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Article(id=1297571007133406169, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1297570992835023717, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20260164, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1772208000000, receivedDateStr=2026-02-28, revisedDate=null, revisedDateStr=null, acceptedDate=1773676800000, acceptedDateStr=2026-03-17, onlineDate=1787294635803, onlineDateStr=2026-08-21, pubDate=1785772800000, pubDateStr=2026-08-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1787294635803, onlineIssueDateStr=2026-08-21, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1787294635803, creator=13701087609, updateTime=1787294635803, updator=13701087609, issue=Issue{id=1297570992835023717, tenantId=1146029695717560320, journalId=1192105938417971205, year='2026', volume='66', issue='8', pageStart='3681', pageEnd='4288', issueExtLink='null', onlineDate='null', pubDate='1785772800000', pubDateStr='2026-08-04', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1787294632395, creator='13701087609', updateTime=1787294931551, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1297572247670124783, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1297570992835023717, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1297572247670124784, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1297570992835023717, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=3889, endPage=3901, ext={EN=ArticleExt(id=1297571007296984026, articleId=1297571007133406169, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Molecular mechanism of arginine methyltransferase BcHMT2 in regulating the growth, development, and pathogenicity of Botrytis cinerea, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

[Objective] To elucidate the role of arginine methyltransferase in the growth, development, and pathogenicity of Botrytis cinerea. [Methods] Bioinformatics approaches were used to perform sequence alignment, phylogenetic analysis, and conserved domain prediction of the arginine methyltransferase HMT2 from eight fungal species, including B. cinerea, Saccharomyces cerevisiae, Fusarium graminearum, and Magnaporthe oryzae. The gene encoding this enzyme in B. cinerea was identified as BcHMT2. The BcHMT2-deleted mutant was constructed via homologous recombination, followed by phenotypic characterization and pathogenicity assays. [Results] The BcHMT2-deleted mutant exhibited a significantly reduced growth rate, significantly weakend cell wall-degrading enzyme activity, and decreased sclerotial production and conidiation. Meanwhile, the hyphal cells became smaller and the conidial morphology was abnormal. In addition, the mutant displayed markedly attenuated pathogenicity on tomato fruits and tobacco leaves. [Conclusion] This study reveals the regulatory role of BcHMT2 in the growth, development, and pathogenicity of B. cinerea, providing novel insights into the research on the prevention and control of gray mold.

, authors=Yubo LI1, 2, Siqi DENG1, Guang’en DONG1, Jinping ZANG1, Helong SI1, Jingao DONG1, 2, Jihong XING1, 2, Kang ZHANG1, 2, authorsList=Yubo LI, Siqi DENG, Guang’en DONG, Jinping ZANG, Helong SI, Jingao DONG, Jihong XING, Kang ZHANG, authorCompany=null, correspAuthors=Jihong XING, Kang ZHANG, authorNote=

These authors contributed equally to this work.

, correspAuthorsNote=
E-mail: XING Jihong, ;
ZHANG Kang,
, 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=1297571010480460777, articleId=1297571007133406169, tenantId=1146029695717560320, journalId=1192105938417971205, language=CN, title=精氨酸甲基转移酶BcHMT2调控灰葡萄孢生长发育及致病力的分子机制, columnId=1192149544164012138, journalTitle=微生物学报, columnName=研究报告, runingTitle=null, highlight=null, articleAbstract=

【目的】 阐明精氨酸甲基转移酶在灰葡萄孢(Botrytis cinerea)生长发育及致病过程中的作用。 【方法】 利用生物信息学手段对灰葡萄孢、酿酒酵母(Saccharomyces cerevisiae)、禾谷镰孢(Fusarium graminearum)、稻瘟病菌(Magnaporthe oryzae)等8种真菌的精氨酸甲基转移酶HMT2进行序列比对、系统发育分析与保守结构域预测，明确了灰葡萄孢中编码该酶的基因为BcHMT2。通过同源重组技术构建BcHMT2基因敲除突变体，并进行表型鉴定与致病力分析。 【结果】 BcHMT2敲除突变体生长速率显著减慢，菌核产量和产孢量均降低；菌丝细胞变小，细胞壁降解酶活性降低，分生孢子形态异常。此外，该突变体对番茄果实和烟草叶片的致病力明显减弱。 【结论】 本研究揭示了BcHMT2在调控灰葡萄孢生长发育及致病过程中的作用，为灰葡萄孢防控研究提供了新见解。

, authors=李宇波1, 2, 邓思琪1, 董广恩1, 藏金萍1, 司贺龙1, 董金皋1, 2, 邢继红1, 2, 张康1, 2, authorsList=李宇波, 邓思琪, 董广恩, 藏金萍, 司贺龙, 董金皋, 邢继红, 张康, authorCompany=null, correspAuthors=邢继红, 张康, authorNote=

作者贡献声明

李宇波：试验操作、数据分析及文章撰写；邓思琪：数据分析及文章撰写；董广恩：部分试验操作、数据收集及分析；臧金萍：提供经费支持；司贺龙：参与文章编辑和审阅；董金皋：参与技术指导及文章审阅；邢继红：提供技术指导，研究构思和设计及文章审阅与修改；张康：项目申领、资源提供等贡献。

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G3 Genes|Genomes|Genetics, 2019, 9(12): 4087-4096., articleTitle=rmtA-dependent transcriptome and its role in secondary metabolism, environmental stress, and virulence in Aspergillus flavus, refAbstract=null)], funds=[Fund(id=1297571016746750005, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, awardId=32072369, language=EN, fundingSource=National Natural Science Foundation of China(32072369), fundOrder=null, country=null), Fund(id=1297571018650964025, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, awardId=32072369, language=CN, fundingSource=国家自然科学基金(32072369), fundOrder=null, country=null), Fund(id=1297571018780987450, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, awardId=246Z6506G, language=EN, fundingSource=Central Guidance for Local Technology Development Funding(246Z6506G), fundOrder=null, country=null), Fund(id=1297571019070394427, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, awardId=246Z6506G, language=CN, fundingSource=中央引导地方科技发展资金(246Z6506G), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1297571010711147498, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, xref=1., ext=[AuthorCompanyExt(id=1297571010719536107, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, companyId=1297571010711147498, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1.Hebei Key Laboratory of Plant Physiology and Molecular Pathology, Baoding, Hebei, China), AuthorCompanyExt(id=1297571010727924716, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, companyId=1297571010711147498, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1.河北省植物生理与分子病理学重点实验室，河北 保定)]), AuthorCompany(id=1297571010799227885, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, xref=2., ext=[AuthorCompanyExt(id=1297571010807616494, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, companyId=1297571010799227885, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2.State Key Laboratory of North China Crop Improvement and Regulation, Baoding, Hebei, China), AuthorCompanyExt(id=1297571010816005103, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, companyId=1297571010799227885, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2.华北作物改良与调控国家重点实验室，河北 保定)])], figs=[ArticleFig(id=1297571015312298021, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=EN, label=Figure 1, caption=Phylogenetic analysis of BcHMT2 and conserved domains of BcHMT2. A: Multiple sequence alignment of BcHMT2 homologous sequences; B: Phylogenetic tree and conserved domains of BcHMT2., figureFileSmall=trkLIY02kx+N6MN9yEnbZA==, figureFileBig=aZ7W9HBAC3Iny8YoeysHeA==, tableContent=null), ArticleFig(id=1297571015404572710, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=CN, label=图1, caption=BcHMT2 的系统发育分析与保守结构域, figureFileSmall=trkLIY02kx+N6MN9yEnbZA==, figureFileBig=aZ7W9HBAC3Iny8YoeysHeA==, tableContent=null), ArticleFig(id=1297571015563956263, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=EN, label=Figure 2, caption=Molecular identification of the BcHMT2 knockout mutants. A: Schematic diagram of the knockout strategy and primer design; B: DNA-level identification of transformants (W: B05.10, 1: Transformant); C: qPCR analysis of transformants; D: Western blotting analysis of histone methylation levels in transformants (** indicates P<0.01)., figureFileSmall=jxpap1bnTWMY7ls0pEj67A==, figureFileBig=wun4RJ9G1255b84UTbAY9Q==, tableContent=null), ArticleFig(id=1297571015626870824, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=CN, label=图2, caption=BcHMT2 敲除突变体的分子鉴定, figureFileSmall=jxpap1bnTWMY7ls0pEj67A==, figureFileBig=wun4RJ9G1255b84UTbAY9Q==, tableContent=null), ArticleFig(id=1297571015706562601, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=EN, label=Figure 3, caption=Colony morphology and growth rate of wild-type and mutant strains. A: Colony morphology on PDA medium; B: Determination of growth rate., figureFileSmall=YXgkgK3PuMv4X+yER3SDkA==, figureFileBig=TIuziTGibEdMqHn5Vy6EEg==, tableContent=null), ArticleFig(id=1297571015777865770, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=CN, label=图3, caption=野生型和突变体的菌落形态及生长速率, figureFileSmall=YXgkgK3PuMv4X+yER3SDkA==, figureFileBig=TIuziTGibEdMqHn5Vy6EEg==, tableContent=null), ArticleFig(id=1297571015903694891, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=EN, label=Figure 4, caption=Hyphal morphology in wild-type and mutant strains. A: Hyphal morphology under light microscopy (scale bar: 100 μm); B: Statistical analysis of hyphal cell length and width (**: P<0.01)., figureFileSmall=iznchJrt9F1UXts6C1c6Kw==, figureFileBig=l4tdIgWXVAzpZMtRmK1ApA==, tableContent=null), ArticleFig(id=1297571015979192364, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=CN, label=图4, caption=野生型和突变体的菌丝形态观察, figureFileSmall=iznchJrt9F1UXts6C1c6Kw==, figureFileBig=l4tdIgWXVAzpZMtRmK1ApA==, tableContent=null), ArticleFig(id=1297571016050495533, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=EN, label=Figure 5, caption=Analysis of cell wall degrading enzyme activity in ΔBcHMT2 mutants. A: Intracellular cell wall degrading enzyme activity assay; B: Extracellular cell wall degrading enzyme activity assay (**: P<0.01)., figureFileSmall=tXxtjLEtdNk4afOBbTlChQ==, figureFileBig=2OfMkfzwj6dV6JYw4nff3A==, tableContent=null), ArticleFig(id=1297571016134381614, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=CN, label=图5, caption=ΔBcHMT2 突变体胞壁降解酶活性测定, figureFileSmall=tXxtjLEtdNk4afOBbTlChQ==, figureFileBig=2OfMkfzwj6dV6JYw4nff3A==, tableContent=null), ArticleFig(id=1297571016209879087, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=EN, label=Figure 6, caption=Conidial morphology and conidiation yield in wild-type and mutant strains. A: Conidial morphology (scale bar: 100 μm); B: Scatter plot of conidial length-width ratio; C: Statistical analysis of conidial production (** indicates P<0.01)., figureFileSmall=axyRIMElrYqWc7rgNhXlhQ==, figureFileBig=zz8gzO/3Pm+0wP8uJLjXIQ==, tableContent=null), ArticleFig(id=1297571016289570864, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=CN, label=图6, caption=野生型和突变体分生孢子形态及产孢量, figureFileSmall=axyRIMElrYqWc7rgNhXlhQ==, figureFileBig=zz8gzO/3Pm+0wP8uJLjXIQ==, tableContent=null), ArticleFig(id=1297571016352485425, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=EN, label=Figure 7, caption=Pathogenicity analysis of wild-type and mutant strains. A: Pathogenic symptoms on tomato fruits; B: Statistical analysis of lesion area on tomato fruits; C: Pathogenic symptoms on tobacco leaves; D: Statistical analysis of lesion area on tobacco leaves. **: P<0.01., figureFileSmall=vR0f6aIvcW51AvSgJ684rA==, figureFileBig=zh+myysVSsGTairyqrTm1g==, tableContent=null), ArticleFig(id=1297571016448954418, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=CN, label=图7, caption=突变体和野生型的致病力分析, figureFileSmall=vR0f6aIvcW51AvSgJ684rA==, figureFileBig=zh+myysVSsGTairyqrTm1g==, tableContent=null), ArticleFig(id=1297571016516063283, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=EN, label=Table 1, caption=

Primers for construction of the BcHMT2 knockout vector

, figureFileSmall=null, figureFileBig=null, tableContent=
Gene namesPrimer namesPrimer sequences (5′→3′)
BcHMT2BcHMT2-I-FGCGGCCGCGAAAATTCTGAGCGGTTAAGGATT
BcHMT2-I-RACTAGTGATTATGTTTGTATTGTGCTTGGAA
BcHMT2BcHMT2-Ⅱ-FCTCGAGAGAGCTCTTCCAAATTCCTAAGTAA
BcHMT2-Ⅱ-RGGTACCATCCCCCGGGTATACTTTCCA
BcHMT2, hygromycin BBcHMT2-P1CCAGGGCCATCGAGCTTCAATATC
BcHMT2-P2AGGCGATGTTCGGGGATTC
BcHMT2, hygromycin BBcHMT2-P3GTAGAAACCATCGGCGCAG
BcHMT2-P4AAACCCAACCTGTCTGCAGACCAAA
BcHMT2, hygromycin BBcHMT2-P5CTATTCCTTTGCCCTCGGA
BcHMT2-P6ATGAAAAAGCCTGAACTCACCGC
BcHMT2BcHMT2-P7TCCTTGCAATGTTGAATCATAGCAT
BcHMT2-P8AGGTGACACATTGGAAACAAGGTGT
BcHMT2BcHMT2-P9ATATAAATTAACGGCGAGGAGGTAA
BcHMT2-P10TCTTCAACATATAATTTAGTGCCGG
BcHMT2BcHMT2-qRT-FGTACCGATGCCTACCGTGAC
BcHMT2-qRT-RCACCAGCTTTGGCACAGAAC
TubulinTubulin-FTCTGGCGAGCACGGTCTTGACGGTT
Tubulin-RTGGCTCCAAATCGACGAGGACGGCA
), ArticleFig(id=1297571016595755060, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571007133406169, language=CN, label=表1, caption=

BcHMT2 敲除载体构建引物

, figureFileSmall=null, figureFileBig=null, tableContent=
Gene namesPrimer namesPrimer sequences (5′→3′)
BcHMT2BcHMT2-I-FGCGGCCGCGAAAATTCTGAGCGGTTAAGGATT
BcHMT2-I-RACTAGTGATTATGTTTGTATTGTGCTTGGAA
BcHMT2BcHMT2-Ⅱ-FCTCGAGAGAGCTCTTCCAAATTCCTAAGTAA
BcHMT2-Ⅱ-RGGTACCATCCCCCGGGTATACTTTCCA
BcHMT2, hygromycin BBcHMT2-P1CCAGGGCCATCGAGCTTCAATATC
BcHMT2-P2AGGCGATGTTCGGGGATTC
BcHMT2, hygromycin BBcHMT2-P3GTAGAAACCATCGGCGCAG
BcHMT2-P4AAACCCAACCTGTCTGCAGACCAAA
BcHMT2, hygromycin BBcHMT2-P5CTATTCCTTTGCCCTCGGA
BcHMT2-P6ATGAAAAAGCCTGAACTCACCGC
BcHMT2BcHMT2-P7TCCTTGCAATGTTGAATCATAGCAT
BcHMT2-P8AGGTGACACATTGGAAACAAGGTGT
BcHMT2BcHMT2-P9ATATAAATTAACGGCGAGGAGGTAA
BcHMT2-P10TCTTCAACATATAATTTAGTGCCGG
BcHMT2BcHMT2-qRT-FGTACCGATGCCTACCGTGAC
BcHMT2-qRT-RCACCAGCTTTGGCACAGAAC
TubulinTubulin-FTCTGGCGAGCACGGTCTTGACGGTT
Tubulin-RTGGCTCCAAATCGACGAGGACGGCA
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精氨酸甲基转移酶BcHMT2调控灰葡萄孢生长发育及致病力的分子机制
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李宇波 1, 2 , 邓思琪 1 , 董广恩 1 , 藏金萍 1 , 司贺龙 1 , 董金皋 1, 2 , 邢继红 1, 2 , 张康 1, 2
微生物学报 | 研究报告 2026,66(8): 3889-3901
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微生物学报 |研究报告 2026 , 66 (8) : 3889 -3901
精氨酸甲基转移酶BcHMT2调控灰葡萄孢生长发育及致病力的分子机制
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李宇波1, 2, 邓思琪1, 董广恩1, 藏金萍1, 司贺龙1, 董金皋1, 2, 邢继红1, 2 , 张康1, 2
作者信息
  • 1.河北省植物生理与分子病理学重点实验室，河北 保定
  • 2.华北作物改良与调控国家重点实验室，河北 保定
通讯作者:
邢继红, 张康
作者简介:

作者贡献声明

李宇波：试验操作、数据分析及文章撰写；邓思琪：数据分析及文章撰写；董广恩：部分试验操作、数据收集及分析；臧金萍：提供经费支持；司贺龙：参与文章编辑和审阅；董金皋：参与技术指导及文章审阅；邢继红：提供技术指导，研究构思和设计及文章审阅与修改；张康：项目申领、资源提供等贡献。

Molecular mechanism of arginine methyltransferase BcHMT2 in regulating the growth, development, and pathogenicity of Botrytis cinerea
Yubo LI1, 2, Siqi DENG1, Guang’en DONG1, Jinping ZANG1, Helong SI1, Jingao DONG1, 2, Jihong XING1, 2 , Kang ZHANG1, 2
Affiliations
  • 1.Hebei Key Laboratory of Plant Physiology and Molecular Pathology, Baoding, Hebei, China
  • 2.State Key Laboratory of North China Crop Improvement and Regulation, Baoding, Hebei, China
    • Corresponding Author:
    E-mail: XING Jihong, ;
    ZHANG Kang,
出版时间: 2026-08-04 doi: 10.13343/j.cnki.wsxb.20260164
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【目的】 阐明精氨酸甲基转移酶在灰葡萄孢(Botrytis cinerea)生长发育及致病过程中的作用。 【方法】 利用生物信息学手段对灰葡萄孢、酿酒酵母(Saccharomyces cerevisiae)、禾谷镰孢(Fusarium graminearum)、稻瘟病菌(Magnaporthe oryzae)等8种真菌的精氨酸甲基转移酶HMT2进行序列比对、系统发育分析与保守结构域预测，明确了灰葡萄孢中编码该酶的基因为BcHMT2。通过同源重组技术构建BcHMT2基因敲除突变体，并进行表型鉴定与致病力分析。 【结果】 BcHMT2敲除突变体生长速率显著减慢，菌核产量和产孢量均降低；菌丝细胞变小，细胞壁降解酶活性降低，分生孢子形态异常。此外，该突变体对番茄果实和烟草叶片的致病力明显减弱。 【结论】 本研究揭示了BcHMT2在调控灰葡萄孢生长发育及致病过程中的作用，为灰葡萄孢防控研究提供了新见解。

灰葡萄孢  /  精氨酸甲基转移酶  /  BcHMT2  /  生长发育  /  致病力

[Objective] To elucidate the role of arginine methyltransferase in the growth, development, and pathogenicity of Botrytis cinerea. [Methods] Bioinformatics approaches were used to perform sequence alignment, phylogenetic analysis, and conserved domain prediction of the arginine methyltransferase HMT2 from eight fungal species, including B. cinerea, Saccharomyces cerevisiae, Fusarium graminearum, and Magnaporthe oryzae. The gene encoding this enzyme in B. cinerea was identified as BcHMT2. The BcHMT2-deleted mutant was constructed via homologous recombination, followed by phenotypic characterization and pathogenicity assays. [Results] The BcHMT2-deleted mutant exhibited a significantly reduced growth rate, significantly weakend cell wall-degrading enzyme activity, and decreased sclerotial production and conidiation. Meanwhile, the hyphal cells became smaller and the conidial morphology was abnormal. In addition, the mutant displayed markedly attenuated pathogenicity on tomato fruits and tobacco leaves. [Conclusion] This study reveals the regulatory role of BcHMT2 in the growth, development, and pathogenicity of B. cinerea, providing novel insights into the research on the prevention and control of gray mold.

Botrytis cinerea  /  arginine methyltransferase  /  BcHMT2  /  growth and development  /  pathogenicity
李宇波, 邓思琪, 董广恩, 藏金萍, 司贺龙, 董金皋, 邢继红, 张康. 精氨酸甲基转移酶BcHMT2调控灰葡萄孢生长发育及致病力的分子机制. 微生物学报, 2026 , 66 (8) : 3889 -3901 . DOI: 10.13343/j.cnki.wsxb.20260164
Yubo LI, Siqi DENG, Guang’en DONG, Jinping ZANG, Helong SI, Jingao DONG, Jihong XING, Kang ZHANG. Molecular mechanism of arginine methyltransferase BcHMT2 in regulating the growth, development, and pathogenicity of Botrytis cinerea[J]. Acta Microbiologica Sinica, 2026 , 66 (8) : 3889 -3901 . DOI: 10.13343/j.cnki.wsxb.20260164
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灰葡萄孢(Botrytis cinerea)是一种危害性较强的植物病原真菌，该菌种有性繁殖阶段归类于子囊菌门葡萄孢盘菌属，无性繁殖阶段则属于盘菌亚门葡萄孢属[1-2]。在自然界中，灰葡萄孢主要以无性态形式存在和传播。该菌分生孢子梗通常丛生，细长且呈灰色至褐色，顶端膨大并产生小突起，其上着生球形或椭球形的分生孢子[3]。灰葡萄孢分布广泛，传播迅速，不仅是真菌分子生物学研究的重要模式生物，也是农业生产中的毁灭性病原物。它主要通过分生孢子、菌丝、侵染垫及菌核侵染寄主，可侵染超过2 000种植物，引发灰霉病，造成巨大的经济损失[4]。
灰葡萄孢的致病机制复杂，主要包括3个阶段：穿透寄主表皮、降解寄主细胞壁以及诱导寄主细胞死亡[5]。当寄主出现伤口时，灰葡萄孢的菌丝和分生孢子可通过表皮损伤直接侵入寄主组织，或利用侵染垫和附着胞等特化结构入侵植物幼嫩部位[6]。分生孢子附着于寄主表皮后会迅速形成附着胞，进而侵入寄主细胞。在这一过程中，侵染垫和附着胞起着至关重要的作用[7-8]。侵入后，灰葡萄孢可分泌多种细胞壁降解酶，涵盖多聚半乳糖醛酸酶、果胶酯酶、纤维素酶等，通过分解植物细胞壁的核心组分实现侵染[9-10]。
蛋白质翻译后修饰是调控蛋白功能的重要方式，其中组蛋白甲基化修饰尤为关键。甲基化修饰多发生于赖氨酸、精氨酸残基的侧链区域：赖氨酸位点可形成单甲基化、二甲基化及三甲基化修饰，精氨酸则以单甲基化、对称性二甲基化或非对称性二甲基化修饰为主[11]。精氨酸甲基化由蛋白质精氨酸甲基转移酶(protein arginine methyltransferases, PRMTs)催化，并以S-腺苷甲硫氨酸(S-adenosylmethionine, SAM)为甲基供体，使蛋白质精氨酸残基的胍氮原子甲基化[12]。目前，植物中PRMTs的研究主要集中在拟南芥和水稻等模式植物中，拟南芥基因组中已鉴定出9种PRMTs[13]，水稻中鉴定出8种[14]。在酿酒酵母中，已鉴定出Hmt1、Hsl7和Rmt2共3种PRMTs，其中Hmt1作为主要的精氨酸甲基转移酶，参与核质转运、转录激活与延伸、mRNA剪接及信号转导等多种生物学过程。在禾谷镰孢中，AMT1被证实发挥主要生物学功能，其缺失会导致病菌侵染力与致病力显著下降[15]。尽管精氨酸甲基转移酶在多种丝状真菌中已有研究，但其在灰葡萄孢中的功能尚无报道。因此，本研究利用生物信息学方法从灰葡萄孢中鉴定出精氨酸甲基转移酶基因BcHMT2，通过构建其敲除突变体分析其对灰葡萄孢生长、发育及致病力的影响。本研究旨在明确精氨酸甲基转移酶在灰葡萄孢生长发育及致病过程中的功能与调控机制，一方面为深入解析该病原菌的致病分子机理夯实理论基础，另一方面也为筛选新型防治靶点、攻克灰霉病防控难题提供科学依据。
1 材料与方法
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1.1 材料
本研究所使用的同源重组载体为pBS-pUC载体。供试菌株为灰葡萄孢(Botrytis cinerea)野生型B05.10，番茄品种为‘京樊502’，烟草品种为本氏烟草(Nicotiana benthamiana)。以上材料均由本实验室提供。
1.2 灰葡萄孢 BcHMT2 基因的生物信息学分析
利用Ensembl Fungi数据库(http://fungi.ensembl.org/index.html)和NCBI数据库(http://www.ncbi.nlm.nih.gov)，获取灰葡萄孢(Botrytis cinerea)、酿酒酵母(Saccharomyces cerevisiae)、禾谷镰孢(Fusarium graminearum)、稻瘟病菌(Magnaporthe oryzae)、粗糙脉孢菌(Neurospora crassa)、黄曲霉(Aspergillus flavus)、构巢曲霉(Aspergillus nidulans)和白色念珠菌(Candida albicans) 8种真菌的HMT2蛋白序列。借助ClustalX2软件完成上述蛋白序列的多序列比对，再利用MEGA 7.0软件采用邻接法(neighbor-joining method)构建系统发育树，设置bootstrap重复次数为500次，以验证发育关系的可信度。
1.3 BcHMT2 基因敲除突变体的构建
根据灰葡萄孢全基因组DNA序列设计BcHMT2基因前后800 bp分别为前、后同源臂。将前、后同源臂分别与pBS-pUC载体连接，构建重组载体。通过原生质体转化技术将重组载体转入灰葡萄孢B05.10的原生质体中。转化成功后，用含0.1%潮霉素和0.2%氨苄青霉素(ampicillin, Amp)的RPDA培养基覆盖长出的菌丝。待单菌落长出后，将其转移至相同抗性的PDA培养基上，于20 ℃黑暗条件下培养。挑取菌落边缘菌丝转接至双抗PDA培养基，重复筛选至少3次。
1.4 敲除转化子的分子鉴定
采用CTAB法分别提取灰葡萄孢野生型B05.10和3个转化子的基因组DNA，作为PCR模板进行DNA水平鉴定。分别提取灰葡萄孢野生型B05.10和3个转化子的RNA，然后用七步法反转录试剂盒(Biosharp公司)合成cDNA，具体步骤详见试剂盒说明书。以Tubulin基因为内参基因，以野生型B05.10和3个转化子的cDNA为模板，利用qPCR技术检测3个转化子中BcHMT2基因的表达水平。扩增体系(20 μL)：cDNA 3 μL，2×Hieff qPCR SYBR Green Master Mix (No Rox) 7 μL，上、下游引物(10 µmol/L)各0.4 μL，ddH2O 9.2 μL。反应程序：95 ℃预变性6 min；95 ℃变性10 s，59.5 ℃退火20 s，72 ℃延伸20 s，共40个循环。此外，将野生型和3个转化子接种到PDA培养基中培养14 d后，提取各菌株的总蛋白：取研磨好的样品1 g，加入蛋白提取液1 mL，冰浴10 min (每2 min混匀1次)，4 ℃、12 000 r/min离心20 min，将上清分装后置于-80 ℃冰箱保存。然后利用Western blotting技术进行组蛋白H3及H3R2me2a的杂交检测。本研究所用引物见表1。
1.5 ΔBcHMT2 突变体的菌落形态观察及生长速率测定
采用8 mm打孔器在同一生长阶段的野生型菌株与ΔBcHMT2突变体菌株菌落边缘打取菌饼，转接至无抗性PDA培养基，置于20 ℃黑暗环境培养4 d。每日固定时段测定并记录菌落直径，各菌株均设置3次生物学重复。利用GraphPad Prism软件对数据进行单因素方差分析，*：P<0.05差异显著；**：P<0.01差异极显著。
1.6 ΔBcHMT2 突变体的菌丝形态观察
分别将同时期的野生型和ΔBcHMT2突变体菌株接种到无抗性PDA培养基上，并在每个培养基中斜插3片灭菌盖玻片。于20 ℃黑暗条件下培养5 d后，利用光学显微镜观察菌丝形态，测量并记录野生型B05.10及突变体菌丝细胞的长度和宽度。每株菌株设置3个重复。
1.7 ΔBcHMT2 突变体胞壁降解酶活性测定
取野生型和ΔBcHMT2突变体的菌盘放入液体PDB中，在黑暗条件下培养15 d，取出菌片并用液氮速冻。称取0.1 g菌片进行研磨，提取菌株胞内细胞壁降解酶；采用过滤去除菌丝后的发酵液，检测对应菌株的胞外酶活性。利用果胶酶活性检测试剂盒(Biopike公司)和纤维素酶活性检测试剂盒(北京索莱宝科技有限公司)，结合酶标仪检测各菌株细胞内、果胶酶(polygalacturonase, PG)和纤维素酶(cellulase, Cx)的活性。设置3组重复。
1.8 ΔBcHMT2 突变体分生孢子形态和产孢量测定
将同一生长期的野生型、ΔBcHMT2突变体菌株接种至无抗性PDA培养基，置于真菌培养箱培养15 d。用适量的ddH2O冲洗平皿中的菌丝2-3次，通过4层纱布过滤制备分生孢子悬浮液，并使用血球计数板统计分生孢子数量。将分生孢子悬浮液吹吸混匀后，在显微镜下观察分生孢子的形态。每株菌株设置3个重复。
1.9 ΔBcHMT2 突变体的致病力测定
分别将同一生长时期的野生型和ΔBcHMT2突变体菌株接种到无抗性PDA培养基上，培养7 d后，各自取直径为10 mm的菌盘，均匀放置于番茄和烟草叶片上。将接种后的植物置于保湿缸中，于25 ℃黑暗条件下培养，接种2 d后记录病斑面积并拍照。每种处理设置3组重复。
2 结果与分析
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2.1 灰葡萄孢精氨酸甲基转移酶基因 BcHMT2 的鉴定
通过Ensembl Fungi和NCBI数据库获取灰葡萄孢、酿酒酵母、禾谷镰孢、稻瘟病菌、粗糙脉孢菌、黄曲霉、构巢曲霉和白色念珠菌共8种真菌的HMT2蛋白序列，利用MEGA 7.0软件进行多序列比对及系统发育分析。结果显示，8种真菌的HMT2蛋白序列相似性较高(图1A)，其中BcHMT2与其余7种蛋白的序列相似性分别为57.05%、47.39%、56.82%、56.88%、60.04%、57.67%、50.17%，表明HMT2在真菌界具有高度保守性且同源关系密切。进一步利用SMART和Pfam数据库分析保守结构域，并通过IBS软件绘制结构域图谱，发现所有供试真菌的HMT2蛋白均含有典型的甲基转移酶(methyltransferase)结构域(图1B)。基于上述生物信息学分析，初步推断BcHMT2为灰葡萄孢中编码精氨酸甲基转移酶的基因。
2.2 BcHMT2敲除突变体的构建与分子鉴定
依据BcHMT2基因敲除策略设计引物(图2A)，对获得的3株转化子进行分子鉴定。DNA水平鉴定显示，利用同源臂引物P1/P2和P3/P4扩增，3株转化子均获得约1 400 bp的目标条带，而野生型B05.10未扩增出相应条带；利用潮霉素特异性引物P5/P6扩增，转化子获得约800 bp条带，野生型无条带。进一步利用验证引物P1/P7、P8/P4、P9/P7和P8/P10进行PCR鉴定，结果显示野生型B05.10扩增出约800 bp (P1/P7和P8/P4)和1 500 bp (P9/P7和P8/P10)的特异性条带，而转化子未获得相应条带(图2B)，证明BcHMT2基因已被成功替换。
转录水平检测显示，3株转化子中BcHMT2基因的表达量较野生型均显著降低(图2C)。蛋白水平上，Western blotting检测组蛋白甲基化修饰水平发现，与野生型相比，转化子中组蛋白H3R2me2a的甲基化水平显著下降，而组蛋白H3作为内参保持一致(图2D)。上述结果证实成功构建了BcHMT2基因敲除突变体，将其命名为ΔBcHMT2-1、ΔBcHMT2-2、ΔBcHMT2-3。
2.3 ΔBcHMT2 突变体菌落形态观察及生长速率测定
将野生型B05.10和3株ΔBcHMT2突变体接种于无抗性PDA培养基上，在相同条件下培养观察。结果显示，ΔBcHMT2突变体菌落颜色较野生型(灰色)略浅，且菌核产量明显减少(图3A)。生长速率测定表明，突变体的生长受到显著抑制：培养4 d后，野生型菌落直径达到9.00 cm，而3株突变体的菌落直径分别仅为7.41、7.50、8.00 cm (图3B)。这表明BcHMT2基因对灰葡萄孢的营养生长至关重要。
2.4 ΔBcHMT2 突变体菌丝形态观察
显微镜观察发现，ΔBcHMT2突变体的菌丝形态发生明显变化，表现为菌丝直径变细、分支减少(图4A)。形态测量结果显示，突变体菌丝细胞的长度和宽度均极显著减小(图4B)，表明BcHMT2基因缺失影响了菌丝的细胞骨架构建或细胞壁合成。
2.5 ΔBcHMT2 突变体胞壁降解酶活性测定
利用果胶酶和纤维素酶活性检测试剂盒检测野生型和3个ΔBcHMT2突变体胞内(图5A)、胞外(图5B)果胶酶(PG)和纤维素酶(Cx)的酶活性。结果发现，与野生型相比，3个ΔBcHMT2突变体的2种酶活性均显著降低，表明BcHMT2基因在该过程中发挥正调控作用。
2.6 ΔBcHMT2 突变体分生孢子形态及产孢量分析
为探究BcHMT2基因对灰葡萄孢分生孢子发育的影响，利用光学显微镜观察野生型和突变体菌株的分生孢子形态。结果显示，野生型分生孢子呈饱满的椭圆形或圆形，而突变体的分生孢子体积明显偏小，形态多为不规则椭圆形(图6A)。散点图统计进一步证实了突变体分生孢子长宽比的异常(图6B)。产孢量测定结果显示，3株突变体的产孢量均极显著低于野生型(图6C)，表明BcHMT2基因是灰葡萄孢正常产孢所必需的。
2.7 BcHMT2 基因对病菌致病力的影响
致病力测定结果显示，接种野生型菌株的番茄果实和烟草叶片均出现明显的腐烂症状和病斑扩展；而接种ΔBcHMT2突变体的组织仅出现小型水浸状斑点，侵染能力明显减弱(图7A、7C)。分析表明，突变体在番茄和烟草上的病斑面积均极显著小于野生型(图7B、7D)。上述结果表明，BcHMT2基因的缺失导致灰葡萄孢致病力大幅下降。
3 讨论与结论
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蛋白质精氨酸甲基转移酶(PRMTs)在真菌界具有高度保守性，且在真菌生长发育调控及致病过程中具有重要生物学功能[16]。在稻瘟病菌(Magnaporthe oryzae)中，基因组编码的4种精氨酸甲基转移酶中MoHMT1基因被证实是菌丝发育和分生孢子形成所必需的，敲除MoHMT1基因导致菌落颜色变浅、生长速率减缓及产孢量显著下降[17]。在禾谷镰孢(Fusarium graminearum)中，组蛋白精氨酸甲基转移酶AMT1调控病菌的营养生长，其缺失突变体表现出明显的生长抑制表型[18]。在黄曲霉(Aspergillus flavus)中，rmtA不仅调节菌核的形成，还是黄曲霉毒素B1合成及其他代谢产物合成的关键调控因子[19]。此外，在白色念珠菌(Candida albicans)中，CaHMT1基因的缺失导致不对称二甲基精氨酸和ω-单甲基精氨酸水平显著降低，表明其在精氨酸甲基化修饰中占据主导地位[20]。
近年来的研究进一步揭示了PRMTs在病原菌致病过程中的关键调控功能。稻瘟病菌作为全球性的主要作物病原菌，其MoHMT1基因敲除突变体在接种水稻和大麦幼苗后未能形成明显病斑，证实了MoHMT1在致病过程中的不可或缺性[16-17]。在黄曲霉中，rmtA通过正调控veA基因的表达进而影响黄曲霉毒素B1的合成及aflJ等基因的表达，深度参与了致病与毒素合成的调控网络[21]。
本研究利用同源重组技术，获得了灰葡萄孢BcHMT2基因的敲除突变体(ΔBcHMT2-1、ΔBcHMT2-2和ΔBcHMT2-3)。表型分析结果显示，与野生型B05.10相比，BcHMT2缺失导致菌落颜色变浅、生长速率显著减慢、菌核产量降低。微观形态观察发现，突变体菌丝细胞变小、分生孢子形态异常(体积偏小且呈不规则椭圆形)。更重要的是，致病力测定表明，突变体对番茄果实和烟草叶片的致病力显著减弱，病斑扩展受到明显抑制。这些表型特征与稻瘟病菌MoHMT1和禾谷镰孢AMT1的缺失表型具有高度相似性，进一步证实了HMT2家族基因在丝状真菌生长发育和致病过程中的保守功能。Western blotting结果显示，BcHMT2的缺失导致组蛋白H3R2me2a (不对称二甲基化)水平显著降低，这与白色念珠菌中CaHMT1的功能一致，表明BcHMT2主要通过催化组蛋白H3R2的不对称二甲基化修饰，进而调控下游基因的表达，最终影响灰葡萄孢的形态建成和致病力。
综上所述，本研究明确了精氨酸甲基转移酶基因BcHMT2在灰葡萄孢中的关键生物学功能。BcHMT2不仅调控灰葡萄孢的营养生长、菌丝形态和分生孢子发育，还对其致病力具有重要的正向调控作用。本研究不仅丰富了对灰葡萄孢表观遗传调控机制的理解，也为开发以PRMTs为靶标的新型杀菌剂提供了理论依据和潜在的基因资源。后续研究将进一步解析BcHMT2调控的下游靶基因网络及其在灰霉病防治中的应用潜力。
基金
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  • 国家自然科学基金(32072369)
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2026年第66卷第8期
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doi: 10.13343/j.cnki.wsxb.20260164
  • 接收时间：2026-02-28
  • 首发时间：2026-08-21
  • 出版时间：2026-08-04
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  • 收稿日期：2026-02-28
  • 录用日期：2026-03-17
基金
National Natural Science Foundation of China(32072369)
国家自然科学基金(32072369)
Central Guidance for Local Technology Development Funding(246Z6506G)
中央引导地方科技发展资金(246Z6506G)
作者信息
    1.河北省植物生理与分子病理学重点实验室，河北 保定
    2.华北作物改良与调控国家重点实验室，河北 保定

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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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