Article(id=1304414728781062720, tenantId=1146029695717560320, journalId=1302319053441957962, issueId=1304414700964443026, articleNumber=null, orderNo=null, doi=10.7501/j.issn.0253-2670.2026.04.022, pmid=null, cstr=null, oa=null, hot=0, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1759334400000, receivedDateStr=2025-10-02, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1788926306194, onlineDateStr=2026-09-09, pubDate=null, pubDateStr=null, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1788926306194, onlineIssueDateStr=2026-09-09, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1788926306194, creator=13701087609, updateTime=1788926306194, updator=13701087609, issue=Issue{id=1304414700964443026, tenantId=1146029695717560320, journalId=1302319053441957962, year='2026', volume='57', issue='4', pageStart='1209', pageEnd='1596', issueExtLink='null', onlineDate='null', pubDate='1772208000000', pubDateStr='2026-02-28', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1788926299563, creator='13701087609', updateTime=1788926573099, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1304415848316297970, tenantId=1146029695717560320, journalId=1302319053441957962, issueId=1304414700964443026, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1304415848316297971, tenantId=1146029695717560320, journalId=1302319053441957962, issueId=1304414700964443026, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=1460, endPage=1472, ext={EN=ArticleExt(id=1304414730605584962, articleId=1304414728781062720, tenantId=1146029695717560320, journalId=1302319053441957962, language=EN, title=Identification of R2R3-MYB gene family in Chrysanthemum indicum var. aromaticum and analysis of their expression patterns in response to UV-B stress, columnId=null, journalTitle=Chinese Traditional and Herbal Drugs, columnName=null, runingTitle=null, highlight=null, articleAbstract=Objective To identify members of the R2R3-MYB gene family (named CiaMYB) in Chrysanthemum indicum var. aromaticum, a folk medicinal plant from Shennongjia, analyze their genetic evolutionary characteristics and expression patterns in response to UV-B stress, and explore the molecular mechanisms by which they regulate flavonoid synthesis. Methods Based on the whole-genome sequence of C. indicum var. aromaticum, bioinformatics analyses were performed to comprehensively identify CiaMYB members, and systematic analyses were conducted on their protein physicochemical properties, chromosomal localization, gene structure, collinearity relationships, and composition of cis-acting elements. Combined with transcriptome sequencing and correlation analysis, CiaMYB genes involved in flavonoid biosynthesis were screened. Quantitative reverse transcription polymerase chain reaction (qRT-PCR) was used to verify the expression patterns of key genes, and the functional localization of the proteins was analyzed. Results A total of 140 CiaMYB genes were identified, encoding hydrophilic proteins consisting of 206—868 amino acids, which are unevenly distributed on nine chromosomes. The promoter regions are enriched with cis-acting elements related to light response, abscisic acid (ABA) and methyl jasmonate (MeJA) signaling pathways. UV-B irradiation significantly induces differential expression of CiaMYB genes and accumulation of total flavonoids in leaves. Among them, CiaMYB040 and CiaMYB066 are highly homologous to the S7 subfamily, which is critical for flavonol synthesis in Arabidopsis thaliana, and their expression levels show a strong positive correlation with total flavonoid content, suggesting that they play a key transcriptional regulatory role in the flavonoid synthesis pathway. Subcellular localization confirmed that CiaMYB040 and CiaMYB066 proteins are localized in the nucleus. Conclusion This study is the first to systematically reveal the evolutionary characteristics of R2R3-MYB family members in C. indicum var. aromaticum, and preliminarily screen CiaMYB genes that respond to UV-B stress and are involved in regulating flavonoid synthesis. It provides important candidate gene resources for elucidating the molecular mechanism of plant high-altitude stress adaptation, as well as for flavonoid metabolism regulation and stress-resistant genetic improvement in Asteraceae plants., authors=HE Ye, MEI Yang, WANG Miaomiao, YANG Jingyue, ZHANG Jingjing, LIU Yifei, authorsList=HE Ye, MEI Yang, WANG Miaomiao, YANG Jingyue, ZHANG Jingjing, LIU Yifei, authorCompany=null, correspAuthors=null, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=null), CN=ArticleExt(id=1304414730509115969, articleId=1304414728781062720, tenantId=1146029695717560320, journalId=1302319053441957962, language=CN, title=神农香菊R2R3-MYB基因家族鉴定及其响应UV-B胁迫的表达模式分析, columnId=1304140203967410414, journalTitle=中草药, columnName=药材与资源, runingTitle=null, highlight=null, articleAbstract=目的 旨在鉴定神农架民间药用植物神农香菊Chrysanthemum indicum var. aromaticumR2R3-MYB基因家族成员(命名为CiaMYB),解析其基因进化特征及响应UV-B胁迫的表达特征模式,探讨其调控黄酮类化合物合成的分子机制。方法 基于神农香菊全基因组序列,通过生物信息学分析全面鉴定CiaMYB家族成员,系统分析其蛋白理化性质、染色体定位、基因结构、共线性关系及顺式作用元件组成;结合转录组测序与相关性分析,筛选参与黄酮类化合物生物合成的CiaMYB基因;利用实时荧光定量PCR(quantitative reverse transcription polymerase chain reaction,qRT-PCR)验证关键基因表达模式,并解析蛋白功能定位。结果 共鉴定出140个CiaMYB基因,其编码蛋白为206~868个氨基酸组成的亲水性蛋白,不均匀分布于9条染色体中。启动子区富集光响应、脱落酸(abscisic acid,ABA)和茉莉酸甲酯(methyl jasmonate,MeJA)信号通路相关顺式作用元件。UV-B辐照显著诱导CiaMYB基因差异表达和叶片总黄酮积累,其中CiaMYB040CiaMYB066与拟南芥黄酮醇合成关键的S7亚家族高度同源,且表达量与总黄酮含量呈极强正相关,提示其在黄酮合成途径中发挥关键转录调控作用。亚细胞定位证实CiaMYB040和CiaMYB066蛋白定位于细胞核。结论 首次系统揭示神农香菊R2R3-MYB家族成员的进化特征,初步筛选了响应UV-B胁迫参与调控黄酮类物质合成的CiaMYB基因,为解析植物高海拔抗逆适应的分子机制,菊科植物黄酮代谢调控及抗逆遗传改良提供了重要的候选基因资源。, authors=何叶1, 梅洋1, 汪苗苗1, 杨静月1, 张景景1,2, 刘义飞1,2, authorsList=何叶, 梅洋, 汪苗苗, 杨静月, 张景景, 刘义飞, authorCompany=1 湖北中医药大学药学院, 湖北 武汉 430065;
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雷霄飞, 杨学领. 神农香菊化学成分及药理作用研究进展[J]. 现代农业科技, 2014(19): 312-313.
Wang X, Zhang J J, Liu Z G, et al. Comparative transcriptome analysis of three chrysanthemums provides insights into flavonoid and terpenoid biosynthesis [J]. J Plant Biol, 2021, 64(5): 389-401.
Chen K L, Liu Y M, Zhang Y, et al. A special aromatic Chrysanthemum breed with high content of thujone [J]. Phcog Mag, 2020, 16(71): 625.
Feng Y, Li B, Hu J, et al. Sustainable use of Chrysanthemum indicum var. aromaticum as value-added green materials in microemulsion hydrogels [J]. ACS Sustainable Chem Eng, 2023, 11(9): 3820-3831.
Shi C, Liu H T. How plants protect themselves from ultraviolet-B radiation stress [J]. Plant Physiol, 2021, 187(3): 1096-1103.
Sztatelman O, Grzyb J, Gabryś H, et al. The effect of UV-B on Arabidopsis leaves depends on light conditions after treatment [J]. BMC Plant Biol, 2015, 15: 281.
Xiong Y, Yuan S, Xiong Y L, et al. Analysis of allohexaploid wheatgrass genome reveals its Y haplome origin in Triticeae and high-altitude adaptation [J]. Nat Commun, 2025, 16(1): 3104.
Ferreyra M L F, Serra P, Casati P. Recent advances on the roles of flavonoids as plant protective molecules after UV and high light exposure [J]. Physiol Plant, 2021, 173(3): 736-749.
Takshak S, Agrawal S B. Defense potential of secondary metabolites in medicinal plants under UV-B stress [J]. J Photochem Photobiol B, 2019, 193: 51-88.
Liang T, Shi C, Peng Y, et al. Brassinosteroid-activated BRI1-EMS-SUPPRESSOR 1 inhibits flavonoid biosynthesis and coordinates growth and UV-B stress responses in plants [J]. Plant Cell, 2020, 32(10): 3224-3239.
胡雅丹, 伍国强, 刘晨, 等. MYB转录因子在调控植物响应逆境胁迫中的作用[J]. 生物技术通报, 2024, 40(6): 5-22.
Li Y P, Qin W, Fu X Q, et al. Transcriptomic analysis reveals the parallel transcriptional regulation of UV-B-induced artemisinin and flavonoid accumulation in Artemisia annua L [J]. Plant Physiol Biochem, 2021, 163: 189-200.
He J, Liu Y Q, Yuan D Y, et al. An R2R3 MYB transcription factor confers brown planthopper resistance by regulating the phenylalanine ammonia-lyase pathway in rice [J]. Proc Natl Acad Sci USA, 2020, 117(1): 271-277.
Hao X L, Pu Z Q, Cao G, et al. Tanshinone and salvianolic acid biosynthesis are regulated by SmMYB98 in Salvia miltiorrhiza hairy roots [J]. J Adv Res, 2020, 23: 1-12.
Liu M Y, Sun W J, Ma Z T, et al. Integrated network analyses identify MYB4R1 neofunctionalization in the UV-B adaptation of Tartary buckwheat [J]. Plant Commun, 2022, 3(6): 100414.
Shamala L F, Zhou H C, Han Z X, et al. UV-B induces distinct transcriptional re-programing in UVR8-signal transduction, flavonoid, and terpenoids pathways in Camellia sinensis [J]. Front Plant Sci, 2020, 11: 234.
李强, 康璠, 薛晴, 等. 神农香菊R2R3-MYB转录因子CiMYB4在镉胁迫中的功能分析[J]. 草业学报, 2024, 33(5): 128-142.
Potter S C, Luciani A, Eddy S R, et al. HMMER web server: 2018 update [J]. Nucleic Acids Res, 2018, 46(W1): W200-W204.
Luo D F, Mei D S, Wei W L, et al. Identification and phylogenetic analysis of the R2R3-MYB subfamily in Brassica napus [J]. Plants, 2023, 12(4): 886.
Chen C J, Wu Y, Li J W, et al. TBtools-II: A “one for all, all for one” bioinformatics platform for biological big-data mining [J]. Mol Plant, 2023, 16(11): 1733-1742.
Wang Y P, Tang H B, Debarry J D, et al. MCScanX: A toolkit for detection and evolutionary analysis of gene synteny and collinearity [J]. Nucleic Acids Res, 2012, 40(7): e49.
Edgar R C. MUSCLE: Multiple sequence alignment with high accuracy and high throughput [J]. Nucleic Acids Res, 2004, 32(5): 1792-1797.
Minh B Q, Schmidt H A, Chernomor O, et al. IQ-TREE 2: New models and efficient methods for phylogenetic inference in the genomic era [J]. Mol Biol Evol, 2020, 37(5): 1530-1534.
Dubos C, Stracke R, Grotewold E, et al. MYB transcription factors in Arabidopsis [J]. Trends Plant Sci, 2010, 15(10): 573-581.
Gu C S, Chen S M, Liu Z L, et al. Reference gene selection for quantitative real-time PCR in Chrysanthemum subjected to biotic and abiotic stress [J]. Mol Biotechnol, 2011, 49(2): 192-197.
Livak K J, Schmittgen T D. Analysis of relative gene expression data using real-time quantitative PCR and the 2(-Delta Delta C(T)) method [J]. Methods, 2001, 25(4): 402-408.
Chang Y K, Shi M M, Wang X, et al. A CRY1-HY5-MYB signaling cascade fine-tunes guard cell reactive oxygen species levels and triggers stomatal opening [J]. Plant Cell, 2025, 37(4): koaf064.
Zhang X Y, He Y Q, Li L Y, et al. Involvement of the R2R3-MYB transcription factor MYB21 and its homologs in regulating flavonol accumulation in Arabidopsis stamen [J]. J Exp Bot, 2021, 72(12): 4319-4332.
Wu Y, Wen J, Xia Y P, et al. Evolution and functional diversification of R2R3-MYB transcription factors in plants [J]. Hortic Res, 2022, 9: uhac058.
Deng Y N, Yang P, Zhang Q L, et al. Genomic insights into the evolution of flavonoid biosynthesis and O-methyltransferase and glucosyltransferase in Chrysanthemum indicum [J]. Cell Rep, 2024, 43(2): 113725.
Song A P, Su J S, Wang H B, et al. Analyses of a chromosome-scale genome assembly reveal the origin and evolution of cultivated Chrysanthemum [J]. Nat Commun, 2023, 14(1): 2021.
Wen X H, Li J Z, Wang L L, et al. The Chrysanthemum lavandulifolium genome and the molecular mechanism underlying diverse Capitulum types [J]. Hortic Res, 2022, 9: uhab022.
Xu X Y, Yang Y H, Zhang T, et al. Systematic analysis of the R2R3-MYB transcription factor gene family in Stevia rebaudiana [J]. Ind Crops Prod, 2024, 210: 118123.
Arce-Rodríguez M L, Martínez O, Ochoa-Alejo N. Genome-wide identification and analysis of the MYB transcription factor gene family in chili pepper (Capsicum spp.) [J]. Int J Mol Sci, 2021, 22(5): 2229.
Bhatia C, Pandey A, Gaddam S R, et al. Low temperature-enhanced flavonol synthesis requires light-associated regulatory components in Arabidopsis thaliana [J]. Plant Cell Physiol, 2018, 59(10): 2099-2112.
Yao P F, Huang Y J, Dong Q X, et al. FtMYB6, a light-induced SG7 R2R3-MYB transcription factor, promotes flavonol biosynthesis in Tartary buckwheat (Fagopyrum tataricum) [J]. J Agric Food Chem, 2020, 68(47): 13685-13696.
Liu S A, Gu X Y, Jiang Y B, et al. UV-B promotes flavonoid biosynthesis in Ginkgo biloba by inducing the GbHY5-GbMYB1-GbFLS module [J]. Hortic Res, 2023, 10(8): uhad118.
Yu W, Zhou X R, Meng J H, et al. WRKY transcription factors modulate the flavonoid pathway of Rhododendron chrysanthum pall. under UV-B stress [J]. Plants, 2025, 14(1): 133.
Zhao Q Z, Dong M Y, Li M F, et al. Light-induced flavonoid biosynthesis in Sinopodophyllum hexandrum with high-altitude adaptation [J]. Plants, 2023, 12(3): 575.
Liu C Y, Long J M, Zhu K J, et al. Characterization of a Citrus R2R3-MYB transcription factor that regulates the flavonol and hydroxycinnamic acid biosynthesis [J]. Sci Rep, 2016, 6: 25352.
Wang Y G, Ye H Y, Wang K T, et al. CcMYB12 positively regulates flavonoid accumulation during fruit development in Carya cathayensis and has a role in abiotic stress responses [J]. Int J Mol Sci, 2022, 23(24): 15618.)
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神农香菊R2R3-MYB基因家族鉴定及其响应UV-B胁迫的表达模式分析
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何叶, 梅洋, 汪苗苗, 杨静月, 张景景, 刘义飞
中草药 | 药材与资源 2026,57(4): 1460-1472
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中草药 |药材与资源 2026 , 57 (4) : 1460 -1472
神农香菊R2R3-MYB基因家族鉴定及其响应UV-B胁迫的表达模式分析
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何叶, 梅洋, 汪苗苗, 杨静月, 张景景, 刘义飞
作者信息
通讯作者:
张景景
作者简介:
何叶: 何叶(2001—),女,硕士研究生,研究方向为中药资源品质评价及开发利用。E-mail:2477067231@qq.com
Identification of R2R3-MYB gene family in Chrysanthemum indicum var. aromaticum and analysis of their expression patterns in response to UV-B stress
HE Ye, MEI Yang, WANG Miaomiao, YANG Jingyue, ZHANG Jingjing, LIU Yifei
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doi: 10.7501/j.issn.0253-2670.2026.04.022
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目的 旨在鉴定神农架民间药用植物神农香菊Chrysanthemum indicum var. aromaticumR2R3-MYB基因家族成员(命名为CiaMYB),解析其基因进化特征及响应UV-B胁迫的表达特征模式,探讨其调控黄酮类化合物合成的分子机制。方法 基于神农香菊全基因组序列,通过生物信息学分析全面鉴定CiaMYB家族成员,系统分析其蛋白理化性质、染色体定位、基因结构、共线性关系及顺式作用元件组成;结合转录组测序与相关性分析,筛选参与黄酮类化合物生物合成的CiaMYB基因;利用实时荧光定量PCR(quantitative reverse transcription polymerase chain reaction,qRT-PCR)验证关键基因表达模式,并解析蛋白功能定位。结果 共鉴定出140个CiaMYB基因,其编码蛋白为206~868个氨基酸组成的亲水性蛋白,不均匀分布于9条染色体中。启动子区富集光响应、脱落酸(abscisic acid,ABA)和茉莉酸甲酯(methyl jasmonate,MeJA)信号通路相关顺式作用元件。UV-B辐照显著诱导CiaMYB基因差异表达和叶片总黄酮积累,其中CiaMYB040CiaMYB066与拟南芥黄酮醇合成关键的S7亚家族高度同源,且表达量与总黄酮含量呈极强正相关,提示其在黄酮合成途径中发挥关键转录调控作用。亚细胞定位证实CiaMYB040和CiaMYB066蛋白定位于细胞核。结论 首次系统揭示神农香菊R2R3-MYB家族成员的进化特征,初步筛选了响应UV-B胁迫参与调控黄酮类物质合成的CiaMYB基因,为解析植物高海拔抗逆适应的分子机制,菊科植物黄酮代谢调控及抗逆遗传改良提供了重要的候选基因资源。
神农香菊  /  R2R3-MYB  /  转录调控  /  UV-B  /  黄酮代谢
Objective To identify members of the R2R3-MYB gene family (named CiaMYB) in Chrysanthemum indicum var. aromaticum, a folk medicinal plant from Shennongjia, analyze their genetic evolutionary characteristics and expression patterns in response to UV-B stress, and explore the molecular mechanisms by which they regulate flavonoid synthesis. Methods Based on the whole-genome sequence of C. indicum var. aromaticum, bioinformatics analyses were performed to comprehensively identify CiaMYB members, and systematic analyses were conducted on their protein physicochemical properties, chromosomal localization, gene structure, collinearity relationships, and composition of cis-acting elements. Combined with transcriptome sequencing and correlation analysis, CiaMYB genes involved in flavonoid biosynthesis were screened. Quantitative reverse transcription polymerase chain reaction (qRT-PCR) was used to verify the expression patterns of key genes, and the functional localization of the proteins was analyzed. Results A total of 140 CiaMYB genes were identified, encoding hydrophilic proteins consisting of 206—868 amino acids, which are unevenly distributed on nine chromosomes. The promoter regions are enriched with cis-acting elements related to light response, abscisic acid (ABA) and methyl jasmonate (MeJA) signaling pathways. UV-B irradiation significantly induces differential expression of CiaMYB genes and accumulation of total flavonoids in leaves. Among them, CiaMYB040 and CiaMYB066 are highly homologous to the S7 subfamily, which is critical for flavonol synthesis in Arabidopsis thaliana, and their expression levels show a strong positive correlation with total flavonoid content, suggesting that they play a key transcriptional regulatory role in the flavonoid synthesis pathway. Subcellular localization confirmed that CiaMYB040 and CiaMYB066 proteins are localized in the nucleus. Conclusion This study is the first to systematically reveal the evolutionary characteristics of R2R3-MYB family members in C. indicum var. aromaticum, and preliminarily screen CiaMYB genes that respond to UV-B stress and are involved in regulating flavonoid synthesis. It provides important candidate gene resources for elucidating the molecular mechanism of plant high-altitude stress adaptation, as well as for flavonoid metabolism regulation and stress-resistant genetic improvement in Asteraceae plants.
Chrysanthemum indicum var. aromaticum L.  /  R2R3-MYB  /  transcriptional regulation  /  UV-B  /  flavonoid metabolism
何叶, 梅洋, 汪苗苗, 杨静月, 张景景, 刘义飞. 神农香菊R2R3-MYB基因家族鉴定及其响应UV-B胁迫的表达模式分析. 中草药, 2026 , 57 (4) : 1460 -1472 . DOI: 10.7501/j.issn.0253-2670.2026.04.022
HE Ye, MEI Yang, WANG Miaomiao, YANG Jingyue, ZHANG Jingjing, LIU Yifei. Identification of R2R3-MYB gene family in Chrysanthemum indicum var. aromaticum and analysis of their expression patterns in response to UV-B stress[J]. Chinese Traditional and Herbal Drugs, 2026 , 57 (4) : 1460 -1472 . DOI: 10.7501/j.issn.0253-2670.2026.04.022

    国家自然科学基金项目 (32270231); 湖北中医药大学重大项目 (2023ZDXM006)

参考文献 引证文献
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袁仁森, 王旭, 张景景, 等. 神农香菊自然居群遗传变异评价研究及核心种质筛选[J]. 世界科学技术—中医药现代化, 2022, 24(4): 1325-1334.
雷霄飞, 杨学领. 神农香菊化学成分及药理作用研究进展[J]. 现代农业科技, 2014(19): 312-313.
Wang X, Zhang J J, Liu Z G, et al. Comparative transcriptome analysis of three chrysanthemums provides insights into flavonoid and terpenoid biosynthesis [J]. J Plant Biol, 2021, 64(5): 389-401.
Chen K L, Liu Y M, Zhang Y, et al. A special aromatic Chrysanthemum breed with high content of thujone [J]. Phcog Mag, 2020, 16(71): 625.
Feng Y, Li B, Hu J, et al. Sustainable use of Chrysanthemum indicum var. aromaticum as value-added green materials in microemulsion hydrogels [J]. ACS Sustainable Chem Eng, 2023, 11(9): 3820-3831.
Shi C, Liu H T. How plants protect themselves from ultraviolet-B radiation stress [J]. Plant Physiol, 2021, 187(3): 1096-1103.
Sztatelman O, Grzyb J, Gabryś H, et al. The effect of UV-B on Arabidopsis leaves depends on light conditions after treatment [J]. BMC Plant Biol, 2015, 15: 281.
Xiong Y, Yuan S, Xiong Y L, et al. Analysis of allohexaploid wheatgrass genome reveals its Y haplome origin in Triticeae and high-altitude adaptation [J]. Nat Commun, 2025, 16(1): 3104.
Ferreyra M L F, Serra P, Casati P. Recent advances on the roles of flavonoids as plant protective molecules after UV and high light exposure [J]. Physiol Plant, 2021, 173(3): 736-749.
Takshak S, Agrawal S B. Defense potential of secondary metabolites in medicinal plants under UV-B stress [J]. J Photochem Photobiol B, 2019, 193: 51-88.
Liang T, Shi C, Peng Y, et al. Brassinosteroid-activated BRI1-EMS-SUPPRESSOR 1 inhibits flavonoid biosynthesis and coordinates growth and UV-B stress responses in plants [J]. Plant Cell, 2020, 32(10): 3224-3239.
胡雅丹, 伍国强, 刘晨, 等. MYB转录因子在调控植物响应逆境胁迫中的作用[J]. 生物技术通报, 2024, 40(6): 5-22.
Li Y P, Qin W, Fu X Q, et al. Transcriptomic analysis reveals the parallel transcriptional regulation of UV-B-induced artemisinin and flavonoid accumulation in Artemisia annua L [J]. Plant Physiol Biochem, 2021, 163: 189-200.
He J, Liu Y Q, Yuan D Y, et al. An R2R3 MYB transcription factor confers brown planthopper resistance by regulating the phenylalanine ammonia-lyase pathway in rice [J]. Proc Natl Acad Sci USA, 2020, 117(1): 271-277.
Hao X L, Pu Z Q, Cao G, et al. Tanshinone and salvianolic acid biosynthesis are regulated by SmMYB98 in Salvia miltiorrhiza hairy roots [J]. J Adv Res, 2020, 23: 1-12.
Liu M Y, Sun W J, Ma Z T, et al. Integrated network analyses identify MYB4R1 neofunctionalization in the UV-B adaptation of Tartary buckwheat [J]. Plant Commun, 2022, 3(6): 100414.
Shamala L F, Zhou H C, Han Z X, et al. UV-B induces distinct transcriptional re-programing in UVR8-signal transduction, flavonoid, and terpenoids pathways in Camellia sinensis [J]. Front Plant Sci, 2020, 11: 234.
李强, 康璠, 薛晴, 等. 神农香菊R2R3-MYB转录因子CiMYB4在镉胁迫中的功能分析[J]. 草业学报, 2024, 33(5): 128-142.
Potter S C, Luciani A, Eddy S R, et al. HMMER web server: 2018 update [J]. Nucleic Acids Res, 2018, 46(W1): W200-W204.
Luo D F, Mei D S, Wei W L, et al. Identification and phylogenetic analysis of the R2R3-MYB subfamily in Brassica napus [J]. Plants, 2023, 12(4): 886.
Chen C J, Wu Y, Li J W, et al. TBtools-II: A “one for all, all for one” bioinformatics platform for biological big-data mining [J]. Mol Plant, 2023, 16(11): 1733-1742.
Wang Y P, Tang H B, Debarry J D, et al. MCScanX: A toolkit for detection and evolutionary analysis of gene synteny and collinearity [J]. Nucleic Acids Res, 2012, 40(7): e49.
Edgar R C. MUSCLE: Multiple sequence alignment with high accuracy and high throughput [J]. Nucleic Acids Res, 2004, 32(5): 1792-1797.
Minh B Q, Schmidt H A, Chernomor O, et al. IQ-TREE 2: New models and efficient methods for phylogenetic inference in the genomic era [J]. Mol Biol Evol, 2020, 37(5): 1530-1534.
Dubos C, Stracke R, Grotewold E, et al. MYB transcription factors in Arabidopsis [J]. Trends Plant Sci, 2010, 15(10): 573-581.
Gu C S, Chen S M, Liu Z L, et al. Reference gene selection for quantitative real-time PCR in Chrysanthemum subjected to biotic and abiotic stress [J]. Mol Biotechnol, 2011, 49(2): 192-197.
Livak K J, Schmittgen T D. Analysis of relative gene expression data using real-time quantitative PCR and the 2(-Delta Delta C(T)) method [J]. Methods, 2001, 25(4): 402-408.
Chang Y K, Shi M M, Wang X, et al. A CRY1-HY5-MYB signaling cascade fine-tunes guard cell reactive oxygen species levels and triggers stomatal opening [J]. Plant Cell, 2025, 37(4): koaf064.
Zhang X Y, He Y Q, Li L Y, et al. Involvement of the R2R3-MYB transcription factor MYB21 and its homologs in regulating flavonol accumulation in Arabidopsis stamen [J]. J Exp Bot, 2021, 72(12): 4319-4332.
Wu Y, Wen J, Xia Y P, et al. Evolution and functional diversification of R2R3-MYB transcription factors in plants [J]. Hortic Res, 2022, 9: uhac058.
Deng Y N, Yang P, Zhang Q L, et al. Genomic insights into the evolution of flavonoid biosynthesis and O-methyltransferase and glucosyltransferase in Chrysanthemum indicum [J]. Cell Rep, 2024, 43(2): 113725.
Song A P, Su J S, Wang H B, et al. Analyses of a chromosome-scale genome assembly reveal the origin and evolution of cultivated Chrysanthemum [J]. Nat Commun, 2023, 14(1): 2021.
Wen X H, Li J Z, Wang L L, et al. The Chrysanthemum lavandulifolium genome and the molecular mechanism underlying diverse Capitulum types [J]. Hortic Res, 2022, 9: uhab022.
Xu X Y, Yang Y H, Zhang T, et al. Systematic analysis of the R2R3-MYB transcription factor gene family in Stevia rebaudiana [J]. Ind Crops Prod, 2024, 210: 118123.
Arce-Rodríguez M L, Martínez O, Ochoa-Alejo N. Genome-wide identification and analysis of the MYB transcription factor gene family in chili pepper (Capsicum spp.) [J]. Int J Mol Sci, 2021, 22(5): 2229.
Bhatia C, Pandey A, Gaddam S R, et al. Low temperature-enhanced flavonol synthesis requires light-associated regulatory components in Arabidopsis thaliana [J]. Plant Cell Physiol, 2018, 59(10): 2099-2112.
Yao P F, Huang Y J, Dong Q X, et al. FtMYB6, a light-induced SG7 R2R3-MYB transcription factor, promotes flavonol biosynthesis in Tartary buckwheat (Fagopyrum tataricum) [J]. J Agric Food Chem, 2020, 68(47): 13685-13696.
Liu S A, Gu X Y, Jiang Y B, et al. UV-B promotes flavonoid biosynthesis in Ginkgo biloba by inducing the GbHY5-GbMYB1-GbFLS module [J]. Hortic Res, 2023, 10(8): uhad118.
Yu W, Zhou X R, Meng J H, et al. WRKY transcription factors modulate the flavonoid pathway of Rhododendron chrysanthum pall. under UV-B stress [J]. Plants, 2025, 14(1): 133.
Zhao Q Z, Dong M Y, Li M F, et al. Light-induced flavonoid biosynthesis in Sinopodophyllum hexandrum with high-altitude adaptation [J]. Plants, 2023, 12(3): 575.
Liu C Y, Long J M, Zhu K J, et al. Characterization of a Citrus R2R3-MYB transcription factor that regulates the flavonol and hydroxycinnamic acid biosynthesis [J]. Sci Rep, 2016, 6: 25352.
Wang Y G, Ye H Y, Wang K T, et al. CcMYB12 positively regulates flavonoid accumulation during fruit development in Carya cathayensis and has a role in abiotic stress responses [J]. Int J Mol Sci, 2022, 23(24): 15618.
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doi: 10.7501/j.issn.0253-2670.2026.04.022
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https://castjournals.cast.org.cn/joweb/zcy/CN/10.7501/j.issn.0253-2670.2026.04.022
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2种不同金属材料的力学参数

Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)

Genus
种数
Number of
species
占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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