{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T02:45:11Z","timestamp":1791773111165,"version":"4.3.4"},"reference-count":29,"publisher":"Wiley","issue":"5","license":[{"start":{"date-parts":[[2026,10,11]],"date-time":"2026-10-11T00:00:00Z","timestamp":1791676800000},"content-version":"vor","delay-in-days":40,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100009708","name":"Novo Nordisk Fonden","doi-asserted-by":"publisher","award":["2019OC53580"],"award-info":[{"award-number":["2019OC53580"]}],"id":[{"id":"10.13039\/501100009708","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100009708","name":"Novo Nordisk Fonden","doi-asserted-by":"publisher","award":["NNF18OC0034226"],"award-info":[{"award-number":["NNF18OC0034226"]}],"id":[{"id":"10.13039\/501100009708","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100009708","name":"Novo Nordisk Fonden","doi-asserted-by":"publisher","award":["NNF20OC0061440"],"award-info":[{"award-number":["NNF20OC0061440"]}],"id":[{"id":"10.13039\/501100009708","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100009708","name":"Novo Nordisk Fonden","doi-asserted-by":"publisher","award":["NNF23OC0084195"],"award-info":[{"award-number":["NNF23OC0084195"]}],"id":[{"id":"10.13039\/501100009708","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004836","name":"Danmarks Frie Forskningsfond","doi-asserted-by":"publisher","award":["0136\u201000015B"],"award-info":[{"award-number":["0136\u201000015B"]}],"id":[{"id":"10.13039\/501100004836","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004836","name":"Danmarks Frie Forskningsfond","doi-asserted-by":"publisher","award":["0135\u201000014B"],"award-info":[{"award-number":["0135\u201000014B"]}],"id":[{"id":"10.13039\/501100004836","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100011075","name":"Wallenberg Wood Science Center","doi-asserted-by":"publisher","award":["WIFORCE"],"award-info":[{"award-number":["WIFORCE"]}],"id":[{"id":"10.13039\/501100011075","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Physiologia Plantarum"],"published-print":{"date-parts":[[2026,9]]},"abstract":"ABSTRACT<\/jats:title>\n \n Flowering in plants is controlled by internal and environmental signals, with temperature playing a critical role. Accordingly,\n \n Arabidopsis thaliana<\/jats:italic>\n <\/jats:styled-content>\n plants show temperature\u2010dependent variation in flowering time. Because unexpected cold periods can occur at any time, plants have evolved mechanisms to detect such cold snaps and delay flowering. Plants are more cold\u2010tolerant in the vegetative stage, whereas flowers are sensitive and often show reduced reproductive success due to damage to floral structures and gametes. At the molecular level, delayed flowering can be caused by repression of the\n FLOWERING LOCUS T<\/jats:italic>\n (\n FT<\/jats:italic>\n ) gene, and several MADS\u2010box transcription factors repress\n FT<\/jats:italic>\n expression in response to cold. In this article, we used a forward genetic screen to investigate how a cold snap affects the transition to flowering. We germinated an\n \n A. thaliana<\/jats:italic>\n <\/jats:styled-content>\n EMS M2 population at 20\u00b0C, gradually lowered the temperature to 10\u00b0C, and selected early\u2010flowering mutants. Whole\u2010genome sequencing identified seven mutant alleles of\n SHORT VEGETATIVE PHASE<\/jats:italic>\n (\n SVP<\/jats:italic>\n ), which we named\n frosty<\/jats:italic>\n . Yeast two\u2010hybrid assays showed that the corresponding FROSTY protein isoforms differ in their ability to interact with the MADS\u2010box proteins AGL6 and SOC1. These findings establish a central role for\n SVP<\/jats:italic>\n in repressing flowering in response to a cold snap and provide novel alleles, several of which affect splice junctions. Our study offers new insights into the molecular mechanisms that govern temperature\u2010responsive flowering in Arabidopsis and highlights the dynamic interplay between\n SVP<\/jats:italic>\n and environmental cues.\n <\/jats:p>","DOI":"10.1111\/ppl.71146","type":"journal-article","created":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T02:15:58Z","timestamp":1791771358000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["A Frosty Genetic Screen Unmasks a Major Regulatory Role for\n SHORT VEGETATIVE PHASE<\/scp>\n of Flowering in Response to a Cold Snap"],"prefix":"10.1111","volume":"178","author":[{"given":"Ashleigh","family":"Edwards","sequence":"first","affiliation":[{"name":"Department of Plant & Environmental Sciences University of Copenhagen Frederiksberg Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-5543-8781","authenticated-orcid":false,"given":"Naveen","family":"Shankar","sequence":"additional","affiliation":[{"name":"Ume\u00e5 Plant Science Centre, Department of Plant Physiology Ume\u00e5 University Ume\u00e5 Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Manuela","family":"Jurca","sequence":"additional","affiliation":[{"name":"Ume\u00e5 Plant Science Centre, Department of Plant Physiology Ume\u00e5 University Ume\u00e5 Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1497-9657","authenticated-orcid":false,"given":"Min Kyun","family":"Han","sequence":"additional","affiliation":[{"name":"Department of Systems Biology Institute of Life Science and Biotechnology, Yonsei University Seoul South\u00a0Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8813-0157","authenticated-orcid":false,"given":"Hans","family":"Thordal\u2010Christensen","sequence":"additional","affiliation":[{"name":"Department of Plant & Environmental Sciences University of Copenhagen Frederiksberg Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5764-9423","authenticated-orcid":false,"given":"Stephan","family":"Wenkel","sequence":"additional","affiliation":[{"name":"Department of Plant & Environmental Sciences University of Copenhagen Frederiksberg Denmark"},{"name":"Ume\u00e5 Plant Science Centre, Department of Plant Physiology Ume\u00e5 University Ume\u00e5 Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,10,11]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"crossref","first-page":"627","DOI":"10.1038\/nrg3291","article-title":"The Genetic Basis of Flowering Responses to Seasonal Cues","volume":"13","author":"Andres F.","year":"2012","journal-title":"Nature Reviews. Genetics"},{"key":"e_1_2_9_3_1","first-page":"E2760","article-title":"SHORT VEGETATIVE PHASE Reduces Gibberellin Biosynthesis at the Arabidopsis Shoot Apex to Regulate the Floral Transition","volume":"111","author":"Andr\u00e9s F.","year":"2014","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"key":"e_1_2_9_4_1","doi-asserted-by":"crossref","first-page":"1038","DOI":"10.1105\/tpc.105.039982","article-title":"Salicylic Acid\u2010Independent ENHANCED DISEASE SUSCEPTIBILITY1 Signaling in Arabidopsis Immunity and Cell Death Is Regulated by the Monooxygenase FMO1 and the Nudix Hydrolase NUDT7","volume":"18","author":"Bartsch M.","year":"2006","journal-title":"Plant Cell"},{"key":"e_1_2_9_5_1","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1105\/tpc.15.00910","article-title":"Time\u2010Series Transcriptomics Reveals That AGAMOUS\u2010LIKE22 Affects Primary Metabolism and Developmental Processes in Drought\u2010Stressed Arabidopsis","volume":"28","author":"Bechtold U.","year":"2016","journal-title":"Plant Cell"},{"key":"e_1_2_9_6_1","doi-asserted-by":"crossref","first-page":"168","DOI":"10.1038\/ng1085","article-title":"A Thermosensory Pathway Controlling Flowering Time in \n Arabidopsis thaliana","volume":"33","author":"Bl\u00e1zquez M. A.","year":"2003","journal-title":"Nature Genetics"},{"key":"e_1_2_9_7_1","doi-asserted-by":"crossref","first-page":"1536","DOI":"10.1016\/j.molp.2015.06.009","article-title":"Disruption of the Arabidopsis Defense Regulator Genes SAG101, EDS1, and PAD4 Confers Enhanced Freezing Tolerance","volume":"8","author":"Chen Q. F.","year":"2015","journal-title":"Molecular Plant"},{"key":"e_1_2_9_8_1","doi-asserted-by":"crossref","first-page":"1424","DOI":"10.1105\/tpc.105.031831","article-title":"Comprehensive Interaction Map of the Arabidopsis MADS Box Transcription Factors","volume":"17","author":"Folter S.","year":"2005","journal-title":"Plant Cell"},{"key":"e_1_2_9_9_1","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1038\/ng.806","article-title":"A Framework for Variation Discovery and Genotyping Using Next\u2010Generation DNA Sequencing Data","volume":"43","author":"DePristo M. A.","year":"2011","journal-title":"Nature Genetics"},{"key":"e_1_2_9_10_1","doi-asserted-by":"crossref","first-page":"1349","DOI":"10.1093\/nar\/19.6.1349","article-title":"A Simple and Rapid Method for the Preparation of Plant Genomic DNA for PCR Analysis","volume":"19","author":"Edwards K.","year":"1991","journal-title":"Nucleic Acids Research"},{"key":"e_1_2_9_11_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.108.061531"},{"key":"e_1_2_9_12_1","doi-asserted-by":"crossref","first-page":"993","DOI":"10.1016\/j.jgg.2023.08.004","article-title":"Natural Variation in the SVP Contributes to the Pleiotropic Adaption of \n Arabidopsis thaliana\n Across Contrasted Habitats","volume":"50","author":"Guo X.","year":"2023","journal-title":"Journal of Genetics and Genomics"},{"key":"e_1_2_9_13_1","doi-asserted-by":"crossref","first-page":"351","DOI":"10.1046\/j.1365-313x.2000.00682.x","article-title":"Molecular Cloning of SVP: A Negative Regulator of the Floral Transition in Arabidopsis","volume":"21","author":"Hartmann U.","year":"2000","journal-title":"Plant Journal"},{"key":"e_1_2_9_14_1","doi-asserted-by":"crossref","first-page":"2774","DOI":"10.1101\/gad.1244504","article-title":"PAF1\u2010Complex\u2010Mediated Histone Methylation of FLOWERING LOCUS C Chromatin Is Required for the Vernalization\u2010Responsive, Winter\u2010Annual Habit in Arabidopsis","volume":"18","author":"He Y.","year":"2004","journal-title":"Genes & Development"},{"key":"e_1_2_9_15_1","doi-asserted-by":"crossref","first-page":"1696","DOI":"10.1016\/j.molp.2022.08.007","article-title":"FLOWERING LOCUS M Isoforms Differentially Affect the Subcellular Localization and Stability of SHORT VEGETATIVE PHASE to Regulate Temperature\u2010Responsive Flowering in Arabidopsis","volume":"15","author":"Jin S.","year":"2022","journal-title":"Molecular Plant"},{"key":"e_1_2_9_16_1","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1146\/annurev.cellbio.042308.113411","article-title":"Vernalization: Winter and the Timing of Flowering in Plants","volume":"25","author":"Kim D. H.","year":"2009","journal-title":"Annual Review of Cell and Developmental Biology"},{"key":"e_1_2_9_17_1","doi-asserted-by":"crossref","first-page":"242","DOI":"10.1038\/nature10928","article-title":"Transcription Factor PIF4 Controls the Thermosensory Activation of Flowering","volume":"484","author":"Kumar S. V.","year":"2012","journal-title":"Nature"},{"key":"e_1_2_9_18_1","doi-asserted-by":"crossref","first-page":"946","DOI":"10.1016\/j.csbj.2019.06.014","article-title":"Structural Basis for Plant MADS Transcription Factor Oligomerization","volume":"17","author":"Lai X.","year":"2019","journal-title":"Computational and Structural Biotechnology Journal"},{"key":"e_1_2_9_19_1","doi-asserted-by":"crossref","first-page":"628","DOI":"10.1126\/science.1241097","article-title":"Regulation of Temperature\u2010Responsive Flowering by MADS\u2010Box Transcription Factor Repressors","volume":"342","author":"Lee J. H.","year":"2013","journal-title":"Science"},{"key":"e_1_2_9_20_1","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1101\/gad.1518407","article-title":"Role of SVP in the Control of Flowering Time by Ambient Temperature in Arabidopsis","volume":"21","author":"Lee J. H.","year":"2007","journal-title":"Genes & Development"},{"key":"e_1_2_9_21_1","doi-asserted-by":"crossref","first-page":"1754","DOI":"10.1093\/bioinformatics\/btp324","article-title":"Fast and Accurate Short Read Alignment With Burrows\u2010Wheeler Transform","volume":"25","author":"Li H.","year":"2009","journal-title":"Bioinformatics"},{"key":"e_1_2_9_22_1","doi-asserted-by":"crossref","first-page":"2078","DOI":"10.1093\/bioinformatics\/btp352","article-title":"The Sequence Alignment\/Map Format and SAMtools","volume":"25","author":"Li H.","year":"2009","journal-title":"Bioinformatics"},{"key":"e_1_2_9_23_1","doi-asserted-by":"crossref","first-page":"711","DOI":"10.1016\/j.devcel.2009.03.011","article-title":"Regulation of Floral Patterning by Flowering Time Genes","volume":"16","author":"Liu C.","year":"2009","journal-title":"Developmental Cell"},{"key":"e_1_2_9_24_1","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1186\/s13059-015-0597-1","article-title":"Combinatorial Activities of SHORT VEGETATIVE PHASE and FLOWERING LOCUS C Define Distinct Modes of Flowering Regulation in Arabidopsis","volume":"16","author":"Mateos J. L.","year":"2015","journal-title":"Genome Biology"},{"key":"e_1_2_9_25_1","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1003289","article-title":"The Flowering Repressor SVP Underlies a Novel \n Arabidopsis thaliana\n QTL Interacting With the Genetic Background","volume":"9","author":"M\u00e9ndez\u2010Vigo B.","year":"2013","journal-title":"PLoS Genetics"},{"key":"e_1_2_9_26_1","doi-asserted-by":"crossref","first-page":"414","DOI":"10.1038\/nature12633","article-title":"Temperature\u2010Dependent Regulation of Flowering by Antagonistic FLM Variants","volume":"503","author":"Pos\u00e9 D.","year":"2013","journal-title":"Nature"},{"key":"e_1_2_9_27_1","doi-asserted-by":"publisher","DOI":"10.1023\/a:1025426920923"},{"key":"e_1_2_9_28_1","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1016\/j.plantsci.2018.07.018","article-title":"Overexpression of the MADS\u2010Box Gene K\u2010Domain Increases the Yield Potential of Blueberry","volume":"276","author":"Song G. Q.","year":"2018","journal-title":"Plant Science"},{"key":"e_1_2_9_29_1","doi-asserted-by":"crossref","first-page":"e164","DOI":"10.1093\/nar\/gkq603","article-title":"ANNOVAR: Functional Annotation of Genetic Variants From High\u2010Throughput Sequencing Data","volume":"38","author":"Wang K.","year":"2010","journal-title":"Nucleic Acids Research"},{"key":"e_1_2_9_30_1","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1186\/s13059-017-1378-9","article-title":"Adaptation of \n Arabidopsis thaliana\n to the Yangtze River Basin","volume":"18","author":"Zou Y. P.","year":"2017","journal-title":"Genome Biology"}],"container-title":["Physiologia Plantarum"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/ppl.71146","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1111\/ppl.71146","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/ppl.71146","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T02:16:03Z","timestamp":1791771363000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1111\/ppl.71146"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9]]},"references-count":29,"journal-issue":{"issue":"5","published-print":{"date-parts":[[2026,9]]}},"alternative-id":["10.1111\/ppl.71146"],"URL":"https:\/\/doi.org\/10.1111\/ppl.71146","archive":["Portico"],"relation":{},"ISSN":["0031-9317","1399-3054"],"issn-type":[{"value":"0031-9317","type":"print"},{"value":"1399-3054","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9]]},"assertion":[{"value":"2026-06-26","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-09-09","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-10-11","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e71146"}}