Article(id=1308841457230906274, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1308841198224240856, articleNumber=null, orderNo=null, doi=10.16438/j.0513-4870.2026-0092, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1769011200000, receivedDateStr=2026-01-22, revisedDate=1773936000000, revisedDateStr=2026-03-20, acceptedDate=null, acceptedDateStr=null, onlineDate=1789981720502, onlineDateStr=2026-09-21, pubDate=1781193600000, pubDateStr=2026-06-12, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1789981720502, onlineIssueDateStr=2026-09-21, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1789981720502, creator=13701087609, updateTime=1789981720502, updator=13701087609, issue=Issue{id=1308841198224240856, tenantId=1146029695717560320, journalId=1189982191388893191, year='2026', volume='61', issue='6', pageStart='1881', pageEnd='2376', issueExtLink='null', onlineDate='null', pubDate='1781193600000', pubDateStr='2026-06-12', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1789981658750, creator='13701087609', updateTime=1789984187230, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1308851803500933401, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1308841198224240856, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1308851803500933402, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1308841198224240856, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=2347, endPage=2359, ext={EN=ArticleExt(id=1308841457444815779, articleId=1308841457230906274, tenantId=1146029695717560320, journalId=1189982191388893191, language=EN, title=Identification and expression pattern analysis of the MYC gene family in Lonicera japonica, columnId=1190335348761793317, journalTitle=Acta Pharmaceutica Sinica, columnName=Original Articles, runingTitle=null, highlight=null, articleAbstract=

The MYC transcription factor is a core regulator in the jasmonic acid signaling pathway, widely involved in plant growth and development, secondary metabolism, stress response, and hormone signal transduction. Based on the genomic data of Lonicera japonica, this study identified members of the LjMYC gene family through bioinformatics methods, and analyzed their physicochemical properties, evolutionary relationships, conserved domains and gene structures, as well as promoter cis-acting elements. The expression patterns of this gene family in different tissues, different flowering stages, and under methyl jasmonate (MeJA) treatment were also investigated. The results showed that a total of 25 LjMYC genes were identified in Lonicera japonica, distributed across eight chromosomes. Subcellular localization prediction indicated that 22 LjMYCs proteins were localized in the nucleus, LjMYC13 and LjMYC14 in chloroplasts, and LjMYC4 in the cytoplasm. Cis-acting element analysis revealed that the promoter regions of the LjMYCs gene contain elements responsive to light, growth and development, hormones, and stress. RT-qPCR analysis demonstrated that eight genes, including LjMYC8, exhibited tissue-specific expression in floral organs. The expression trends of ten genes, including LjMYC1, were generally consistent during the flowering process (S1-S6): Expression was slightly lower at S2 than at S1, increased at S3, slightly decreased at S4, and remained high at S5 and S6. Exogenous MeJA treatment promoted flowering, whereas delayed flowering treated with sodium diethyldithiocarbamate trihydrate (DIECA). Except for LjMYC20, MeJA treatment induced the expression of nine members, including LjMYC1, while DIECA treatment suppressed their expression. This study systematically reveals the fundamental characteristics of the LjMYC gene family and its expression patterns in different tissues, flowering stages, and in response to jasmonic acid signaling, providing key insights for exploring the functions of this gene family in jasmonic acid signal transduction in Lonicera japonica.

, authors=Hui-ling GONG1, Miao-miao ZHANG1, Shi-cheng LUO1, Hang WANG1, Jia-peng YANG1, Ai-qing CAO1, Ping XU2, 3, Yao CHEN3, Zai-ping FENG1, *, authorsList=Hui-ling GONG, Miao-miao ZHANG, Shi-cheng LUO, Hang WANG, Jia-peng YANG, Ai-qing CAO, Ping XU, Yao CHEN, Zai-ping FENG, authorCompany=null, correspAuthors=Zai-ping FENG, authorNote=null, correspAuthorsNote=null, copyrightStatement=Copyright ©2026 Acta Pharmaceutica Sinica. All rights reserved., 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=1308841460716372918, articleId=1308841457230906274, tenantId=1146029695717560320, journalId=1189982191388893191, language=CN, title=金银花MYC基因家族的鉴定及表达模式分析, columnId=1190335348896011050, journalTitle=药学学报, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=

MYC转录因子是茉莉酸信号转导通路中的核心调控因子, 广泛参与植物生长发育、次生代谢、逆境应答以及激素信号传导等过程。本研究基于金银花(Lonicera japonica Thunb.) 基因组数据, 通过生物信息学方法对LjMYC基因家族进行鉴定, 对其理化性质、进化关系、保守结构域与基因结构、启动子顺式作用元件等方面进行分析, 并对该基因家族在金银花不同组织、不同开花时期以及茉莉酸甲酯(MeJA) 处理下的表达模式进行分析。结果表明, 共鉴定出25个金银花LjMYCs基因, 分布在8条染色体上, 亚细胞定位预测其中22个LjMYCs蛋白定位于细胞核, LjMYC13和LjMYC14定位于叶绿体中, LjMYC4定位于细胞质中。顺式作用元件分析表明, LjMYCs基因启动子区域含有光响应、生长发育、激素和胁迫响应等作用元件。RT-qPCR方法分析表明, LjMYC8等8个基因在花蕊中的表达具有组织特异性; LjMYC1等10个基因在开花进程(S1~S6) 中的表达变化趋势基本一致, S2时期的表达量略低于S1时期, S3时期表达量升高, S4时期略有下降, S5和S6时期持续高表达。外源MeJA处理促进开花, 二乙基二硫代氨基甲酸钠三水合物(DIECA) 处理延迟开花; 除LjMYC20外, MeJA处理可诱导LjMYC1等9个成员的表达, DIECA处理则抑制其表达。本研究结果揭示了LjMYC基因家族的基本特征以及在不同组织、不同开花时期和茉莉酸信号响应中的表达特征, 为探究该基因家族在金银花茉莉酸信号传导中的功能提供了关键依据。

, authors=巩慧玲1, 张苗苗1, 罗世诚1, 王航1, 杨家鹏1, 曹爱青1, 徐苹2, 3, 陈耀3, 冯再平1, *, authorsList=巩慧玲, 张苗苗, 罗世诚, 王航, 杨家鹏, 曹爱青, 徐苹, 陈耀, 冯再平, authorCompany=null, correspAuthors=冯再平, authorNote=

作者贡献:

第一作者巩慧玲负责论文设计、论文指导、数据分析及论文撰写; 张苗苗负责实验、数据分析及论文初稿的撰写; 罗世诚、王航、杨家鹏和曹爱青参与生物信息学、实验材料的准备以及RT-qPCR实验; 通讯作者冯再平指导文章撰写并提出修改意见; 徐苹和陈耀负责实验材料的种植、日常管理和维护。

, correspAuthorsNote=
冯再平,E-mail:
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A: Gene structure; B: Conserved motif; C: Onserved domain. 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CK: Control; MeJA: Methyl jasmonate; DIECA: Sodium diethyl dithiocarbamate trihydrate; S1: Juvenile floral bud; S2: Three-green floral bud; S3: Two-white floral bud; S4: Complete white floral bud; S5: Silver flower; S6: Gold flower. n = 3, $ \stackrel{-}{x} $ ± s. *P < 0.05, **P < 0.01, ***P < 0.001 vs MeJA; △P < 0.05, △△P < 0.01, △△△P < 0.001 vs DIECA. Compared with the control group at the same time period, figureFileSmall=oDOQRBPCU+QnnGTtvTrInw==, figureFileBig=JzbB5nEz7kiWJHdV3GdHCQ==, tableContent=null), ArticleFig(id=1308841468320645127, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308841457230906274, language=EN, label=null, caption=null, figureFileSmall=fqYtBYIhM/PvGPSIQMLRtA==, figureFileBig=U/3vVbe04keXVS2bwuWMmg==, tableContent=null), ArticleFig(id=1308841468379365384, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308841457230906274, language=CN, label=Fig.9, caption=Treatment with MeJA (A) and DIECA (B) affects the duration of flower buds (left: Control, right: Treatment), figureFileSmall=fqYtBYIhM/PvGPSIQMLRtA==, figureFileBig=U/3vVbe04keXVS2bwuWMmg==, tableContent=null), ArticleFig(id=1308841468450668553, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308841457230906274, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
GeneForward primer (5′-3′)Reverse primer (5′-3′)
LjActinCCCTAAAGCCAACAGAGAGAAGCGACCACTAGCATACAGAGAAAG
LjMYC1TGGGAATTCTTCAAACGGACATCTCTTTCTAGGCTTCCGTTC
LjMYC8GGTATCTGTGCTTGATAACACCCTACCTTGCAGGATTCCAAC
LjMYC11ACCCTCGGAGAATTAGTCCCACTACCTTGCAGGATTCCAAC
LjMYC12TGTTTGTATACCGTCTGCAACCCGGAACAGACATCGAACC
LjMYC14CATAGCGTAACTCAACCTCCCGCTGCCACTGCTTTATCCAC
LjMYC16GTTCTGCTCCTTTCTCCCAATTTCCGTCTTTTGCTTCACC
LjMYC20GATGTTTCTTCTCCTCGGATTCCTGTATTGTGTCGTCCA
LjMYC22CTCCAACATCAACAAGTGCAATGACCCGAAAACAAAGTCCC
LjMYC23GAAGAAATCCCCATCGACCACGTTTCGCTGTTACTTTCCTG
LjMYC24CCATGTATTTCTCCTTTCCTCGCATCGGAAAGCCATAAGTGC
LjMYC25CATCAACGAACTGAAGGCCAAGCCACAATTTTCACATCCACT
), ArticleFig(id=1308841468551331850, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308841457230906274, language=CN, label=Tab.1, caption=

Primer information for RT-qPCR

, figureFileSmall=null, figureFileBig=null, tableContent=
GeneForward primer (5′-3′)Reverse primer (5′-3′)
LjActinCCCTAAAGCCAACAGAGAGAAGCGACCACTAGCATACAGAGAAAG
LjMYC1TGGGAATTCTTCAAACGGACATCTCTTTCTAGGCTTCCGTTC
LjMYC8GGTATCTGTGCTTGATAACACCCTACCTTGCAGGATTCCAAC
LjMYC11ACCCTCGGAGAATTAGTCCCACTACCTTGCAGGATTCCAAC
LjMYC12TGTTTGTATACCGTCTGCAACCCGGAACAGACATCGAACC
LjMYC14CATAGCGTAACTCAACCTCCCGCTGCCACTGCTTTATCCAC
LjMYC16GTTCTGCTCCTTTCTCCCAATTTCCGTCTTTTGCTTCACC
LjMYC20GATGTTTCTTCTCCTCGGATTCCTGTATTGTGTCGTCCA
LjMYC22CTCCAACATCAACAAGTGCAATGACCCGAAAACAAAGTCCC
LjMYC23GAAGAAATCCCCATCGACCACGTTTCGCTGTTACTTTCCTG
LjMYC24CCATGTATTTCTCCTTTCCTCGCATCGGAAAGCCATAAGTGC
LjMYC25CATCAACGAACTGAAGGCCAAGCCACAATTTTCACATCCACT
), ArticleFig(id=1308841468643606539, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308841457230906274, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
RenameGene IDAAMw/DaProtein isoelectricInstability indexAliphatic indexGRAVYSubcellular localization
LjMYC1Lj1C61T4.144049 249.677.5954.8575.55-0.521Nuclear
LjMYC2Lj2A158T89.168975 638.116.3140.6882.74-0.287Nuclear
LjMYC3Lj2C185G6.184993 328.336.1839.0781.06-0.390Nuclear
LjMYC4Lj2P239T15.126830 116.565.9247.8583.66-0.212Cytoplasmic
LjMYC5Lj2A1154T64.121023 102.508.9337.3792.76-0.164Nuclear
LjMYC6Lj2C1174T10.19410 567.854.7542.1362.34-0.353Nuclear
LjMYC7Lj3A964T51.140144 966.095.6353.0159.58-0.670Nuclear
LjMYC8Lj3A1054T81.152158 806.977.8843.7188.77-0.444Nuclear
LjMYC9Lj4A63T85.141545 995.256.0954.9386.89-0.219Nuclear
LjMYC10Lj2A158T89.165672 427.784.8542.9574.83-0.391Nuclear
LjMYC11Lj2C185G6.146752 437.917.9449.3291.78-0.496Nuclear
LjMYC12Lj2P239T15.159865 680.056.1151.0171.12-0.607Nuclear
LjMYC13Lj2A1154T64.164672 451.285.6939.8681.76-0.380Chloroplast
LjMYC14Lj2C1174T10.133436 724.394.3458.0373.29-0.602Chloroplast
LjMYC15Lj3A964T51.142046 287.444.3542.468.95-0.286Nuclear
LjMYC16Lj3A1054T81.147453 016.166.7749.2977.7-0.560Nuclear
LjMYC17Lj4A63T85.134938 865.027.6348.4477.91-0.389Nuclear
LjMYC18Lj6C802T7.133738 556.935.9470.6270.62-0.601Nuclear
LjMYC19Lj7A495T60.148253 227.115.7841.8574.61-0.560Nuclear
LjMYC20Lj1A64T38.160668 091.925.3246.0186.7-0.473Nuclear
LjMYC21Lj7C513T12.117920 444.195.5548.2986.09-0.521Nuclear
LjMYC22Lj8A221T74.166272 524.595.3548.3669.83-0.627Nuclear
LjMYC23Lj8A300T31.143348 422.544.5348.2176.72-0.565Nuclear
LjMYC24Lj9A394T44.120323 053.775.0653.3661.53-0.497Nuclear
LjMYC25Lj9E599T1.134638 754.767.0236.9484.22-0.536Nuclear
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Physicochemical properties of LjMYC gene family. AA: Number of amino acid; Mw: Molecular weight/Da; GRAVY: Grand average of hydropathy

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RenameGene IDAAMw/DaProtein isoelectricInstability indexAliphatic indexGRAVYSubcellular localization
LjMYC1Lj1C61T4.144049 249.677.5954.8575.55-0.521Nuclear
LjMYC2Lj2A158T89.168975 638.116.3140.6882.74-0.287Nuclear
LjMYC3Lj2C185G6.184993 328.336.1839.0781.06-0.390Nuclear
LjMYC4Lj2P239T15.126830 116.565.9247.8583.66-0.212Cytoplasmic
LjMYC5Lj2A1154T64.121023 102.508.9337.3792.76-0.164Nuclear
LjMYC6Lj2C1174T10.19410 567.854.7542.1362.34-0.353Nuclear
LjMYC7Lj3A964T51.140144 966.095.6353.0159.58-0.670Nuclear
LjMYC8Lj3A1054T81.152158 806.977.8843.7188.77-0.444Nuclear
LjMYC9Lj4A63T85.141545 995.256.0954.9386.89-0.219Nuclear
LjMYC10Lj2A158T89.165672 427.784.8542.9574.83-0.391Nuclear
LjMYC11Lj2C185G6.146752 437.917.9449.3291.78-0.496Nuclear
LjMYC12Lj2P239T15.159865 680.056.1151.0171.12-0.607Nuclear
LjMYC13Lj2A1154T64.164672 451.285.6939.8681.76-0.380Chloroplast
LjMYC14Lj2C1174T10.133436 724.394.3458.0373.29-0.602Chloroplast
LjMYC15Lj3A964T51.142046 287.444.3542.468.95-0.286Nuclear
LjMYC16Lj3A1054T81.147453 016.166.7749.2977.7-0.560Nuclear
LjMYC17Lj4A63T85.134938 865.027.6348.4477.91-0.389Nuclear
LjMYC18Lj6C802T7.133738 556.935.9470.6270.62-0.601Nuclear
LjMYC19Lj7A495T60.148253 227.115.7841.8574.61-0.560Nuclear
LjMYC20Lj1A64T38.160668 091.925.3246.0186.7-0.473Nuclear
LjMYC21Lj7C513T12.117920 444.195.5548.2986.09-0.521Nuclear
LjMYC22Lj8A221T74.166272 524.595.3548.3669.83-0.627Nuclear
LjMYC23Lj8A300T31.143348 422.544.5348.2176.72-0.565Nuclear
LjMYC24Lj9A394T44.120323 053.775.0653.3661.53-0.497Nuclear
LjMYC25Lj9E599T1.134638 754.767.0236.9484.22-0.536Nuclear
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Gene originGene1&Gene2KaKsKa/Ks
LjMYC gene familyLjMYC2&LjMYC130.229 926 4840.632 651 3950.363 433 141
LjMYC8&LjMYC110.019 013 4290.030 088 6480.631 913 710
LjMYC11&LjMYC200.522 317 5372.303 045 4740.226 794 279
LjMYC11&LjMYC210.523 238 7702.222 296 1180.235 449 617
LjMYC12&LjMYC220.198 532 5881.930 966 4620.102 815 140
LjMYC9&LjMYC250.508 731 5313.014 668 2200.168 752 079
LjMYC20&LjMYC210.329 392 6351.598 003 9680.206 127 545
LjMYC19&LjMYC250.293 770 6121.656 723 6420.177 320 226
), ArticleFig(id=1308841468865904654, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308841457230906274, language=CN, label=Tab.3, caption=

Ka/Ks analysis of collinear gene pairs of LjMYCs gene. The Ka/Ks ratio is an indicator of genes undergoing positive selection and purification selection during evolution, where Ka represents the substitution rate of non synonymous mutations and Ks represents the substitution rate of synonymous mutations

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Gene originGene1&Gene2KaKsKa/Ks
LjMYC gene familyLjMYC2&LjMYC130.229 926 4840.632 651 3950.363 433 141
LjMYC8&LjMYC110.019 013 4290.030 088 6480.631 913 710
LjMYC11&LjMYC200.522 317 5372.303 045 4740.226 794 279
LjMYC11&LjMYC210.523 238 7702.222 296 1180.235 449 617
LjMYC12&LjMYC220.198 532 5881.930 966 4620.102 815 140
LjMYC9&LjMYC250.508 731 5313.014 668 2200.168 752 079
LjMYC20&LjMYC210.329 392 6351.598 003 9680.206 127 545
LjMYC19&LjMYC250.293 770 6121.656 723 6420.177 320 226
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金银花MYC基因家族的鉴定及表达模式分析
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巩慧玲 1 , 张苗苗 1 , 罗世诚 1 , 王航 1 , 杨家鹏 1 , 曹爱青 1 , 徐苹 2, 3 , 陈耀 3 , 冯再平 1, *
药学学报 | 研究论文 2026,61(6): 2347-2359
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药学学报 |研究论文 2026 , 61 (6) : 2347 -2359
金银花MYC基因家族的鉴定及表达模式分析
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巩慧玲1, 张苗苗1, 罗世诚1, 王航1, 杨家鹏1, 曹爱青1, 徐苹2, 3, 陈耀3, 冯再平1, *
作者信息
  • 1兰州理工大学生命科学与工程学院, 甘肃 兰州 730050
  • 2甘肃省定西市通渭县什川镇人民政府, 甘肃 定西 743000
  • 3甘肃省定西市通渭县清凉沅金银花产业扶贫开发有限公司, 甘肃 定西 743000
通讯作者:
冯再平,E-mail:
作者简介:

作者贡献:

第一作者巩慧玲负责论文设计、论文指导、数据分析及论文撰写; 张苗苗负责实验、数据分析及论文初稿的撰写; 罗世诚、王航、杨家鹏和曹爱青参与生物信息学、实验材料的准备以及RT-qPCR实验; 通讯作者冯再平指导文章撰写并提出修改意见; 徐苹和陈耀负责实验材料的种植、日常管理和维护。

Identification and expression pattern analysis of the MYC gene family in Lonicera japonica
Hui-ling GONG1, Miao-miao ZHANG1, Shi-cheng LUO1, Hang WANG1, Jia-peng YANG1, Ai-qing CAO1, Ping XU2, 3, Yao CHEN3, Zai-ping FENG1, *
Affiliations
  • 1School of Biological Science and Technology, Lanzhou University of Technology, Lanzhou 730050, China
  • 2Shichuan Town People's Government, Tongwei County, Dingxi 743000, China
  • 3Tongwei County Qingliangyuan Honeysuckle Poverty Alleviation Industrial Development Co., Ltd., Dingxi 743000, China
出版时间: 2026-06-12 doi: 10.16438/j.0513-4870.2026-0092
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MYC转录因子是茉莉酸信号转导通路中的核心调控因子, 广泛参与植物生长发育、次生代谢、逆境应答以及激素信号传导等过程。本研究基于金银花(Lonicera japonica Thunb.) 基因组数据, 通过生物信息学方法对LjMYC基因家族进行鉴定, 对其理化性质、进化关系、保守结构域与基因结构、启动子顺式作用元件等方面进行分析, 并对该基因家族在金银花不同组织、不同开花时期以及茉莉酸甲酯(MeJA) 处理下的表达模式进行分析。结果表明, 共鉴定出25个金银花LjMYCs基因, 分布在8条染色体上, 亚细胞定位预测其中22个LjMYCs蛋白定位于细胞核, LjMYC13和LjMYC14定位于叶绿体中, LjMYC4定位于细胞质中。顺式作用元件分析表明, LjMYCs基因启动子区域含有光响应、生长发育、激素和胁迫响应等作用元件。RT-qPCR方法分析表明, LjMYC8等8个基因在花蕊中的表达具有组织特异性; LjMYC1等10个基因在开花进程(S1~S6) 中的表达变化趋势基本一致, S2时期的表达量略低于S1时期, S3时期表达量升高, S4时期略有下降, S5和S6时期持续高表达。外源MeJA处理促进开花, 二乙基二硫代氨基甲酸钠三水合物(DIECA) 处理延迟开花; 除LjMYC20外, MeJA处理可诱导LjMYC1等9个成员的表达, DIECA处理则抑制其表达。本研究结果揭示了LjMYC基因家族的基本特征以及在不同组织、不同开花时期和茉莉酸信号响应中的表达特征, 为探究该基因家族在金银花茉莉酸信号传导中的功能提供了关键依据。

金银花  /  MYC转录因子  /  基因家族  /  表达模式  /  茉莉酸信号

The MYC transcription factor is a core regulator in the jasmonic acid signaling pathway, widely involved in plant growth and development, secondary metabolism, stress response, and hormone signal transduction. Based on the genomic data of Lonicera japonica, this study identified members of the LjMYC gene family through bioinformatics methods, and analyzed their physicochemical properties, evolutionary relationships, conserved domains and gene structures, as well as promoter cis-acting elements. The expression patterns of this gene family in different tissues, different flowering stages, and under methyl jasmonate (MeJA) treatment were also investigated. The results showed that a total of 25 LjMYC genes were identified in Lonicera japonica, distributed across eight chromosomes. Subcellular localization prediction indicated that 22 LjMYCs proteins were localized in the nucleus, LjMYC13 and LjMYC14 in chloroplasts, and LjMYC4 in the cytoplasm. Cis-acting element analysis revealed that the promoter regions of the LjMYCs gene contain elements responsive to light, growth and development, hormones, and stress. RT-qPCR analysis demonstrated that eight genes, including LjMYC8, exhibited tissue-specific expression in floral organs. The expression trends of ten genes, including LjMYC1, were generally consistent during the flowering process (S1-S6): Expression was slightly lower at S2 than at S1, increased at S3, slightly decreased at S4, and remained high at S5 and S6. Exogenous MeJA treatment promoted flowering, whereas delayed flowering treated with sodium diethyldithiocarbamate trihydrate (DIECA). Except for LjMYC20, MeJA treatment induced the expression of nine members, including LjMYC1, while DIECA treatment suppressed their expression. This study systematically reveals the fundamental characteristics of the LjMYC gene family and its expression patterns in different tissues, flowering stages, and in response to jasmonic acid signaling, providing key insights for exploring the functions of this gene family in jasmonic acid signal transduction in Lonicera japonica.

Lonicera japonica  /  MYC transcription factor  /  gene family  /  expression pattern  /  jasmonic acid signaling
巩慧玲, 张苗苗, 罗世诚, 王航, 杨家鹏, 曹爱青, 徐苹, 陈耀, 冯再平. 金银花MYC基因家族的鉴定及表达模式分析. 药学学报, 2026 , 61 (6) : 2347 -2359 . DOI: 10.16438/j.0513-4870.2026-0092
Hui-ling GONG, Miao-miao ZHANG, Shi-cheng LUO, Hang WANG, Jia-peng YANG, Ai-qing CAO, Ping XU, Yao CHEN, Zai-ping FENG. Identification and expression pattern analysis of the MYC gene family in Lonicera japonica[J]. Acta Pharmaceutica Sinica, 2026 , 61 (6) : 2347 -2359 . DOI: 10.16438/j.0513-4870.2026-0092
金银花是忍冬科忍冬属植物忍冬(Lonicera japonica Thunb.) 的干燥花蕾或初开花朵, 作为我国常用的大宗中药材, 具有抗菌、抗病毒、抗肿瘤等多种药理活性[1]。其药用成分主要包括黄酮类、有机酸类、挥发油及环烯醚萜类化合物, 具有重要的药用价值, 临床应用广泛[2]。金银花花蕾的发育过程分为米蕾期(S1)、三青期(S2)、二白期(S3)、大白期(S4)、银花期(S5) 和金花期(S6) 六个时期, 其中三青期至大白期花蕾的药用效果尤为显著[3]。传统金银花品种普遍存在同一植株花蕾发育不同步的问题, 三青期至大白期持续时间短, 这一特性导致采摘时间紧迫, 短期内劳动力需求高度集中, 因而出现因采摘不及时而造成浪费的问题, 严重制约了金银花产业的可持续发展。金银花品种“北花一号”花蕾期长, 大白期持续时间可达10~14天[4], 适宜集中采收, 因此常作为山东、河南和甘肃等地的主栽品种。
MYC转录因子(myelocytomatosis transcription factors) 是碱性螺旋-环-螺旋(basic helix-loop-helix, bHLH) 转录因子家族中一个重要的亚族[5], 除具有bHLH基因家族典型的保守结构域外, 其成员还包含特有的bHLH-MYC-N结构域[6]。MYC转录因子作为茉莉酸(jasmonic acid, JA) 信号转导通路中的核心调控因子, 茉莉酸信号可通过解除茉莉酸ZIM结构域蛋白(jasmonate ZIM domain-containing protein, JAZ) 的抑制来激活MYC的转录活性, 另外被激活的MYC又能直接影响茉莉酸生物合成途径中多个关键基因的表达, 从而促进内源茉莉酸的积累[7]。研究表明, 在拟南芥(Arabidopsis thaliana) 中AtMYC2、AtMYC3、AtMYC4的三突变体在茉莉酸信号响应缺陷的同时, 也表现出开花时间改变以及花器官发育异常的表型[8]; 在水稻(Oryza sativa) 中OsMYC2通过激活下游基因OsMADS1的表达调控小穗发育, 同时促进茉莉酸介导的叶片衰老和调节昼夜开花时间[9,10]; 玫瑰(Rosa hybrida) RhMYC2被茉莉酸诱导后, 直接激活细胞分裂素合成基因RhLOG3并抑制降解基因RhCKX6, 延长了花瓣细胞的增殖期和开花进程[11]。因此MYC转录因子在植物的花器官发育和激素信号传导等生命过程中发挥着核心作用。
茉莉酸类植物激素(jasmonic acid hormones, JAs) 的生物合成与信号传导参与调控植物花器官的发育。内源茉莉酸含量不足会直接影响花器官的正常形成, 常表现为花药发育异常、雄性不育、花瓣发育受阻以及开花延迟等现象[12]。例如, 百合品种“Doubleen”中茉莉酸含量较低导致花器官发育严重缺陷[13]; 拟南芥茉莉酸合成缺陷突变体表现出雄蕊发育异常、花药不开裂、花粉败育以及开花时间改变等表型[14]。茉莉酸甲酯(methyl jasmonate, MeJA) 能够有效模拟内源茉莉酸信号的激活, 可用来研究茉莉酸信号通路的增强对植物生长发育过程的影响。例如外源MeJA对高粱、小麦等禾本科植物颖花的开放均表现出不同程度的诱导作用[15,16], 对谷子雄性不育系颖花的开放也具有较强的诱导效应[17]。此外, 外源MeJA在花的发育中起到调节花丝伸长、雄蕊发育、花药的发育和开裂等作用, 能够恢复Jas合成突变体表现出的花药延迟开裂、花粉粒不育及花延迟开放等缺陷, 调控花期[18]。另外, 二乙基二硫代氨基甲酸钠三水合物(sodium diethyl dithiocarbamate trihydrate, DIECA) 可通过抑制亚麻酸转化为13S-氢过氧亚麻酸这一关键步骤, 阻断茉莉酸前体12-氧代植物二烯酸(12-oxophytodienoic acid, OPDA) 的合成, 有效降低植物内源茉莉酸水平, 从而产生与茉莉酸缺失相关的表型[19]。DIECA一般作为茉莉酸生物合成抑制剂使用, 例如水曲柳(Fraxinus mandshuira) 幼苗经DIECA处理可抑制茉莉酸合成, 影响下游转录调控网络[20]。因此, 茉莉酸信号在调控植物生殖发育, 尤其是花器官形成与开花进程中扮演着重要的角色, 是植物生殖发育的关键调节因子。
目前, 有关金银花的研究多集中于资源评价、化学成分及药效学方面, 在花器官发育调控的分子机制方面的研究较薄弱。随着金银花基因组测序的完成[21], 利用生物信息学手段鉴定并解析LjMYC基因家族的功能, 将为阐明金银花MYC转录因子在茉莉酸信号通路中的功能奠定基础, 对提升金银花生产效率与产业效益具有重要意义。
植物材料  金银花品种“北花一号”种植于甘肃省定西市通渭县李店金银花基地。采摘金银花的叶片、花瓣和花蕊等组织, 以及六个开花时期(S1~S6) 的花蕾。以上材料采摘后立即放入液氮中速冻, 保存于-80 ℃冰箱中, 每组材料3次生物学重复。
试剂  MeJA, 95% (生产批号No.3550407001, 北京索莱宝科技有限公司); DIECA (生产批号JS267982, 上海源叶生物科技有限公司); 总RNA提取试剂盒FastPure Plant Total RNA Isolation Kit (Vazyme, 生产批号7E0212C5); 反转录试剂盒PrimeScriptTM RT Master Mix (Perfect Real Time, 生产批号RR092S); TB Green® Premix Ex TaqTM Ⅱ试剂盒(Tli RNaseH Plus, 生产批号CN830S)。
仪器  实时荧光定量PCR仪(MA-6000, 苏州雅睿生物技术有限公司); 超微量分光光度计(NanoDrop2000, 九方沃德科技有限公司); 高速台式离心机(H/T16MM, 湖南赫西仪器装备有限公司); PCR仪(Genesy 96T, 西安天隆科技有限公司)。
LjMYC基因家族成员鉴定  从National Genomics Data Center下载金银花基因组和gff注释文件(GWHAAZE00000000)[21]。从拟南芥信息网站Tair (https://www.arabidopsis.org/) 中下载拟南芥AtMYCs基因氨基酸序列, 并进行了BLASTp比对。从Pfam蛋白家族数据库(http://pfam.xfam.org) 下载了对应于MYC结构域的隐马尔可夫模型文件(PF14215和PF00010)。使用HMMER 3.0 (http://hmmer.org/) 从金银花基因组数据库中搜索MYC基因, 查询所有包含MYC结构域的基因, 然后使用Pfam和SMART程序来确认MYC的结构域。
LjMYC基因家族成员理化性质分析  使用ProtParam工具(https://web.expasy.org/protparam/) 预测金银花LjMYC蛋白的理化特征, 包括氨基酸序列长度(amino acid sequence length, AA)、理论等电点(theoretical isoelectric point, pI)、分子质量(protein molecular weight, Mw)、不稳定指数(instability index)、脂肪族指数(aliphatic index) 和亲水性平均值(grand average of hydropathicity, GRAVY) 指数等指标。使用Wolf Psort在线网站(https://wolfpsort.hgc.jp/) 进行金银花LjMYC蛋白亚细胞定位预测, 选择最靠前的预测位置作为结果。
多序列比对与系统发育树构建  选取金银花、拟南芥和水稻的MYC的氨基酸序列进行系统发育关系分析。使用Clustal进行多序列比对, 比对完成后使用MEGA 11软件中Neighbor-Joining (NJ) 法构建系统发育树, 参数Bootstrap值设为1 000, 之后使用iTOL (https://itol.embl.de/) 进行系统进化树可视化及美化。
LjMYC家族成员基因结构和蛋白质的结构分析  使用在线MEME工具(https://meme-suite.org/meme/tools/meme) 分析金银花LjMYC成员的基序结构, 其中基序的最大数量设置为10, 基序的最小宽度设置为6, 基序的最大宽度设置为50。从金银花的基因组注释文件中提取金银花MYC结构, 包括CDS和UTR。最后, 使用TBtools v2.138进行可视化分析。
染色体定位及共线性分析  BLASTp程序用于鉴定金银花中的同源MYC基因, 使用MCScanX默认参数分析金银花LjMYC基因之间的共线性关系。使用TBtools对产生的LjMYC共线性基因对进行可视化。
为了进行共线性分析, 从Phytozome v13网站(https://phytozome-next.jgi.doe.gov/) 下载拟南芥和水稻的基因组序列和注释文件。利用MCScanX程序分析金银花与拟南芥、水稻之间的共线性。
LjMYC基因家族启动子顺式调节元件(CREs) 的鉴定  每个金银花LjMYC基因起始密码子(ATG) 上游的2 000 bp区域被认为是启动子序列。使用TBtools提取启动子序列并用plant care (https://bioinformatics.PSB.ugent.be/webtools/plantcare/html/) 分析顺式调控元件。使用TBtools中Basic Biosequence View和HeatMap模块对与胁迫反应、植物生长和发育、植物激素反应和光响应相关的顺式元件进行可视化和总结。
MeJA和DIECA处理金银花的花蕾  选择长势一致的金银花植株, 傍晚喷施400 μmol·L-1的MeJA[22], 24 h后采摘六个开花时期的花蕾; 另选长势一致的金银花植株, 每天傍晚喷施900 μmol·L-1的DIECA, 5天后采摘DIECA处理的六个开花时期的花蕾; 以蒸馏水处理作为对照组, 每个处理重复3次。
RNA提取、cDNA合成及qRT-PCR检测  总RNA提取用试剂盒FastPure Plant Total RNA Isolation Kit (Vazyme), 使用超微量分光光度计检测RNA吸光值。使用反转录试剂盒PrimeScriptTM RT Master Mix (Perfect Real Time) 合成第一链cDNA, 于-20 ℃冰箱中保存备用, 具体操作方法参照说明书。为验证LjMYCs基因在金银花中的表达模式, 选择10个LjMYCs基因。这10个基因特异性引物的设计使用Oligo7软件, 引物由上海生工生物有限公司合成(表1), 采用qRT-PCR方法检测基因的相对表达量。使用金银花Actin作为内参, 每个样品进行3次生物性重复和3次技术性重复, 相对表达量的计算采用2-∆∆CT方法。
数据分析和处理  利用SPSS 27.0.1单因素方差分析进行显著性分析, P < 0.05为差异显著。使用GraphPad Prism 9.5绘制柱状图。
在金银花基因组中鉴定到25个LjMYCs基因, 按照染色体的位置信息分别命名为LjMYC1~LjMYC25。LjMYC蛋白质理化性质分析结果表明(表2), LjMYC蛋白平均氨基酸数为443; 相对分子质量在10 567.85~93 328.33 Da之间; 蛋白质的等电点在4.34~8.93之间; 蛋白质不稳定系数在36.94~70.62之间; 脂溶指数在59.58~92.76之间; 总平均亲水性均为负值, 说明LjMYC蛋白均为亲水性蛋白, 具有较高热稳定性。亚细胞定位预测22个LjMYCs蛋白定位于细胞核, LjMYC13和LjMYC14定位于叶绿体中, LjMYC4定位于细胞质中。
为明确LjMYCs基因的进化关系, 选取拟南芥和水稻MYC基因成员构建系统发育进化树。金银花、拟南芥与水稻系统聚类分析显示(图1), 这些植物的49个MYC基因根据聚类分成7组(Group 1~7)。Group 1包含2个金银花LjMYCs基因、2个拟南芥AtMYCs基因; Group 2包含4个金银花LjMYCs基因; Group 3包含4个金银花LjMYCs基因、2个拟南芥AtMYCs基因和2个水稻OsMYCs基因; Group 4包含5个金银花LjMYCs基因、4个拟南芥AtMYCs基因和3个水稻OsMYCs基因; Group 5包含2个金银花LjMYCs基因、3个拟南芥AtMYCs基因和2个水稻OsMYCs基因; Group 6包含4个金银花LjMYCs基因; Group 7包含4个金银花LjMYCs基因、5个拟南芥AtMYCs基因和1个水稻OsMYC基因。金银花LjMYCs基因与拟南芥AtMYCs基因聚类更近, 表明金银花LjMYCs基因与拟南芥AtMYCs基因进化关系较近, 与水稻的进化关系较远。
为了进一步分析LjMYC蛋白成员的结构, 对25个LjMYCs蛋白进行了基因结构、保守基序和保守结构域分析。基因结构分析表明(图2A), 该家族成员组1、组2和组4中有3~7个内含子, 组3中有8~11个内含子, 组5、组6和组7中有1~3个内含子。处于同亚族内的基因在结构和保守基序组成上呈现高度一致性, 暗示同亚族的基因在金银花特定生理活动中可能具有相似功能。设置10个不同的Motif基序, 对金银花LjMYC蛋白保守基序进行分析(图2B)。25个LjMYCs蛋白中Motif 5和Motif 8分布在18个LjMYCs蛋白中, Motif 7和Motif 9分布在6个LjMYCs蛋白中, Motif 1分布在16个LjMYCs蛋白中, Motif 2分布在22个LjMYCs蛋白中, Motif 3分布在11个LjMYCs蛋白中, Motif 4分布在17个LjMYCs蛋白中, Motif 5分布在18个LjMYCs蛋白中, Motif 6分布在7个LjMYCs蛋白中, Motif 10分布在9个LjMYCs蛋白中, 它们构成了LjMYC蛋白结构域。LjMYCs的保守序列Motif 2、Motif 4、Motif 5、Motif 8均存在于组2、组3和组7中, 表明这三组基因具有更多的同源性, Motif 9是组3和组4特有的保守基序。
通过预测金银花LjMYC家族蛋白可能含有的功能结构域(图2C), 发现所有的LjMYC蛋白均含有bHLH_MYC_N结构域, 大部分LjMYC蛋白含有HLH结构域, 主要集中在第5组和第7组中。一些LjMYC蛋白还含有其他的结构域, 例如LjMYC5除了含有bHLH_MYC_N和HLH结构域外, 还含有Polyoma_ lg_T_C superfamily结构域, LjMYC4还含有DUF4923 superfamily结构域, 这些结构的差异可能是导致LjMYC家族蛋白发生功能分化的原因之一。
对金银花LjMYCs基因成员的转录起始位置上游2 000 bp的各类顺式作用元件序列分析, 在LjMYCs基因启动子区共鉴定出38个不同的顺式作用元件(图3)。这些元件根据其功能响应特性, 可划分为四大类别: 第一类是光响应元件, 包括Box4、GT1-motif、TCT-motif等; 第二类是植物激素响应元件, 包括ABRE、ERE、TCA-element、CGTCA-motif等, 这些顺式作用元件与茉莉酸甲酯(MeJA)、脱落酸(abscisicacid, ABA)、生长素、赤霉素(gibberellicacid, GA) 响应有关; 第三类是与植物生长发育过程相关的元件, 包括O2-site、CAT-box、AAGAA-motif等; 第四类是非生物与生物胁迫响应相关的元件, 包括抗氧化反应元件(ARE)、创伤应答元件(WUN-motif)、应激响应元件(STRE) 等。
25个LjMYCs基因分布在金银花8条染色体上(图4)。第1条和第2条染色体上各有7个LjMYCs基因; 第9条染色体上有3个LjMYCs基因; 第3条、第7条和第8条染色体上各有2个LjMYCs基因; 第4条和第6条染色体上各有一个LjMYC基因; 第5条染色体上没有LjMYC基因。对金银花LjMYCs的基因组内进行共线性分析, 发现金银花基因组内有8对共线性LjMYC基因(图5), 包括LjMYC2/LjMYC13、LjMYC8/LjMYC11、LjMYC11/LjMYC20、LjMYC11/LjMYC21、LjMYC12/LjMYC22、LjMYC9/LjMYC25、LjMYC20/LjMYC21、LjMYC19/LjMYC25。
对LjMYCs基因组内共线性对进行Ka/Ks分析(表3)。金银花8对共线性基因的Ka均小于1, 说明金银花LjMYCs基因在非同义突变的替代率上经历了较强的保守性约束, 表明这些基因在演化过程中可能扮演着重要的功能角色, 不容易发生非同义突变导致氨基酸序列的变化; 除了LjMYC2/LjMYC13和LjMYC8/LjMYC11, 其余6对金银花共线性基因的Ks大于1, 表示同义突变的替代率相对较高, 这可能表明金银花LjMYCs基因在演化中经历了一定的变化和适应; 8对金银花共线性基因的Ka/Ks比值均小于1, 表明基因在进化中受到较强的保守性约束。
此外, 为进一步了解MYC基因的系统进化关系, 构建了金银花基因组与拟南芥、水稻基因组的共线性图谱(图6)。结果显示, 金银花与拟南芥有29对共线性MYC基因, 金银花与水稻有2对共线性MYC基因。这表明相比于水稻, 金银花MYC基因与拟南芥之间的同源关系更近。
外源MeJA可通过AtMYC2/3/4信号通路延迟拟南芥开花, 并抑制FT基因的表达[23]。本研究在系统发育分析的基础上, 在金银花中选出拟南芥AtMYC2/3/4的10个同源基因(LjMYC1、LjMYC8、LjMYC12、LjMYC14、LjMYC16、LjMYC20、LjMYC22、LjMYC23、LjMYC24、LjMYC25), 对其在金银花不同组织中的表达模式进行分析(图7)。结果表明, 除LjMYC1和LjMYC24外, 其余8个成员在花蕊中表达量最高, 花瓣次之, 叶片最低, 表明LjMYCs基因在花蕊中的表达具有组织特异性。其中LjMYC14、LjMYC20与LjMYC22在花蕊中的表达量分别为叶片的3.1倍、3.9倍和4.0倍; LjMYC22与LjMYC24在花瓣中的表达量分别为叶片的2.1倍和2.9倍; LjMYC12和LjMYC16在花瓣与叶片间的表达量无显著差异。
为了探究LjMYCs基因在开花进程中的表达动态, 本研究选取6个关键开花时期(S1~S6) 的花蕾, 通过RT-qPCR方法对该家族10个成员的表达模式进行分析(图8)。结果表明, LjMYC1等10个基因在开花进程(S1~S6) 中的表达变化趋势基本一致, S2时期的表达量略低于S1时期, S3时期表达量升高, 是S1时期表达量的1.8~3.5倍, S4时期略有下降, S5和S6时期表达量均升高, S6时期表达量是S1时期的2.1~5.4倍。
为了探索金银花LjMYCs基因在茉莉酸信号通路中的作用, 利用MeJA和DIECA对金银花的花蕾进行处理, 结果显示外源MeJA处理可促进开花, DIECA处理则使开花延迟(图9)。通过RT-qPCR方法对LjMYCs基因10个成员在MeJA和DIECA处理后六个开花时期(S1~S6) 的表达模式进行分析(图8)。结果表明, 除LjMYC20外, MeJA处理可诱导LjMYC1等9个成员的表达, 而DIECA处理可抑制其表达。LjMYC1等9个成员在MeJA处理下的表达量是对照组的2.1~18.6倍。在S1时期, LjMYC22的表达量是对照组的8.7倍; 在S2时期, LjMYC25的表达量是对照组的18.6倍; 在S4时期, LjMYC12表达量是对照组的8.3倍; 在S5时期, LjMYC8表达量是对照组的4.8倍。值得注意的是, LjMYC20在MeJA和DIECA处理下的表达始终低于对照组, 且在后续4个发育时期(S3~S6) 均呈现显著抑制。
MYC转录因子作为茉莉酸信号途径的核心调控因子, 在植物激素的信号传导、逆境响应、次生代谢产物合成及生长发育方面发挥着重要功能[24]。随着全基因组测序技术的发展, MYC转录因子在多种植物中得到鉴定, 并对其功能进行了研究。在高粱(Sorghum bicolor L.) 全基因组中共鉴定出28个MYC基因[25], 在广藿香(Pogostemon cablin Benth.) 全基因组中共鉴定出21个MYC基因[26], 在艾(Artemisia argyi Lévl. et Vant) 全基因组中共鉴定出24个MYC基因[27]。本研究在金银花中共鉴定出25个LjMYCs基因, 分布在8条染色体上, 亚细胞定位预测22个LjMYCs蛋白定位于细胞核, 这说明其核心功能是在细胞核内作为转录调控因子负责激活或抑制下游靶基因的表达; LjMYC13和LjMYC14定位于叶绿体, 这一非典型定位说明其可能参与叶绿体相关的特殊调控机制; LjMYC4定位于细胞质中, 说明该转录因子的活性可能受到上游信号通路的控制, 值得后续通过实验进一步验证。LjMYCs基因启动子区富含茉莉酸信号通路相关的响应元件, 表明该家族基因在茉莉酸信号转导中可能发挥核心作用。系统进化和共线性分析表明, 相比于水稻, 金银花LjMYC基因与拟南芥AtMYC基因的同源关系更近, 说明二者MYC基因的作用可能具有相似性。
MYC转录因子已被证实广泛参与植物的花发育调控过程, 其功能涉及花器官的发育与形态建成。拟南芥AtMYC2在花器官中激活下游转录因子MYB21和MYB24的表达, 从而进一步直接调控花瓣细胞伸长相关基因的转录, 促进花瓣的正常生长和发育[28]。本研究也表明LjMYCs基因在花蕊中的表达具有组织特异性, 这一表达模式说明其功能可能与花器官的发育有关。MYC基因的表达量常随着开花进程而呈现动态变化, 这一现象暗示其可能并非简单的静态调控因子, 而是作为开花时序网络中的一个动态响应与执行元件[29,30]。比如在拟南芥中AtMYC基因的表达受到发育信号的调控, 并在特定发育阶段被诱导[31]。本研究中LjMYCs基因在6个开花时期(S1~S6) 的表达动态可能与品种“北花一号”开花进程中的阶段特征有关: 在花发育前期(S1~S2) 整体表达量偏低, 在S4时期表达量下调可能延缓了S4向S5时期的过渡, 导致大白期延长与开花延迟。
除了开花时序调控, MYC转录因子还参与花色形成等开花后过程。例如, 拟南芥AtMYC2能直接结合并激活花青素生物合成途径中的关键基因的启动子, 促进花青素的积累, 过表达AtMYC2能使植株呈现更深的紫色[32]。本研究中LjMYCs基因表达量在S5至S6时期持续上升并伴随着花色由白转黄, 说明LjMYCs基因不仅参与开花时序调控, 还可能影响花色合成等开花后发育过程, 其高表达可能具有促进花色加深的作用。值得注意的是, 花发育这一复杂进程受到多层次、多通路的协同调控。例如, 在金银花中, 热休克蛋白Hsp70 (heat shock protein 70) 及其相关miRNA的表达在不同开花时期呈现动态变化, 说明非生物胁迫响应通路可能被整合至花期发育过程中[33]。同时, 参与生物胁迫响应的MLO (mildew resistance locus O) 基因在花器官中的表达也具有组织特异性, 提示生殖发育阶段可能需要特别的抗病保护机制[34]。这些发现共同表明, 开花过程是植物协调生长发育与多重环境胁迫响应的结果。在这一背景下, LjMYCs基因介导的茉莉酸信号通路很可能并非独立运作。在近缘物种灰毡毛忍冬中, 花器官发育关键调控因子AGL12已被鉴定并发现其拥有特定的互作蛋白网络[35]。LjMYCs所调控的次生代谢与防御反应, 可能与AGL12等花发育因子主导的器官建成过程存在交互, 未来研究值得深入探索。此外, 这一调控网络的复杂性在茉莉酸通路内部已有体现。最新研究发现, 金银花中的转录因子LjWRKY50同样响应茉莉酸信号, 并特异性地调控花蕾的持续期[36]。这与本研究中LjMYCs主要调控开花时序转变及花色合成的功能既存在联系又各有分工, 共同揭示了茉莉酸信号通过激活不同的下游转录因子, 对花发育的不同阶段进行调控。此外, 有研究发现MYC基因的表达模式与黄酮类物质的积累呈现正相关, 表明该家族基因可能作为茉莉酸信号通路的关键组分, 参与调控此阶段次生代谢转化。尽管受实验条件所限, 本研究未能对筛选得到的候选LjMYC蛋白进行亚细胞定位实验, 但预测分析显示LjMYC2定位于细胞核。作为茉莉酸信号通路的核心枢纽, LjMYC2在核内可通过结合下游靶基因启动子上的G-box元件, 直接激活黄酮合成关键酶基因的表达, 从而驱动木犀草苷等物质的生物合成[37]。因此, LjMYCs基因很可能同时参与金银花的花发育进程以及黄酮类活性成分的生物合成调控。
MYC转录因子作为茉莉酸信号通路的核心响应元件, 其表达受外源MeJA的正向调控。比如拟南芥幼苗进行外源MeJA处理, 基因AtMYC2、AtMYC3、AtMYC4的表达量在短时间内被快速诱导[38]; 使用DIECA处理拟南芥叶片, 可有效降低内源茉莉酸水平, 从而导致AtMYC2等基因的表达量显著下调[39]; 艾(Artemisia argyi) 在MeJA诱导下, AarMYC3、AarMYC4、AarMYC5等6个基因呈现“先升高后降低”的表达模式, 而AarMYC12却持续上调表达[27]。本研究中LjMYC1等9个成员在MeJA与DIECA处理下总体上呈现一致的表达调控趋势, 即MeJA促进表达而DIECA抑制表达。
综上所述, 本研究系统揭示了金银花LjMYCs基因的基本特征, 以及其在不同组织、不同开花时期和MeJA处理下的表达模式, 为理解茉莉酸信号与LjMYCs基因之间的调控作用提供了依据。

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2026年第61卷第6期
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doi: 10.16438/j.0513-4870.2026-0092
  • 接收时间:2026-01-22
  • 首发时间:2026-09-21
  • 出版时间:2026-06-12
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  • 收稿日期:2026-01-22
  • 修回日期:2026-03-20
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    1兰州理工大学生命科学与工程学院, 甘肃 兰州 730050
    2甘肃省定西市通渭县什川镇人民政府, 甘肃 定西 743000
    3甘肃省定西市通渭县清凉沅金银花产业扶贫开发有限公司, 甘肃 定西 743000

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2种不同金属材料的力学参数
科
Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)
属
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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