{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T23:51:21Z","timestamp":1791589881224,"version":"4.3.4"},"reference-count":66,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,10,10]],"date-time":"2026-10-10T00:00:00Z","timestamp":1791590400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,10,10]],"date-time":"2026-10-10T00:00:00Z","timestamp":1791590400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"Anhui Provincial Natural Science Foundation","award":["2308085MC96"],"award-info":[{"award-number":["2308085MC96"]}]},{"name":"Anhui Provincial Program on Major Basic Research Project","award":["2023z04020005"],"award-info":[{"award-number":["2023z04020005"]}]},{"name":"Project Entrusted by the Enterprise","award":["KJ24578"],"award-info":[{"award-number":["KJ24578"]}]},{"name":"Project Entrusted by the Enterprise","award":["HX20250345"],"award-info":[{"award-number":["HX20250345"]}]},{"name":"Project Entrusted by the Enterprise","award":["HX20260016"],"award-info":[{"award-number":["HX20260016"]}]},{"DOI":"10.13039\/501100005063","name":"Excellent Young Talents Fund Program of Higher Education Institutions of Anhui Province","doi-asserted-by":"publisher","award":["2024AH030017"],"award-info":[{"award-number":["2024AH030017"]}],"id":[{"id":"10.13039\/501100005063","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Mol Horticulture"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Vernalization, a low-temperature-induced transition from vegetative to reproductive growth, is essential for flowering in many\n                    <jats:italic>Brassica<\/jats:italic>\n                    crops, including Wucai (\n                    <jats:italic>Brassica rapa<\/jats:italic>\n                    ssp.\n                    <jats:italic>chinensis<\/jats:italic>\n                    var.\n                    <jats:italic>rosularis<\/jats:italic>\n                    ). To elucidate the molecular basis of vernalization in Wucai, we performed an integrative multi-omics analysis of shoot apices from cold-treated plants. Twenty-eight vernalization pathway-related genes, including homologs of\n                    <jats:italic>FLC<\/jats:italic>\n                    ,\n                    <jats:italic>FLM<\/jats:italic>\n                    ,\n                    <jats:italic>VIN3<\/jats:italic>\n                    ,\n                    <jats:italic>VRN<\/jats:italic>\n                    ,\n                    <jats:italic>SOC1<\/jats:italic>\n                    ,\n                    <jats:italic>SVP<\/jats:italic>\n                    , and\n                    <jats:italic>VIL2<\/jats:italic>\n                    , exhibited conserved cold-responsive expression patterns. Transcription factors from the MYB, bHLH, bZIP, and NAC families showed dynamic regulation, implicating key members such as\n                    <jats:italic>MYB57<\/jats:italic>\n                    ,\n                    <jats:italic>bHLH3<\/jats:italic>\n                    , and\n                    <jats:italic>ABF4<\/jats:italic>\n                    in flowering control and hormonal signaling. Metabolomic profiling revealed significant enrichment of carbohydrate and amino acid metabolism, with increased levels of sucrose, glucose, galactose, glycine, serine, proline, and arginine, suggesting enhanced mobilization of energy and nitrogen during vernalization. Proline accumulation, together with upregulation of\n                    <jats:italic>P5CS<\/jats:italic>\n                    , indicates a potential role in floral transition. Hormonal analysis showed elevated auxin, kinetin, jasmonic acids, and brassinosteroids, with cold-induced\n                    <jats:italic>BcDET2<\/jats:italic>\n                    expression correlating with increased brassinosteroid levels and early flowering. Exogenous brassinolide further promoted flowering by activating\n                    <jats:italic>FT<\/jats:italic>\n                    and\n                    <jats:italic>SOC1<\/jats:italic>\n                    while repressing\n                    <jats:italic>FLC<\/jats:italic>\n                    . These findings provide a comprehensive molecular framework for vernalization-regulated flowering in Wucai.\n                  <\/jats:p>","DOI":"10.1186\/s43897-026-00246-7","type":"journal-article","created":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T23:01:47Z","timestamp":1791586907000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Integrative multi-omics analyses reveal transcriptional, metabolic, and hormonal regulation of vernalization-induced flowering in Wucai (Brassica rapa)"],"prefix":"10.1186","volume":"6","author":[{"given":"Yunfei","family":"Tian","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Danni","family":"Yang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Na","family":"Liao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kang","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuxin","family":"Yang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinghao","family":"Zhou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ting","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wenjie","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoyan","family":"Tang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chenggang","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lingyun","family":"Yuan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6996-425X","authenticated-orcid":false,"given":"Guohu","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,10,10]]},"reference":[{"key":"246_CR1","doi-asserted-by":"publisher","first-page":"58","DOI":"10.1007\/s11032-023-01405-0","volume":"43","author":"A Akter","year":"2023","unstructured":"Akter A, Kakizaki T, Itabashi E, Kunita K, Shimizu M, Akter A, et al. Characterization of FLOWERING LOCUS C 5 in Brassica rapa L. Mol Breed. 2023;43:58.","journal-title":"Mol Breed"},{"key":"246_CR2","doi-asserted-by":"publisher","first-page":"5439","DOI":"10.1093\/jxb\/erx270","volume":"68","author":"R Bloomer","year":"2017","unstructured":"Bloomer R, Dean C. Fine-tuning timing: natural variation informs the mechanistic basis of the switch to flowering in Arabidopsis thaliana. J Exp Bot. 2017;68:5439\u201352.","journal-title":"J Exp Bot"},{"key":"246_CR3","doi-asserted-by":"publisher","first-page":"649","DOI":"10.1016\/j.molp.2016.11.008","volume":"10","author":"C Cai","year":"2017","unstructured":"Cai C, Wang X, Liu B, Wu J, Liang J, Cui Y, et al. Brassicarapa genome 2.0: a reference upgrade through sequence re-assembly and gene re-annotation. Mol Plant. 2017;10:649\u201351.","journal-title":"Mol Plant"},{"key":"246_CR4","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1016\/j.jplph.2017.03.018","volume":"214","author":"G Campos-Rivero","year":"2017","unstructured":"Campos-Rivero G, Osorio-Montalvo P, S\u00e1nchez-Borges R, Duarte-Ak\u00e9 F, De-la-Pe\u00f1a C. Plant hormone signaling in flowering: an epigenetic point of view. J Plant Physiol. 2017;214:16\u201327.","journal-title":"J Plant Physiol"},{"key":"246_CR5","doi-asserted-by":"publisher","first-page":"241","DOI":"10.3389\/fpls.2019.00241","volume":"10","author":"P Carillo","year":"2019","unstructured":"Carillo P, Arena C, Modarelli G, Pascale S, Paradiso R. Photosynthesis in Ranunculus asiaticus L.: the influence of the hybrid and the preparation procedure of tuberous roots. Front Plant Sci. 2019;10:241.","journal-title":"Front Plant Sci"},{"key":"246_CR6","doi-asserted-by":"crossref","unstructured":"Chen G, Ye X, Zhang S, Zhu S, Yuan L, Hou J, et al. Comparative transcriptome analysis between fertile and CMS flower buds in Wucai (Brassica campestris L.). BMC Genomics. 2018;19:908.","DOI":"10.1186\/s12864-018-5331-4"},{"key":"246_CR7","doi-asserted-by":"crossref","unstructured":"Chen G, Ye X, Zeng F, Wang J, Yuan L, Zhu S, et al. Characterization and utilization of a cytoplasmic male sterility line of Wucai (Brassica campestris L.). Hortic Environ Biote. 2019;60:373\u201482.","DOI":"10.1007\/s13580-019-00127-7"},{"key":"246_CR8","doi-asserted-by":"publisher","first-page":"401","DOI":"10.1186\/s12864-020-06802-9","volume":"21","author":"G Chen","year":"2020","unstructured":"Chen G, Wang J, Wang H, Wang C, Tang X, Li J, et al. Genome-wide analysis of proline-rich extension-like receptor protein kinase (PERK) in Brassica rapa and its association with the pollen development. BMC Genomics. 2020;21:401.","journal-title":"BMC Genomics"},{"key":"246_CR9","doi-asserted-by":"publisher","first-page":"77","DOI":"10.1186\/s12870-025-06080-1","volume":"25","author":"H Chen","year":"2025","unstructured":"Chen H, Zhang S, Chang J, Wei H, Li H, Li C, et al. Foliar application of 24-epibrassinolide enhances leaf nicotine content under low temperature conditions during the mature stage of flue-cured tobacco by regulating cold stress tolerance. BMC Plant Biol. 2025;25:77.","journal-title":"BMC Plant Biol"},{"key":"246_CR10","doi-asserted-by":"publisher","first-page":"479","DOI":"10.1093\/plphys\/kiae022","volume":"195","author":"L Chu","year":"2024","unstructured":"Chu L, Zheng W, Liu H, Sheng X, Wang Q, Wang Y, et al. ACC SYNTHASE4 inhibits gibberellin biosynthesis and FLOWERING LOCUS T expression during citrus flowering. Plant Physiol. 2024;195:479\u2013501.","journal-title":"Plant Physiol"},{"key":"246_CR11","doi-asserted-by":"publisher","first-page":"1605","DOI":"10.1111\/jipb.13507","volume":"65","author":"G Cui","year":"2023","unstructured":"Cui G, Li D, Zhang L, Xia C, Kong X, Liu X. GSK3 regulates VRN1 to control flowering time in wheat. J Integr Plant Biol. 2023;65:1605\u20138.","journal-title":"J Integr Plant Biol"},{"key":"246_CR12","doi-asserted-by":"crossref","unstructured":"Dai Y, Zhang S, Guan J, Wang S, Zhang H, Li G, et al. Single-cell transcriptomic analysis of flowering regulation and vernalization in Chinese cabbage shoot apex. Hortic Res. 2024;11:uhae214.","DOI":"10.1093\/hr\/uhae214"},{"key":"246_CR13","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1016\/j.pbi.2017.08.003","volume":"41","author":"T Do","year":"2018","unstructured":"Do T, Martinoia E, Lee Y. Functions of ABC transporters in plant growth and development. Curr Opin Plant Biol. 2018;41:32\u20138.","journal-title":"Curr Opin Plant Biol"},{"key":"246_CR14","doi-asserted-by":"publisher","first-page":"102049","DOI":"10.1016\/j.pbi.2021.102049","volume":"63","author":"S Freytes","year":"2021","unstructured":"Freytes S, Canelo M, Cerd\u00e1n P. Regulation of flowering time: when and where? Curr Opin Plant Biol. 2021;63:102049.","journal-title":"Curr Opin Plant Biol"},{"key":"246_CR15","doi-asserted-by":"crossref","first-page":"3288","DOI":"10.3390\/plants12183288","volume":"12","author":"G Fusco","year":"2023","unstructured":"Fusco G, Carillo P, Nicastro R, Pagkiaro L, Pascale S, Paradiso R. Metabolic profiling in tuberous roots of Ranunculu sasiaticus L. as influenced by vernalization procedure. Plants. 2023;12:3288.","journal-title":"Plants"},{"key":"246_CR16","doi-asserted-by":"publisher","first-page":"1048","DOI":"10.1111\/tpj.15613","volume":"109","author":"X Gao","year":"2022","unstructured":"Gao X, Zhang J, Li J, Wang Y, Zhang R, Du H, et al. The phosphoproteomic and interactomic landscape of qGL3\/OsPPKL1-mediated brassinosteroid signaling in rice. Plant J. 2022;109:1048\u201363.","journal-title":"Plant J"},{"key":"246_CR17","doi-asserted-by":"publisher","first-page":"3092","DOI":"10.3390\/ijms21093092","volume":"21","author":"J Gu","year":"2020","unstructured":"Gu J, Zeng Z, Wang Y, Lyu Y. Transcriptome analysis of carbohydrate metabolism genes and molecular regulation of sucrose transport gene LoSUT on the flowering process of developing oriental hybrid Lily \u201cSorbonne\u201d bulb. Int J Mol Sci. 2020;21:3092.","journal-title":"Int J Mol Sci"},{"key":"246_CR18","doi-asserted-by":"publisher","first-page":"639","DOI":"10.1038\/s41467-018-03065-7","volume":"9","author":"J Hepworth","year":"2018","unstructured":"Hepworth J, Antoniou-Kourounioti R, Bloomer R, Selga C, Berggren K, Cox D, et al. Absence of warmth permits epigenetic memory of winter in Arabidopsis. Nat Commun. 2018;9:639.","journal-title":"Nat Commun"},{"key":"246_CR19","doi-asserted-by":"publisher","first-page":"1286","DOI":"10.3390\/agriculture11121286","volume":"11","author":"J Hong","year":"2021","unstructured":"Hong J, Suh E, Park S, Park J, Lee Y. Multiplex CRISPR\/Cas9 mutagenesis of BrVRN1 delays flowering time in Chinese cabbage (Brassica rapa L. ssp. pekinensis). Agriculture. 2021a;11:1286.","journal-title":"Agriculture"},{"key":"246_CR20","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1111\/jipb.13071","volume":"63","author":"L Hong","year":"2021","unstructured":"Hong L, Niu F, Lin Y, Wang S, Chen L, Jiang L. MYB106 is a negative regulator and a substrate for CRL3BPM E3 ligase in regulating flowering time in Arabidopsis thaliana. J Integr Plant Biol. 2021b;63:16.","journal-title":"J Integr Plant Biol"},{"key":"246_CR21","doi-asserted-by":"publisher","first-page":"108198","DOI":"10.1016\/j.plaphy.2023.108198","volume":"205","author":"M Hosseini","year":"2023","unstructured":"Hosseini M, Saidi A, Maali-Amiri R, Khosravi-Nejad F, Abbasi A. Low-temperature acclimation related with developmental regulations of polyamines and ethylene metabolism in wheat recombinant inbred lines. Plant Physiol Biochem. 2023;205:108198.","journal-title":"Plant Physiol Biochem"},{"key":"246_CR22","doi-asserted-by":"publisher","first-page":"947","DOI":"10.1111\/nph.19680","volume":"242","author":"P Huang","year":"2024","unstructured":"Huang P, Schmitt J, Runcie D. Exploring the molecular regulation of vernalization-induced flowering synchrony in Arabidopsis. New Phytol. 2024;242:947\u201359.","journal-title":"New Phytol"},{"key":"246_CR23","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1016\/j.jplph.2016.04.006","volume":"203","author":"N H\u00fcner","year":"2016","unstructured":"H\u00fcner N, Dahal K, Bode R, Kurepin L, Ivanov A. Photosynthetic acclimation, vernalization, crop productivity and \u2018the grand design of photosynthesis.\u2019 J Plant Physiol. 2016;203:29\u201343.","journal-title":"J Plant Physiol"},{"key":"246_CR24","doi-asserted-by":"publisher","first-page":"489","DOI":"10.1016\/j.molp.2019.01.002","volume":"12","author":"K Hwang","year":"2019","unstructured":"Hwang K, Susila H, Nasim Z, Jung J, Ahn J. Arabidopsis ABF3 and ABF4 transcription factors act with the NF-YC complex to regulate SOC1 expression and mediate drought-accelerated flowering. Mol Plant. 2019;12:489\u2013505.","journal-title":"Mol Plant"},{"key":"246_CR25","doi-asserted-by":"publisher","first-page":"e70287","DOI":"10.1111\/ppl.70287","volume":"177","author":"M James","year":"2025","unstructured":"James M, Masclaux-Daubresse C, Goux D, Jing L, Etienne P, Trouverie J. The taproot acts as a storage organ during rapeseed vernalization. Physiol Plant. 2025;177:e70287.","journal-title":"Physiol Plant"},{"key":"246_CR26","doi-asserted-by":"publisher","first-page":"157","DOI":"10.1007\/s12374-021-09344-z","volume":"65","author":"H Kang","year":"2022","unstructured":"Kang H, Nugroho A, Park M, Kim J, Lee S, Moon H, et al. Vernalization regulates flowering genes and modulates glucosinolates biosynthesis in Chinese cabbage. J Plant Biol. 2022;65:157\u201373.","journal-title":"J Plant Biol"},{"key":"246_CR27","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1007\/s11103-021-01231-y","volume":"108","author":"S Kim","year":"2022","unstructured":"Kim S, Kim J, Kang H, Kim D. A premature stop codon in BrFLC2 transcript results in early flowering in oilseed-type Brassica rapa plants. Plant Mol Biol. 2022;108:241\u201355.","journal-title":"Plant Mol Biol"},{"key":"246_CR28","doi-asserted-by":"publisher","first-page":"949","DOI":"10.1093\/jxb\/ery423","volume":"70","author":"A Lazaro","year":"2019","unstructured":"Lazaro A, Zhou Y, Giesguth M, Nawaz K, Bergonzi S, Pecinka A, et al. PERPETUAL FLOWERING2 coordinates the vernalization response and perennial flowering in Arabis alpina. J Exp Bot. 2019;70:949\u201361.","journal-title":"J Exp Bot"},{"key":"246_CR29","doi-asserted-by":"publisher","first-page":"2269","DOI":"10.1073\/pnas.1423585112","volume":"112","author":"J Lee","year":"2015","unstructured":"Lee J, Yun J, Zhao W, Shen W, Amasino R. A methyltransferase required for proper timing of the vernalization response in Arabidopsis. Proc Natl Acad Sci U S A. 2015;112:2269\u201374.","journal-title":"Proc Natl Acad Sci U S A"},{"key":"246_CR30","doi-asserted-by":"publisher","first-page":"1135","DOI":"10.1016\/j.molp.2018.06.007","volume":"11","author":"Z Li","year":"2018","unstructured":"Li Z, Ou Y, Zhang Z, Li J, He Y. Brassinosteroid signaling recruits histone 3 Lysine-27 demethylation activity to FLOWERING LOCUS C chromatin to inhibit the floral transition in Arabidopsis. Mol Plant. 2018;11:1135\u201346.","journal-title":"Mol Plant"},{"key":"246_CR31","doi-asserted-by":"publisher","first-page":"334","DOI":"10.1007\/s11105-019-01161-6","volume":"37","author":"L Li","year":"2019","unstructured":"Li L, Wu P, Wu X, Hou X, Wang H, Liu T. Molecular mechanism, evolution, expression, and functional analyses of MAF\/FLC clade members in Pak-Choi (Brassica rapa ssp. chinensis). Plant Mol Biol Rep. 2019;37:334\u201346.","journal-title":"Plant Mol Biol Rep"},{"key":"246_CR32","doi-asserted-by":"publisher","first-page":"331","DOI":"10.1186\/s12870-020-02539-5","volume":"20","author":"Y Li","year":"2020","unstructured":"Li Y, Xiong H, Guo H, Zhou C, Xie Y, Zhao L, et al. Identification of the vernalization gene VRN-B1 responsible for heading date variation by QTL mapping using a RIL population in wheat. BMC Plant Biol. 2020;20:331.","journal-title":"BMC Plant Biol"},{"key":"246_CR33","doi-asserted-by":"publisher","first-page":"7940","DOI":"10.3390\/ijms23147940","volume":"23","author":"X Li","year":"2022","unstructured":"Li X, Wu Y, Chi H, Wei H, Wang H, Yu S. Genomewide identification and characterization of the genes involved in the flowering of cotton. Int J Mol Sci. 2022;23:7940.","journal-title":"Int J Mol Sci"},{"key":"246_CR34","doi-asserted-by":"publisher","first-page":"4180","DOI":"10.1093\/jxb\/erae105","volume":"75","author":"X Li","year":"2024","unstructured":"Li X, Lin C, Lan C, Tao Z. Genetic and epigenetic basis of phytohormones control of floral transition in plants. J Exp Bot. 2024;75:4180\u201394.","journal-title":"J Exp Bot"},{"key":"246_CR35","doi-asserted-by":"publisher","first-page":"933923","DOI":"10.3389\/fpls.2022.933923","volume":"13","author":"Q Liang","year":"2022","unstructured":"Liang Q, Song K, Lu M, Dai T, Yang J, Wan J, et al. Transcriptome and metabolome analyses reveal the involvement of multiple pathways in flowering intensity in mango. Front Plant Sci. 2022;13:933923.","journal-title":"Front Plant Sci"},{"key":"246_CR36","doi-asserted-by":"publisher","first-page":"2000125","DOI":"10.1002\/sstr.202000125","volume":"2","author":"L Liu","year":"2021","unstructured":"Liu L, Xuan L, Jiang Y, Hu H. Regulation by FLOWERING LOCUS T and TERMINAL FLOWER 1 in flowering time and plant architecture. Small Struct. 2021;2:2000125.","journal-title":"Small Struct"},{"key":"246_CR37","doi-asserted-by":"publisher","first-page":"853444","DOI":"10.3389\/fpls.2022.853444","volume":"13","author":"X Liu","year":"2022","unstructured":"Liu X, Luo M, Li M, Wei J. Depicting precise temperature and duration of vernalization and inhibiting early bolting and flowering of Angelica sinensis by freezing storage. Front Plant Sci. 2022;13:853444.","journal-title":"Front Plant Sci"},{"key":"246_CR38","doi-asserted-by":"publisher","first-page":"2251","DOI":"10.1007\/s11427-024-2664-0","volume":"67","author":"X Liu","year":"2024","unstructured":"Liu X, Deng M, Shi B, Zhu K, Chen J, Xu S, et al. Distinct roles of H3K27me3 and H3K36me3 in vernalization response, maintenance, and resetting in winter wheat. Sci China Life Sci. 2024a;67:2251\u201366.","journal-title":"Sci China Life Sci"},{"key":"246_CR39","doi-asserted-by":"publisher","first-page":"306","DOI":"10.1186\/s13059-024-03342-3","volume":"25","author":"Y Liu","year":"2024","unstructured":"Liu Y, Liu P, Gao L, Li Y, Ren X, Jia J, et al. Epigenomic identification of vernalization cis-regulatory elements in winter wheat. Genome Biol. 2024b;25:306.","journal-title":"Genome Biol"},{"key":"246_CR40","doi-asserted-by":"publisher","first-page":"105398","DOI":"10.1016\/j.jprot.2025.105398","volume":"314","author":"X Liu","year":"2025","unstructured":"Liu X, Liao N, Tang X, Wang K, Wang W, Khan A, et al. TMT-label comparative proteomics reveals the vernalization mechanism in Wucai (Brassica campestris L.). J Proteomics. 2025;314:105398.","journal-title":"J Proteomics"},{"key":"246_CR41","doi-asserted-by":"publisher","first-page":"e1010766","DOI":"10.1371\/journal.pgen.1010766","volume":"19","author":"D Martignago","year":"2023","unstructured":"Martignago D, Falavigna V, Lombardi A, Gao H, Krukowski P, Galbiati M, et al. The bZIP transcription factor AREB3 mediates FT signalling and floral transition at the Arabidopsis shoot apical meristem. PLoS Genet. 2023;19:e1010766.","journal-title":"PLoS Genet"},{"key":"246_CR42","doi-asserted-by":"publisher","first-page":"1016","DOI":"10.4161\/psb.4.11.9797","volume":"4","author":"R Mattioli","year":"2009","unstructured":"Mattioli R, Costantino P, Trovato M. Proline accumulation in plants not only stress. Plant Signal Behav. 2009;4:1016\u20138.","journal-title":"Plant Signal Behav"},{"key":"246_CR43","doi-asserted-by":"publisher","first-page":"2348","DOI":"10.3390\/plants11182348","volume":"11","author":"R Mattioli","year":"2022","unstructured":"Mattioli R, Francioso A, Trovato M. Proline affects flowering time in Arabidopsis by modulating FLC expression: a clue of epigenetic regulation? Plants. 2022;11:2348.","journal-title":"Plants"},{"key":"246_CR44","doi-asserted-by":"publisher","first-page":"108842","DOI":"10.1016\/j.plaphy.2024.108842","volume":"213","author":"F Niu","year":"2024","unstructured":"Niu F, Rehmani M, Yan J. Multilayered regulation and implication of flowering time in plants. Plant Physiol Bioch. 2024;213:108842.","journal-title":"Plant Physiol Bioch"},{"key":"246_CR45","doi-asserted-by":"publisher","first-page":"24023","DOI":"10.1038\/s41598-021-03557-5","volume":"11","author":"A Nugroho","year":"2021","unstructured":"Nugroho A, Lee S, Pervitasari A, Moon H, Choi D, Kim J, et al. Transcriptomic and metabolic analyses revealed the modulatory effect of vernalization on glucosinolate metabolism in radish (Raphanu ssativus L.). Sci Rep. 2021;11:24023.","journal-title":"Sci Rep"},{"key":"246_CR46","doi-asserted-by":"publisher","first-page":"e14199","DOI":"10.1111\/ppl.14199","volume":"176","author":"S Patra","year":"2024","unstructured":"Patra S, Chatterjee D, Dutta R, Mandal A. Abiotic and biotic factors regulate the timing of floral induction: a review. Physiol Plantarum. 2024;176:e14199.","journal-title":"Physiol Plantarum"},{"key":"246_CR47","doi-asserted-by":"publisher","first-page":"188","DOI":"10.1186\/s12870-024-04887-y","volume":"24","author":"C Qiu","year":"2024","unstructured":"Qiu C, Wang T, Wang H, Tao Z, Wang C, Ma J, et al. SISTER OF FCA physically associates with SKB1 to regulate flowering time in Arabidopsis thaliana. BMC Plant Biol. 2024;24:188.","journal-title":"BMC Plant Biol"},{"key":"246_CR48","doi-asserted-by":"publisher","first-page":"3206","DOI":"10.1105\/tpc.20.00072","volume":"32","author":"C Rodrigues","year":"2020","unstructured":"Rodrigues C, M\u00fcdsam C, Keller I, Zierer W, Czarnecki O, Corral J, et al. Vernalization alters sink and source identities and reverses phloem translocation from taproots to shoots in sugar beet. Plant Cell. 2020;32:3206\u201323.","journal-title":"Plant Cell"},{"key":"246_CR49","doi-asserted-by":"publisher","first-page":"951","DOI":"10.1093\/plcell\/koab308","volume":"34","author":"P Salom\u00e9","year":"2022","unstructured":"Salom\u00e9 P. The circadian clock reaches vernalization: how CCA1 and LHY induce VIN3 transcription during winter. Plant Cell. 2022;34:951\u20132.","journal-title":"Plant Cell"},{"key":"246_CR50","doi-asserted-by":"publisher","first-page":"1533","DOI":"10.3389\/fpls.2017.01533","volume":"8","author":"M Shang","year":"2017","unstructured":"Shang M, Wang X, Zhang J, Qi X, Ping A, Hou L, et al. Genetic regulation of GA metabolism during vernalization, floral bud initiation and development in Pak Choi (Brassica rapa ssp. chinensis Makino). Front Plant Sci. 2017;8:1533.","journal-title":"Front Plant Sci"},{"key":"246_CR51","doi-asserted-by":"publisher","first-page":"818","DOI":"10.1093\/plcell\/koab286","volume":"34","author":"L Shen","year":"2022","unstructured":"Shen L, Zhang Y, Sawettalake N. A molecular switch for FLOWERING LOCUS C activation determines flowering time in Arabidopsis. Plant Cell. 2022;34:818\u201333.","journal-title":"Plant Cell"},{"key":"246_CR52","doi-asserted-by":"publisher","first-page":"819","DOI":"10.1186\/s12864-021-08110-2","volume":"22","author":"F Shi","year":"2021","unstructured":"Shi F, Xu H, Liu C, Tan C, Ren J, Ye X, et al. Whole-transcriptome sequencing reveals a vernalization-related ceRNA regulatory network in Chinese cabbage (Brassica campestris L. ssp. pekinensis). BMC Genomics. 2021;22:819.","journal-title":"BMC Genomics"},{"key":"246_CR53","doi-asserted-by":"publisher","first-page":"uhad174","DOI":"10.1093\/hr\/uhad174","volume":"10","author":"M Shi","year":"2023","unstructured":"Shi M, Wang C, Wang P, Yun F, Liu Z, Ye F, et al. Role of methylation in vernalization and photoperiod pathway: a potential flowering regulator? Hortic Res. 2023;10:uhad174.","journal-title":"Hortic Res"},{"key":"246_CR54","doi-asserted-by":"publisher","first-page":"160","DOI":"10.1104\/pp.107.102632","volume":"145","author":"T Slotte","year":"2007","unstructured":"Slotte T, Holm K, McIntyre L, Lagercrantz U, Lascoux M. Differential expression of genes important for adaptation in Capsella bursa-pastoris (Brassicaceae). Plant Physiol. 2007;145:160\u201373.","journal-title":"Plant Physiol"},{"key":"246_CR55","doi-asserted-by":"publisher","first-page":"e0141446","DOI":"10.1371\/journal.pone.0141446","volume":"10","author":"M Sun","year":"2015","unstructured":"Sun M, Qi X, Hou L, Xu X, Zhu Z, Li M. Gene expression analysis of pak choi in response to vernalization. PLoS ONE. 2015;10:e0141446.","journal-title":"PLoS ONE"},{"key":"246_CR56","doi-asserted-by":"publisher","first-page":"1549","DOI":"10.1104\/pp.19.00021","volume":"180","author":"V Tilmes","year":"2019","unstructured":"Tilmes V, Mateos J, Madrid E, Vincent C, Severing E, Carrera E, et al. Gibberellins act downstream of Arabis PERPETUAL FLOWERING1 to accelerate floral induction during vernalization. Plant Physiol. 2019;180:1549\u201363.","journal-title":"Plant Physiol"},{"key":"246_CR57","doi-asserted-by":"publisher","first-page":"2679","DOI":"10.1007\/s12298-021-01110-2","volume":"27","author":"P Wang","year":"2021","unstructured":"Wang P, Liu D, Yang F, Ge H, Zhao X, Chen H, et al. Identification of key gene networks controlling vernalization development characteristics of Isatis indigotica by full-length transcriptomes and gene expression profiles. Physiol Mol Biol Plants. 2021;27:2679\u201393.","journal-title":"Physiol Mol Biol Plants"},{"key":"246_CR58","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1186\/s12863-023-01144-3","volume":"24","author":"J Wang","year":"2023","unstructured":"Wang J, Sun L, Zhang H, Jiao B, Wang H, Zhou S. Transcriptome analysis during vernalization in wheat (Triticum aestivum L.). BMC Genom Data. 2023a;24:43.","journal-title":"BMC Genom Data"},{"key":"246_CR59","doi-asserted-by":"publisher","first-page":"1516","DOI":"10.1111\/nph.18779","volume":"238","author":"Y Wang","year":"2023","unstructured":"Wang Y, Zhu Y, Jiang H, Mao Z, Zhang J, Fang H, et al. The regulatory module MdBZR1\u2013MdCOL6 mediates brassinosteroid- and light-regulated anthocyanin synthesis in apple. New Phytol. 2023b;238:1516\u201333.","journal-title":"New Phytol"},{"key":"246_CR60","doi-asserted-by":"publisher","first-page":"uhae013","DOI":"10.1093\/hr\/uhae013","volume":"11","author":"T Wu","year":"2024","unstructured":"Wu T, Liu Z, Yu T, Zhou R, Yang Q, Cao R, et al. Flowering genes identification, network analysis, and database construction for 837 plants. Hortic Res. 2024;11:uhae013.","journal-title":"Hortic Res"},{"key":"246_CR61","doi-asserted-by":"publisher","first-page":"1576519","DOI":"10.3389\/fpls.2025.1576519","volume":"16","author":"L Zeng","year":"2025","unstructured":"Zeng L, Meng W, Zhong X, Peng J, Ynag X, Sun L. Transcriptome analyses reveal the flowering regulatory networks in the desert ephemeral plant Eremopyrum triticeum. Front Plant Sci. 2025;16:1576519.","journal-title":"Front Plant Sci"},{"key":"246_CR62","doi-asserted-by":"publisher","first-page":"134","DOI":"10.1111\/tpj.15783","volume":"111","author":"C Zhang","year":"2022","unstructured":"Zhang C, Zhou Q, Liu W, Wu X, Li Z, Xu Y. BrABF3 promotes flowering through the direct activation of CONSTANS transcription in pak choi. Plant J. 2022;111:134\u201348.","journal-title":"Plant J"},{"key":"246_CR63","doi-asserted-by":"publisher","first-page":"3470","DOI":"10.1093\/plcell\/koad166","volume":"35","author":"C Zhang","year":"2023","unstructured":"Zhang C, Jian M, Li W, Yao X, Tan C, Qian Q, et al. Gibberellin signaling modulates flowering via the DELLA-BRAHMA-NF-YC module in Arabidopsis. Plant Cell. 2023a;35:3470\u201384.","journal-title":"Plant Cell"},{"key":"246_CR64","doi-asserted-by":"publisher","first-page":"E27","DOI":"10.1038\/s41586-023-06189-z","volume":"619","author":"Z Zhang","year":"2023","unstructured":"Zhang Z, Luo X, Yang Y, He Y. Cold induction of nuclear FRIGIDA condensation in Arabidopsis. Nature. 2023b;619:E27\u201332.","journal-title":"Nature"},{"key":"246_CR65","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1093\/jxb\/erab418","volume":"73","author":"L Zhao","year":"2022","unstructured":"Zhao L, Li X, Chen W, Xu Z, Chen M, Wang H, et al. The emerging role of jasmonate in the control of flowering time. J Exp Bot. 2022;73:11\u201321.","journal-title":"J Exp Bot"},{"key":"246_CR66","doi-asserted-by":"publisher","first-page":"1651","DOI":"10.1111\/nph.18837","volume":"238","author":"W Zhao","year":"2023","unstructured":"Zhao W, Liang J, Huang H, Yang J, Feng J, Sun L, et al. Tomato defence against Meloidogyne incognita by jasmonic acid-mediated fine-tuning of kaempferol homeostasis. New Phytol. 2023;238:1651\u201370.","journal-title":"New Phytol"}],"container-title":["Molecular Horticulture"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s43897-026-00246-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s43897-026-00246-7","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s43897-026-00246-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T23:01:50Z","timestamp":1791586910000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s43897-026-00246-7"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,10]]},"references-count":66,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["246"],"URL":"https:\/\/doi.org\/10.1186\/s43897-026-00246-7","relation":{},"ISSN":["2730-9401"],"issn-type":[{"value":"2730-9401","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,10]]},"assertion":[{"value":"9 October 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 April 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"10 October 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Not applicable.","order":1,"name":"Ethics","label":"Ethics approval and consent to participate","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"All authors approve the manuscript and consent to the publication of the work.","order":2,"name":"Ethics","label":"Consent for publication","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare that they have no competing interests.","order":3,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"72"}}