-page":"2237","DOI":"10.1016\/j.jmb.2017.12.007","article-title":"A completely reimplemented MPI bioinformatics toolkit with a new HHpred server at its core","volume":"430","author":"Zimmermann","year":"2018","journal-title":"J Mol Biol."}],"container-title":["The Plant Cell"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/plcell\/advance-article-pdf\/doi\/10.1093\/plcell\/koag267\/70975972\/koag267.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/plcell\/article-pdf\/38\/9\/koag267\/70975972\/koag267.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/plcell\/article-pdf\/38\/9\/koag267\/70975972\/koag267.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T08:46:43Z","timestamp":1791276403000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/plcell\/article\/doi\/10.1093\/plcell\/koag267\/8786537"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,1]]},"references-count":113,"journal-issue":{"issue":"9","published-print":{"date-parts":[[2026,9,1]]}},"URL":"https:\/\/doi.org\/10.1093\/plcell\/koag267","relation":{},"ISSN":["1040-4651","1532-298X"],"issn-type":[{"value":"1040-4651","type":"print"},{"value":"1532-298X","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,9]]},"published":{"date-parts":[[2026,9,1]]},"article-number":"koag267"}}