首页 期刊 论文 学科刊群 资讯 加盟 关于
EN
image
Article(id=1226956549617267460, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226956547847275311, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250131, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1740153600000, receivedDateStr=2025-02-22, revisedDate=null, revisedDateStr=null, acceptedDate=1745337600000, acceptedDateStr=2025-04-23, onlineDate=1770458837179, onlineDateStr=2026-02-07, pubDate=1756915200000, pubDateStr=2025-09-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1770458837179, onlineIssueDateStr=2026-02-07, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1770458837179, creator=13701087609, updateTime=1770458837179, updator=13701087609, issue=Issue{id=1226956547847275311, tenantId=1146029695717560320, journalId=1192105938417971205, year='2025', volume='65', issue='9', pageStart='3821', pageEnd='4232', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1770458836757, creator=13701087609, updateTime=1770459153781, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1226957877613605816, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226956547847275311, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1226957877613605817, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226956547847275311, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=3848, endPage=3858, ext={EN=ArticleExt(id=1226956550317716233, articleId=1226956549617267460, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Research progress of bacterial DNA methyltransferase CcrM involved in epigenetic regulation, columnId=1192149543727808575, journalTitle=Acta Microbiologica Sinica, columnName=Review, runingTitle=null, highlight=null, articleAbstract=

DNA methylation is an important way of epigenetic regulation in bacteria. Alphaproteobacteria methylate DNA by using the cell cycle-regulated DNA methyltransferase (CcrM). CcrM does not contain a functional unit of restriction endonuclease, thus belonging to an orphan methyltransferase. By methylating adenine in DNA sequences, CcrM influences the interaction between DNA and proteins, regulates the expression of numerous genes, and is crucial for the regulation of processes such as the cell cycle of Alphaproteobacteria. We reviewed the function, structure, and epigenetic regulation of CcrM, clarified the mechanisms of CcrM in DNA recognition, catalysis, and activity regulation, summarized the mechanism by which the global cell-cycle regulator (GcrA) utilizes the methylation signals produced by CcrM to regulate gene expression, and provided an outlook on the potential future research directions of CcrM, providing a reference for further in-depth study of the epigenetic regulation mechanisms in bacteria.

, correspAuthors=null, authorNote=null, correspAuthorsNote=
*E-mail:
, 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, authorCompany=null, fund=null, authors=null, authorsList=Hao WANG, Erya WANG, Minliang GUO), CN=ArticleExt(id=1226956552670720805, articleId=1226956549617267460, tenantId=1146029695717560320, journalId=1192105938417971205, language=CN, title=参与表观遗传调控的细菌DNA甲基转移酶CcrM的研究进展, columnId=1192149543882997826, journalTitle=微生物学报, columnName=综述, runingTitle=null, highlight=null, articleAbstract=

DNA甲基化是细菌进行表观遗传调控的重要方式，α变形菌利用受细胞周期调控的DNA甲基转移酶(cell cycle-regulated DNA methyltransferase, CcrM)对DNA进行甲基化。CcrM不含限制性内切酶功能单元，属于孤儿甲基转移酶。CcrM通过对序列中腺嘌呤进行甲基化影响DNA与蛋白质的相互作用，从而调节大量基因的表达，对α变形菌细胞周期等过程的调控至关重要。本文综述了CcrM的功能、结构及其表观调控机制，阐明了CcrM对DNA的识别、催化及活性调控的机理，总结了细胞周期全局性调控因子(global cell-cycle regulator, GcrA)利用CcrM的甲基化信号调节基因表达的机制，并展望了CcrM未来的潜在研究方向，为进一步深入研究细菌表观遗传调控机制提供了参考。

, correspAuthors=null, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=63mUAIgMl2qrsl3w0dUcEg==, magXml=LE8pZfL1HuJCnjM0qkI99A==, pdfUrl=null, pdf=mQGaP9gIp+K8MxH+56uFUQ==, pdfFileSize=1725582, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=vWvOoAt5yrUqm+enbpNbkQ==, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=+TgyMaDGt2RjhBAnLaQc8A==, mapNumber=null, authorCompany=null, fund=null, authors=

作者贡献声明

王浩：论文的撰写和修改；王尔雅：资料的收集与整理、协助论文撰写；郭敏亮：论文整体构思与设计。

, authorsList=王浩, 王尔雅, 郭敏亮)}, authors=[Author(id=1226964048550478239, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, orderNo=0, firstName=null, middleName=null, lastName=null, nameCn=null, orcid=null, stid=null, country=null, authorPic=null, dead=0, email=null, emailSecond=null, emailThird=null, correspondingAuthor=0, authorType=1, ext={EN=AuthorExt(id=1226964048688890279, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, authorId=1226964048550478239, language=EN, stringName=Hao WANG, firstName=Hao, middleName=null, lastName=WANG, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=College of Bioscience and Biotechnology, Yangzhou University, Yangzhou, Jiangsu, China, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null), CN=AuthorExt(id=1226964048844079533, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, authorId=1226964048550478239, language=CN, stringName=王浩, firstName=null, middleName=null, lastName=null, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=扬州大学 生物科学与技术学院，江苏 扬州, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null)}, companyList=[AuthorCompany(id=1226964048282042771, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, xref=null, ext=[AuthorCompanyExt(id=1226964048412066199, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, companyId=1226964048282042771, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=College of Bioscience and Biotechnology, Yangzhou University, Yangzhou, Jiangsu, China), AuthorCompanyExt(id=1226964048420454810, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, companyId=1226964048282042771, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=扬州大学 生物科学与技术学院，江苏 扬州)])]), Author(id=1226964048957325748, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, orderNo=1, firstName=null, middleName=null, lastName=null, nameCn=null, orcid=null, stid=null, country=null, authorPic=null, dead=0, email=null, emailSecond=null, emailThird=null, correspondingAuthor=0, authorType=1, ext={EN=AuthorExt(id=1226964049175429572, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, authorId=1226964048957325748, language=EN, stringName=Erya WANG, firstName=Erya, middleName=null, lastName=WANG, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=College of Bioscience and Biotechnology, Yangzhou University, Yangzhou, Jiangsu, China, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null), CN=AuthorExt(id=1226964050551161299, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, authorId=1226964048957325748, language=CN, stringName=王尔雅, firstName=null, middleName=null, lastName=null, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=扬州大学 生物科学与技术学院，江苏 扬州, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null)}, companyList=[AuthorCompany(id=1226964048282042771, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, xref=null, ext=[AuthorCompanyExt(id=1226964048412066199, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, companyId=1226964048282042771, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=College of Bioscience and Biotechnology, Yangzhou University, Yangzhou, Jiangsu, China), AuthorCompanyExt(id=1226964048420454810, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, companyId=1226964048282042771, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=扬州大学 生物科学与技术学院，江苏 扬州)])]), Author(id=1226964050689573341, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, orderNo=2, firstName=null, middleName=null, lastName=null, nameCn=null, orcid=null, stid=null, country=null, authorPic=null, dead=0, email=guoml@yzu.edu.cn, emailSecond=null, emailThird=null, correspondingAuthor=0, authorType=1, ext={EN=AuthorExt(id=1226964050819596778, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, authorId=1226964050689573341, language=EN, stringName=Minliang GUO, firstName=Minliang, middleName=null, lastName=GUO, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=College of Bioscience and Biotechnology, Yangzhou University, Yangzhou, Jiangsu, China, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null), CN=AuthorExt(id=1226964050941231603, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, authorId=1226964050689573341, language=CN, stringName=郭敏亮, firstName=null, middleName=null, lastName=null, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=扬州大学 生物科学与技术学院，江苏 扬州, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null)}, companyList=[AuthorCompany(id=1226964048282042771, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, xref=null, ext=[AuthorCompanyExt(id=1226964048412066199, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, companyId=1226964048282042771, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=College of Bioscience and Biotechnology, Yangzhou University, Yangzhou, Jiangsu, China), AuthorCompanyExt(id=1226964048420454810, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, companyId=1226964048282042771, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=扬州大学 生物科学与技术学院，江苏 扬州)])])], keywords=[Keyword(id=1226964051247415814, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=EN, orderNo=1, keyword=CcrM), Keyword(id=1226964051373244948, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=EN, orderNo=2, keyword=bacterial DNA methylation), Keyword(id=1226964051482296866, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=EN, orderNo=3, keyword=epigenetic regulation), Keyword(id=1226964051591348775, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=EN, orderNo=4, keyword=GcrA), Keyword(id=1226964051712983601, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=CN, orderNo=1, keyword=受细胞周期调控的DNA甲基转移酶(CcrM)), Keyword(id=1226964051830424126, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=CN, orderNo=2, keyword=细菌DNA甲基化), Keyword(id=1226964051973030478, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=CN, orderNo=3, keyword=表观遗传调控), Keyword(id=1226964052086276703, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=CN, orderNo=4, keyword=细胞周期全局性调控因子(GcrA))], refs=[Reference(id=1226964053654946558, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2021, volume=29, issue=1, pageStart=28, pageEnd=40, url=null, language=null, rfNumber=[1], rfOrder=0, authorNames=OLIVEIRA PH, FANG G, journalName=Trends in Microbiology, refType=null, unstructuredReference=OLIVEIRA PH, FANG G. Conserved DNA methyltransferases: a window into fundamental mechanisms of epigenetic regulation in bacteria[J]. Trends in Microbiology, 2021, 29(1): 28-40., articleTitle=Conserved DNA methyltransferases: a window into fundamental mechanisms of epigenetic regulation in bacteria, refAbstract=null), Reference(id=1226964055051649800, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2017, volume=400, issue=null, pageStart=105, pageEnd=127, url=null, language=null, rfNumber=[2], rfOrder=1, authorNames=GORRELL R, KWOK T, journalName=Current Topics in Microbiology and Immunology, refType=null, unstructuredReference=GORRELL R, KWOK T. The Helicobacter pylori methylome: roles in gene regulation and virulence[J]. Current Topics in Microbiology and Immunology, 2017, 400: 105-127., articleTitle=The Helicobacter pylori methylome: roles in gene regulation and virulence, refAbstract=null), Reference(id=1226964055227810582, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2006, volume=4, issue=3, pageStart=183, pageEnd=192, url=null, language=null, rfNumber=[3], rfOrder=2, authorNames=WION D, CASADESÚS J, journalName=Nature Reviews Microbiology, refType=null, unstructuredReference=WION D, CASADESÚS J. N6-methyl-adenine: an epigenetic signal for DNA-protein interactions[J]. Nature Reviews Microbiology, 2006, 4(3): 183-192., articleTitle=N6-methyl-adenine: an epigenetic signal for DNA-protein interactions, refAbstract=null), Reference(id=1226964055320085279, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2013, volume=9, issue=5, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[4], rfOrder=3, authorNames=FIORAVANTI A, FUMEAUX C, MOHAPATRA SS, BOMPARD C, BRILLI M, FRANDI A, CASTRIC V, VILLERET V, VIOLLIER PH, BIONDI EG, journalName=PLoS Genetics, refType=null, unstructuredReference=FIORAVANTI A, FUMEAUX C, MOHAPATRA SS, BOMPARD C, BRILLI M, FRANDI A, CASTRIC V, VILLERET V, VIOLLIER PH, BIONDI EG. DNA binding of the cell cycle transcriptional regulator GcrA depends on N6-adenosine methylation in Caulobacter crescentus and other Alphaproteobacteria [J]. PLoS Genetics, 2013, 9(5): e1003541., articleTitle=DNA binding of the cell cycle transcriptional regulator GcrA depends on N6-adenosine methylation in Caulobacter crescentus and other Alphaproteobacteria, refAbstract=null), Reference(id=1226964055403971367, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2011, volume=7, issue=null, pageStart=528, pageEnd=null, url=null, language=null, rfNumber=[5], rfOrder=4, authorNames=CHRISTEN B, ABELIUK E, COLLIER JM, KALOGERAKI VS, PASSARELLI B, COLLER JA, FERO MJ, McADAMS HH, SHAPIRO L, journalName=Molecular Systems Biology, refType=null, unstructuredReference=CHRISTEN B, ABELIUK E, COLLIER JM, KALOGERAKI VS, PASSARELLI B, COLLER JA, FERO MJ, McADAMS HH, SHAPIRO L. The essential genome of a bacterium[J]. Molecular Systems Biology, 2011, 7: 528., articleTitle=The essential genome of a bacterium, refAbstract=null), Reference(id=1226964055508828977, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2014, volume=93, issue=4, pageStart=713, pageEnd=735, url=null, language=null, rfNumber=[6], rfOrder=5, authorNames=CURTIS PD, BRUN YV, journalName=Molecular Microbiology, refType=null, unstructuredReference=CURTIS PD, BRUN YV. Identification of essential alphaproteobacterial genes reveals operational variability in conserved developmental and cell cycle systems[J]. Molecular Microbiology, 2014, 93(4): 713-735., articleTitle=Identification of essential alphaproteobacterial genes reveals operational variability in conserved developmental and cell cycle systems, refAbstract=null), Reference(id=1226964055601103673, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=1994, volume=235, issue=2, pageStart=472, pageEnd=485, url=null, language=null, rfNumber=[7], rfOrder=6, authorNames=ZWEIGER G, MARCZYNSKI G, SHAPIRO L, journalName=Journal of Molecular Biology, refType=null, unstructuredReference=ZWEIGER G, MARCZYNSKI G, SHAPIRO L. A Caulobacter DNA methyltransferase that functions only in the predivisional cell[J]. Journal of Molecular Biology, 1994, 235(2): 472-485., articleTitle=A Caulobacter DNA methyltransferase that functions only in the predivisional cell, refAbstract=null), Reference(id=1226964055689184063, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2014, volume=42, issue=6, pageStart=3720, pageEnd=3735, url=null, language=null, rfNumber=[8], rfOrder=7, authorNames=GONZALEZ D, KOZDON JB, McADAMS HH, SHAPIRO L, COLLIER J, journalName=Nucleic Acids Research, refType=null, unstructuredReference=GONZALEZ D, KOZDON JB, McADAMS HH, SHAPIRO L, COLLIER J. The functions of DNA methylation by CcrM in Caulobacter crescentus: a global approach[J]. Nucleic Acids Research, 2014, 42(6): 3720-3735., articleTitle=The functions of DNA methylation by CcrM in Caulobacter crescentus: a global approach, refAbstract=null), Reference(id=1226964055806624584, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=1997, volume=179, issue=18, pageStart=5869, pageEnd=5877, url=null, language=null, rfNumber=[9], rfOrder=8, authorNames=WRIGHT R, STEPHENS C, SHAPIRO L, journalName=Journal of Bacteriology, refType=null, unstructuredReference=WRIGHT R, STEPHENS C, SHAPIRO L. The CcrM DNA methyltransferase is widespread in the alpha subdivision of proteobacteria, and its essential functions are conserved in Rhizobium meliloti and Caulobacter crescentus [J]. Journal of Bacteriology, 1997, 179(18): 5869-5877., articleTitle=The CcrM DNA methyltransferase is widespread in the alpha subdivision of proteobacteria, and its essential functions are conserved in Rhizobium meliloti and Caulobacter crescentus, refAbstract=null), Reference(id=1226964055907287893, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2010, volume=4, issue=null, pageStart=52, pageEnd=null, url=null, language=null, rfNumber=[10], rfOrder=9, authorNames=BRILLI M, FONDI M, FANI R, MENGONI A, FERRI L, BAZZICALUPO M, BIONDI EG, journalName=BMC Systems Biology, refType=null, unstructuredReference=BRILLI M, FONDI M, FANI R, MENGONI A, FERRI L, BAZZICALUPO M, BIONDI EG. The diversity and evolution of cell cycle regulation in alpha-proteobacteria: a comparative genomic analysis[J]. BMC Systems Biology, 2010, 4: 52., articleTitle=The diversity and evolution of cell cycle regulation in alpha-proteobacteria: a comparative genomic analysis, refAbstract=null), Reference(id=1226964056054088539, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2001, volume=183, issue=10, pageStart=3065, pageEnd=3075, url=null, language=null, rfNumber=[11], rfOrder=10, authorNames=KAHNG LS, SHAPIRO L, journalName=Journal of Bacteriology, refType=null, unstructuredReference=KAHNG LS, SHAPIRO L. The CcrM DNA methyltransferase of Agrobacterium tumefaciens is essential, and its activity is cell cycle regulated[J]. Journal of Bacteriology, 2001, 183(10): 3065-3075., articleTitle=The CcrM DNA methyltransferase of Agrobacterium tumefaciens is essential, and its activity is cell cycle regulated, refAbstract=null), Reference(id=1226964056154751842, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2000, volume=182, issue=12, pageStart=3482, pageEnd=3489, url=null, language=null, rfNumber=[12], rfOrder=11, authorNames=ROBERTSON GT, REISENAUER A, WRIGHT R, JENSEN RB, JENSEN A, SHAPIRO L, ROOP RM 2nd, journalName=Journal of Bacteriology, refType=null, unstructuredReference=ROBERTSON GT, REISENAUER A, WRIGHT R, JENSEN RB, JENSEN A, SHAPIRO L, ROOP RM 2nd. The Brucella abortus CcrM DNA methyltransferase is essential for viability, and its overexpression attenuates intracellular replication in murine macrophages[J]. Journal of Bacteriology, 2000, 182(12): 3482-3489., articleTitle=The Brucella abortus CcrM DNA methyltransferase is essential for viability, and its overexpression attenuates intracellular replication in murine macrophages, refAbstract=null), Reference(id=1226964056305746791, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2024, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[13], rfOrder=12, authorNames=王尔雅, journalName=null, refType=null, unstructuredReference=王尔雅. 根癌农杆菌细胞周期调节因子GcrA调节vir基因表达的机制研究[D]. 扬州: 扬州大学硕士学位论文, 2024., articleTitle=根癌农杆菌细胞周期调节因子GcrA调节vir基因表达的机制研究, refAbstract=null), Reference(id=1226964056423187311, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2024, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[13], rfOrder=13, authorNames=WANG EY, journalName=null, refType=null, unstructuredReference=WANG EY. Mechanism of vir gene expression regulated by Agrobacterium tumefaciens cell cycle regulator GcrA[D]. Yangzhou: Master’s Thesis of Yangzhou University, 2024 (in Chinese)., articleTitle=null, refAbstract=null), Reference(id=1226964056549016436, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2018, volume=293, issue=49, pageStart=19038, pageEnd=19046, url=null, language=null, rfNumber=[14], rfOrder=14, authorNames=REICH NO, DANG E, KURNIK M, PATHURI S, WOODCOCK CB, journalName=Journal of Biological Chemistry, refType=null, unstructuredReference=REICH NO, DANG E, KURNIK M, PATHURI S, WOODCOCK CB. The highly specific, cell cycle-regulated methyltransferase from Caulobacter crescentus relies on a novel DNA recognition mechanism[J]. Journal of Biological Chemistry, 2018, 293(49): 19038-19046., articleTitle=The highly specific, cell cycle-regulated methyltransferase from Caulobacter crescentus relies on a novel DNA recognition mechanism, refAbstract=null), Reference(id=1226964056683234175, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2017, volume=56, issue=30, pageStart=3913, pageEnd=3922, url=null, language=null, rfNumber=[15], rfOrder=15, authorNames=WOODCOCK CB, YAKUBOV AB, REICH NO, journalName=Biochemistry, refType=null, unstructuredReference=WOODCOCK CB, YAKUBOV AB, REICH NO. Caulobacter crescentus cell cycle-regulated DNA methyltransferase uses a novel mechanism for substrate recognition[J]. Biochemistry, 2017, 56(30): 3913-3922., articleTitle=Caulobacter crescentus cell cycle-regulated DNA methyltransferase uses a novel mechanism for substrate recognition, refAbstract=null), Reference(id=1226964056783897477, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=1998, volume=95, issue=6, pageStart=2874, pageEnd=2879, url=null, language=null, rfNumber=[16], rfOrder=16, authorNames=BERDIS AJ, LEE I, COWARD JK, STEPHENS C, WRIGHT R, SHAPIRO L, BENKOVIC SJ, journalName=Proceedings of the National Academy of Sciences of the United States of America, refType=null, unstructuredReference=BERDIS AJ, LEE I, COWARD JK, STEPHENS C, WRIGHT R, SHAPIRO L, BENKOVIC SJ. A cell cycle-regulated adenine DNA methyltransferase from Caulobacter crescentus processively methylates GANTC sites on hemimethylated DNA[J]. Proceedings of the National Academy of Sciences of the United States of America, 1998, 95(6): 2874-2879., articleTitle=A cell cycle-regulated adenine DNA methyltransferase from Caulobacter crescentus processively methylates GANTC sites on hemimethylated DNA, refAbstract=null), Reference(id=1226964056871977868, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2019, volume=10, issue=null, pageStart=4600, pageEnd=null, url=null, language=null, rfNumber=[17], rfOrder=17, authorNames=HORTON JR, WOODCOCK CB, OPOT SB, REICH NO, ZHANG X, CHENG XD, journalName=Nature Communications, refType=null, unstructuredReference=HORTON JR, WOODCOCK CB, OPOT SB, REICH NO, ZHANG X, CHENG XD. The cell cycle-regulated DNA adenine methyltransferase CcrM opens a bubble at its DNA recognition site[J]. Nature Communications, 2019, 10: 4600., articleTitle=The cell cycle-regulated DNA adenine methyltransferase CcrM opens a bubble at its DNA recognition site, refAbstract=null), Reference(id=1226964056951669651, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2020, volume=48, issue=20, pageStart=11589, pageEnd=11601, url=null, language=null, rfNumber=[18], rfOrder=18, authorNames=KONTTINEN O, CARMODY J, PATHURI S, ANDERSON K, ZHOU XF, REICH N, journalName=Nucleic Acids Research, refType=null, unstructuredReference=KONTTINEN O, CARMODY J, PATHURI S, ANDERSON K, ZHOU XF, REICH N. Cell cycle regulated DNA methyltransferase: fluorescent tracking of a DNA strand-separation mechanism and identification of the responsible protein motif[J]. Nucleic Acids Research, 2020, 48(20): 11589-11601., articleTitle=Cell cycle regulated DNA methyltransferase: fluorescent tracking of a DNA strand-separation mechanism and identification of the responsible protein motif, refAbstract=null), Reference(id=1226964057052332951, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2023, volume=51, issue=13, pageStart=6883, pageEnd=6898, url=null, language=null, rfNumber=[19], rfOrder=19, authorNames=KONTTINEN O, CARMODY J, KURNIK M, JOHNSON KA, REICH N, journalName=Nucleic Acids Research, refType=null, unstructuredReference=KONTTINEN O, CARMODY J, KURNIK M, JOHNSON KA, REICH N. High fidelity DNA strand-separation is the major specificity determinant in DNA methyltransferase CcrM’s catalytic mechanism[J]. Nucleic Acids Research, 2023, 51(13): 6883-6898., articleTitle=High fidelity DNA strand-separation is the major specificity determinant in DNA methyltransferase CcrM’s catalytic mechanism, refAbstract=null), Reference(id=1226964057157190558, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=1995, volume=253, issue=4, pageStart=618, pageEnd=632, url=null, language=null, rfNumber=[20], rfOrder=20, authorNames=MALONE T, BLUMENTHAL RM, CHENG X, journalName=Journal of Molecular Biology, refType=null, unstructuredReference=MALONE T, BLUMENTHAL RM, CHENG X. Structure-guided analysis reveals nine sequence motifs conserved among DNA amino-methyltransferases, and suggests a catalytic mechanism for these enzymes[J]. Journal of Molecular Biology, 1995, 253(4): 618-632., articleTitle=Structure-guided analysis reveals nine sequence motifs conserved among DNA amino-methyltransferases, and suggests a catalytic mechanism for these enzymes, refAbstract=null), Reference(id=1226964057245270948, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2003, volume=28, issue=6, pageStart=329, pageEnd=335, url=null, language=null, rfNumber=[21], rfOrder=21, authorNames=SCHUBERT HL, BLUMENTHAL RM, CHENG XD, journalName=Trends in Biochemical Sciences, refType=null, unstructuredReference=SCHUBERT HL, BLUMENTHAL RM, CHENG XD. Many paths to methyltransfer: a chronicle of convergence[J]. Trends in Biochemical Sciences, 2003, 28(6): 329-335., articleTitle=Many paths to methyltransfer: a chronicle of convergence, refAbstract=null), Reference(id=1226964057337545643, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=1996, volume=10, issue=12, pageStart=1532, pageEnd=1542, url=null, language=null, rfNumber=[22], rfOrder=22, authorNames=WRIGHT R, STEPHENS C, ZWEIGER G, SHAPIRO L, ALLEY MR, journalName=Genes & Development, refType=null, unstructuredReference=WRIGHT R, STEPHENS C, ZWEIGER G, SHAPIRO L, ALLEY MR. Caulobacter Lon protease has a critical role in cell-cycle control of DNA methylation[J]. Genes & Development, 1996, 10(12): 1532-1542., articleTitle=Caulobacter Lon protease has a critical role in cell-cycle control of DNA methylation, refAbstract=null), Reference(id=1226964057471763376, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=1999, volume=181, issue=8, pageStart=2430, pageEnd=2439, url=null, language=null, rfNumber=[23], rfOrder=23, authorNames=REISENAUER A, QUON K, SHAPIRO L, journalName=Journal of Bacteriology, refType=null, unstructuredReference=REISENAUER A, QUON K, SHAPIRO L. The CtrA response regulator mediates temporal control of gene expression during the Caulobacter cell cycle[J]. Journal of Bacteriology, 1999, 181(8): 2430-2439., articleTitle=The CtrA response regulator mediates temporal control of gene expression during the Caulobacter cell cycle, refAbstract=null), Reference(id=1226964057605981107, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2002, volume=43, issue=4, pageStart=945, pageEnd=960, url=null, language=null, rfNumber=[24], rfOrder=24, authorNames=BELLEFONTAINE AF, PIERREUX CE, MERTENS P, VANDENHAUTE J, LETESSON JJ, de BOLLE X, journalName=Molecular Microbiology, refType=null, unstructuredReference=BELLEFONTAINE AF, PIERREUX CE, MERTENS P, VANDENHAUTE J, LETESSON JJ, de BOLLE X. Plasticity of a transcriptional regulation network among alpha-proteobacteria is supported by the identification of CtrA targets in Brucella abortus [J]. Molecular Microbiology, 2002, 43(4): 945-960., articleTitle=Plasticity of a transcriptional regulation network among alpha-proteobacteria is supported by the identification of CtrA targets in Brucella abortus, refAbstract=null), Reference(id=1226964057782141883, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2017, volume=103, issue=5, pageStart=780, pageEnd=797, url=null, language=null, rfNumber=[25], rfOrder=25, authorNames=FRANCIS N, PONCIN K, FIORAVANTI A, VASSEN V, WILLEMART K, ONG TAP, RAPPEZ L, LETESSON JJ, BIONDI EG, de BOLLE X, journalName=Molecular Microbiology, refType=null, unstructuredReference=FRANCIS N, PONCIN K, FIORAVANTI A, VASSEN V, WILLEMART K, ONG TAP, RAPPEZ L, LETESSON JJ, BIONDI EG, de BOLLE X. CtrA controls cell division and outer membrane composition of the pathogen Brucella abortus [J]. Molecular Microbiology, 2017, 103(5): 780-797., articleTitle=CtrA controls cell division and outer membrane composition of the pathogen Brucella abortus, refAbstract=null), Reference(id=1226964057933136835, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2016, volume=30, issue=null, pageStart=107, pageEnd=113, url=null, language=null, rfNumber=[26], rfOrder=26, authorNames=COLLIER J, journalName=Current Opinion in Microbiology, refType=null, unstructuredReference=COLLIER J. Cell cycle control in Alphaproteobacteria [J]. Current Opinion in Microbiology, 2016, 30: 107-113., articleTitle=Cell cycle control in Alphaproteobacteria, refAbstract=null), Reference(id=1226964058042188745, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2010, volume=39, issue=3, pageStart=455, pageEnd=467, url=null, language=null, rfNumber=[27], rfOrder=27, authorNames=GORA KG, TSOKOS CG, CHEN YE, SRINIVASAN BS, PERCHUK BS, LAUB MT, journalName=Molecular Cell, refType=null, unstructuredReference=GORA KG, TSOKOS CG, CHEN YE, SRINIVASAN BS, PERCHUK BS, LAUB MT. A cell-type-specific protein-protein interaction modulates transcriptional activity of a master regulator in Caulobacter crescentus [J]. Molecular Cell, 2010, 39(3): 455-467., articleTitle=A cell-type-specific protein-protein interaction modulates transcriptional activity of a master regulator in Caulobacter crescentus, refAbstract=null), Reference(id=1226964058155434960, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=1995, volume=177, issue=7, pageStart=1662, pageEnd=1669, url=null, language=null, rfNumber=[28], rfOrder=28, authorNames=STEPHENS CM, ZWEIGER G, SHAPIRO L, journalName=Journal of Bacteriology, refType=null, unstructuredReference=STEPHENS CM, ZWEIGER G, SHAPIRO L. Coordinate cell cycle control of a Caulobacter DNA methyltransferase and the flagellar genetic hierarchy[J]. Journal of Bacteriology, 1995, 177(7): 1662-1669., articleTitle=Coordinate cell cycle control of a Caulobacter DNA methyltransferase and the flagellar genetic hierarchy, refAbstract=null), Reference(id=1226964059560526811, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2024, volume=206, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[29], rfOrder=29, authorNames=CAMPBELL M, BARTON IS, ROOP RM 2nd, CHIEN P, journalName=Journal of Bacteriology, refType=null, unstructuredReference=CAMPBELL M, BARTON IS, ROOP RM 2nd, CHIEN P. Comparison of CcrM-dependent methylation in Caulobacter crescentus and Brucella abortus by nanopore sequencing[J]. Journal of Bacteriology, 2024, 206: e00083-24., articleTitle=Comparison of CcrM-dependent methylation in Caulobacter crescentus and Brucella abortus by nanopore sequencing, refAbstract=null), Reference(id=1226964059694744547, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2019, volume=116, issue=31, pageStart=15661, pageEnd=15670, url=null, language=null, rfNumber=[30], rfOrder=30, authorNames=ZHOU XF, WANG JR, HERRMANN J, MOERNER WE, SHAPIRO L, journalName=Proceedings of the National Academy of Sciences of the United States of America, refType=null, unstructuredReference=ZHOU XF, WANG JR, HERRMANN J, MOERNER WE, SHAPIRO L. Asymmetric division yields progeny cells with distinct modes of regulating cell cycle-dependent chromosome methylation[J]. Proceedings of the National Academy of Sciences of the United States of America, 2019, 116(31): 15661-15670., articleTitle=Asymmetric division yields progeny cells with distinct modes of regulating cell cycle-dependent chromosome methylation, refAbstract=null), Reference(id=1226964059816379371, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2018, volume=110, issue=1, pageStart=1, pageEnd=10, url=null, language=null, rfNumber=[31], rfOrder=31, authorNames=MOUAMMINE A, COLLIER J, journalName=Molecular Microbiology, refType=null, unstructuredReference=MOUAMMINE A, COLLIER J. The impact of DNA methylation in Alphaproteobacteria [J]. Molecular Microbiology, 2018, 110(1): 1-10., articleTitle=The impact of DNA methylation in Alphaproteobacteria, refAbstract=null), Reference(id=1226964059929625585, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2016, volume=12, issue=12, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[32], rfOrder=32, authorNames=ARDISSONE S, REDDER P, RUSSO G, FRANDI A, FUMEAUX C, PATRIGNANI A, SCHLAPBACH R, FALQUET L, VIOLLIER PH, journalName=PLoS Genetics, refType=null, unstructuredReference=ARDISSONE S, REDDER P, RUSSO G, FRANDI A, FUMEAUX C, PATRIGNANI A, SCHLAPBACH R, FALQUET L, VIOLLIER PH. Cell cycle constraints and environmental control of local DNA hypomethylation in α-proteobacteria [J]. PLoS Genetics, 2016, 12(12): e1006499., articleTitle=Cell cycle constraints and environmental control of local DNA hypomethylation in α-proteobacteria, refAbstract=null), Reference(id=1226964060038677496, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2007, volume=104, issue=43, pageStart=17111, pageEnd=17116, url=null, language=null, rfNumber=[33], rfOrder=33, authorNames=COLLIER J, McADAMS HH, SHAPIRO L, journalName=Proceedings of the National Academy of Sciences of the United States of America, refType=null, unstructuredReference=COLLIER J, McADAMS HH, SHAPIRO L. A DNA methylation ratchet governs progression through a bacterial cell cycle[J]. Proceedings of the National Academy of Sciences of the United States of America, 2007, 104(43): 17111-17116., articleTitle=A DNA methylation ratchet governs progression through a bacterial cell cycle, refAbstract=null), Reference(id=1226964060172895232, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2024, volume=52, issue=19, pageStart=11519, pageEnd=11535, url=null, language=null, rfNumber=[34], rfOrder=34, authorNames=MARTIN S, FOURNES F, AMBROSINI G, ISELI C, BOJKOWSKA K, MARQUIS J, GUEX N, COLLIER J, journalName=Nucleic Acids Research, refType=null, unstructuredReference=MARTIN S, FOURNES F, AMBROSINI G, ISELI C, BOJKOWSKA K, MARQUIS J, GUEX N, COLLIER J. DNA methylation by CcrM contributes to genome maintenance in the Agrobacterium tumefaciens plant pathogen[J]. Nucleic Acids Research, 2024, 52(19): 11519-11535., articleTitle=DNA methylation by CcrM contributes to genome maintenance in the Agrobacterium tumefaciens plant pathogen, refAbstract=null), Reference(id=1226964060315500546, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2022, volume=7, issue=1, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[35], rfOrder=35, authorNames=DICENZO GC, CANGIOLI L, NICOUD Q, CHENG JHT, BLOW MJ, SHAPIRO N, WOYKE T, BIONDI EG, ALUNNI B, MENGONI A, MERGAERT P, journalName=mSystems, refType=null, unstructuredReference=DICENZO GC, CANGIOLI L, NICOUD Q, CHENG JHT, BLOW MJ, SHAPIRO N, WOYKE T, BIONDI EG, ALUNNI B, MENGONI A, MERGAERT P. DNA methylation in Ensifer species during free-living growth and during nitrogen-fixing symbiosis with Medicago spp.[J]. mSystems, 2022, 7(1): e0109221., articleTitle=DNA methylation in Ensifer species during free-living growth and during nitrogen-fixing symbiosis with Medicago spp., refAbstract=null), Reference(id=1226964060479078411, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2002, volume=56, issue=null, pageStart=625, pageEnd=656, url=null, language=null, rfNumber=[36], rfOrder=36, authorNames=MARCZYNSKI GT, SHAPIRO L, journalName=Annual Review of Microbiology, refType=null, unstructuredReference=MARCZYNSKI GT, SHAPIRO L. Control of chromosome replication in Caulobacter crescentus [J]. Annual Review of Microbiology, 2002, 56: 625-656., articleTitle=Control of chromosome replication in Caulobacter crescentus, refAbstract=null), Reference(id=1226964060592324627, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2015, volume=11, issue=5, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[37], rfOrder=37, authorNames=PINI F, de NISCO NJ, FERRI L, PENTERMAN J, FIORAVANTI A, BRILLI M, MENGONI A, BAZZICALUPO M, VIOLLIER PH, WALKER GC, BIONDI EG, journalName=PLoS Genetics, refType=null, unstructuredReference=PINI F, de NISCO NJ, FERRI L, PENTERMAN J, FIORAVANTI A, BRILLI M, MENGONI A, BAZZICALUPO M, VIOLLIER PH, WALKER GC, BIONDI EG. Cell cycle control by the master regulator CtrA in Sinorhizobium meliloti [J]. PLoS Genetics, 2015, 11(5): e1005232., articleTitle=Cell cycle control by the master regulator CtrA in Sinorhizobium meliloti, refAbstract=null), Reference(id=1226964060713959451, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2013, volume=11, issue=12, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[38], rfOrder=38, authorNames=MURRAY SM, PANIS G, FUMEAUX C, VIOLLIER PH, HOWARD M, journalName=PLoS Biology, refType=null, unstructuredReference=MURRAY SM, PANIS G, FUMEAUX C, VIOLLIER PH, HOWARD M. Computational and genetic reduction of a cell cycle to its simplest, primordial components[J]. PLoS Biology, 2013, 11(12): e1001749., articleTitle=Computational and genetic reduction of a cell cycle to its simplest, primordial components, refAbstract=null), Reference(id=1226964060831399971, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2013, volume=88, issue=1, pageStart=203, pageEnd=218, url=null, language=null, rfNumber=[39], rfOrder=39, authorNames=GONZALEZ D, COLLIER J, journalName=Molecular Microbiology, refType=null, unstructuredReference=GONZALEZ D, COLLIER J. DNA methylation by CcrM activates the transcription of two genes required for the division of Caulobacter crescentus [J]. Molecular Microbiology, 2013, 88(1): 203-218., articleTitle=DNA methylation by CcrM activates the transcription of two genes required for the division of Caulobacter crescentus, refAbstract=null), Reference(id=1226964060986589228, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2021, volume=17, issue=3, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[40], rfOrder=40, authorNames=ADHIKARI S, ERILL I, CURTIS PD, journalName=PLoS Genetics, refType=null, unstructuredReference=ADHIKARI S, ERILL I, CURTIS PD. Transcriptional rewiring of the GcrA/CcrM bacterial epigenetic regulatory system in closely related bacteria[J]. PLoS Genetics, 2021, 17(3): e1009433., articleTitle=Transcriptional rewiring of the GcrA/CcrM bacterial epigenetic regulatory system in closely related bacteria, refAbstract=null), Reference(id=1226964061112418354, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2015, volume=29, issue=11, pageStart=1175, pageEnd=1187, url=null, language=null, rfNumber=[41], rfOrder=41, authorNames=NARAYANAN S, JANAKIRAMAN B, KUMAR L, RADHAKRISHNAN SK, journalName=Genes & Development, refType=null, unstructuredReference=NARAYANAN S, JANAKIRAMAN B, KUMAR L, RADHAKRISHNAN SK. A cell cycle-controlled redox switch regulates the topoisomerase Ⅳ activity[J]. Genes & Development, 2015, 29(11): 1175-1187., articleTitle=A cell cycle-controlled redox switch regulates the topoisomerase Ⅳ activity, refAbstract=null), Reference(id=1226964061271801918, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2015, volume=6, issue=null, pageStart=7005, pageEnd=null, url=null, language=null, rfNumber=[42], rfOrder=42, authorNames=SANSELICIO S, BERGÉ M, THÉRAULAZ L, RADHAKRISHNAN SK, VIOLLIER PH, journalName=Nature Communications, refType=null, unstructuredReference=SANSELICIO S, BERGÉ M, THÉRAULAZ L, RADHAKRISHNAN SK, VIOLLIER PH. Topological control of the Caulobacter cell cycle circuitry by a polarized single-domain PAS protein[J]. Nature Communications, 2015, 6: 7005., articleTitle=Topological control of the Caulobacter cell cycle circuitry by a polarized single-domain PAS protein, refAbstract=null), Reference(id=1226964061401825351, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2020, volume=114, issue=1, pageStart=127, pageEnd=139, url=null, language=null, rfNumber=[43], rfOrder=43, authorNames=MOHAPATRA SS, FIORAVANTI A, VANDAME P, SPRIET C, PINI F, BOMPARD C, BLOSSEY R, VALETTE O, BIONDI EG, journalName=Molecular Microbiology, refType=null, unstructuredReference=MOHAPATRA SS, FIORAVANTI A, VANDAME P, SPRIET C, PINI F, BOMPARD C, BLOSSEY R, VALETTE O, BIONDI EG. Methylation-dependent transcriptional regulation of crescentin gene (creS) by GcrA in Caulobacter crescentus [J]. Molecular Microbiology, 2020, 114(1): 127-139., articleTitle=Methylation-dependent transcriptional regulation of crescentin gene (creS) by GcrA in Caulobacter crescentus, refAbstract=null), Reference(id=1226964061523460173, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2011, volume=21, issue=13, pageStart=1092, pageEnd=1101, url=null, language=null, rfNumber=[44], rfOrder=44, authorNames=JONAS K, CHEN YE, LAUB MT, journalName=Current Biology, refType=null, unstructuredReference=JONAS K, CHEN YE, LAUB MT. Modularity of the bacterial cell cycle enables independent spatial and temporal control of DNA replication[J]. Current Biology, 2011, 21(13): 1092-1101., articleTitle=Modularity of the bacterial cell cycle enables independent spatial and temporal control of DNA replication, refAbstract=null), Reference(id=1226964061640900690, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2004, volume=304, issue=5673, pageStart=983, pageEnd=987, url=null, language=null, rfNumber=[45], rfOrder=45, authorNames=HOLTZENDORFF J, HUNG D, BRENDE P, REISENAUER A, VIOLLIER PH, McADAMS HH, SHAPIRO L, journalName=Science, refType=null, unstructuredReference=HOLTZENDORFF J, HUNG D, BRENDE P, REISENAUER A, VIOLLIER PH, McADAMS HH, SHAPIRO L. Oscillating global regulators control the genetic circuit driving a bacterial cell cycle[J]. Science, 2004, 304(5673): 983-987., articleTitle=Oscillating global regulators control the genetic circuit driving a bacterial cell cycle, refAbstract=null), Reference(id=1226964061728981077, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2013, volume=110, issue=48, pageStart=E4658, pageEnd=E4667, url=null, language=null, rfNumber=[46], rfOrder=46, authorNames=KOZDON JB, MELFI MD, LUONG K, CLARK TA, BOITANO M, WANG S, ZHOU B, GONZALEZ D, COLLIER J, TURNER SW, KORLACH J, SHAPIRO L, McADAMS HH, journalName=Proceedings of the National Academy of Sciences of the United States of America, refType=null, unstructuredReference=KOZDON JB, MELFI MD, LUONG K, CLARK TA, BOITANO M, WANG S, ZHOU B, GONZALEZ D, COLLIER J, TURNER SW, KORLACH J, SHAPIRO L, McADAMS HH. Global methylation state at base-pair resolution of the Caulobacter genome throughout the cell cycle[J]. Proceedings of the National Academy of Sciences of the United States of America, 2013, 110(48): E4658-E4667., articleTitle=Global methylation state at base-pair resolution of the Caulobacter genome throughout the cell cycle, refAbstract=null), Reference(id=1226964061892558941, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2022, volume=18, issue=1, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[47], rfOrder=47, authorNames=XU CR, HOLLIS H, DAI M, YAO XY, WATSON LT, CAO Y, CHEN MH, journalName=PLoS Computational Biology, refType=null, unstructuredReference=XU CR, HOLLIS H, DAI M, YAO XY, WATSON LT, CAO Y, CHEN MH. Modeling the temporal dynamics of master regulators and CtrA proteolysis in Caulobacter crescentus cell cycle[J]. PLoS Computational Biology, 2022, 18(1): e1009847., articleTitle=Modeling the temporal dynamics of master regulators and CtrA proteolysis in Caulobacter crescentus cell cycle, refAbstract=null), Reference(id=1226964062018388068, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2002, volume=21, issue=18, pageStart=4969, pageEnd=4977, url=null, language=null, rfNumber=[48], rfOrder=48, authorNames=REISENAUER A, SHAPIRO L, journalName=EMBO Journal, refType=null, unstructuredReference=REISENAUER A, SHAPIRO L. DNA methylation affects the cell cycle transcription of the CtrA global regulator in Caulobacter [J]. EMBO Journal, 2002, 21(18): 4969-4977., articleTitle=DNA methylation affects the cell cycle transcription of the CtrA global regulator in Caulobacter, refAbstract=null), Reference(id=1226964062173577324, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2015, volume=29, issue=21, pageStart=2272, pageEnd=2286, url=null, language=null, rfNumber=[49], rfOrder=49, authorNames=HAAKONSEN DL, YUAN AH, LAUB MT, journalName=Genes & Development, refType=null, unstructuredReference=HAAKONSEN DL, YUAN AH, LAUB MT. The bacterial cell cycle regulator GcrA is a σ70 cofactor that drives gene expression from a subset of methylated promoters[J]. Genes & Development, 2015, 29(21): 2272-2286., articleTitle=The bacterial cell cycle regulator GcrA is a σ70 cofactor that drives gene expression from a subset of methylated promoters, refAbstract=null), Reference(id=1226964062303600756, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2015, volume=6, issue=4, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[50], rfOrder=50, authorNames=GONZALEZ D, COLLIER J, journalName=mBio, refType=null, unstructuredReference=GONZALEZ D, COLLIER J. Genomic adaptations to the loss of a conserved bacterial DNA methyltransferase[J]. mBio, 2015, 6(4): e00952., articleTitle=Genomic adaptations to the loss of a conserved bacterial DNA methyltransferase, refAbstract=null), Reference(id=1226964062429429883, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2018, volume=46, issue=6, pageStart=3245, pageEnd=3256, url=null, language=null, rfNumber=[51], rfOrder=51, authorNames=WU XX, HAAKONSEN DL, SANDERLIN AG, LIU YJ, SHEN LQ, ZHUANG NN, LAUB MT, ZHANG Y, journalName=Nucleic Acids Research, refType=null, unstructuredReference=WU XX, HAAKONSEN DL, SANDERLIN AG, LIU YJ, SHEN LQ, ZHUANG NN, LAUB MT, ZHANG Y. Structural insights into the unique mechanism of transcription activation by Caulobacter crescentus GcrA[J]. Nucleic Acids Research, 2018, 46(6): 3245-3256., articleTitle=Structural insights into the unique mechanism of transcription activation by Caulobacter crescentus GcrA, refAbstract=null), Reference(id=1226964062534287489, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2023, volume=51, issue=4, pageStart=1960, pageEnd=1970, url=null, language=null, rfNumber=[52], rfOrder=52, authorNames=WU XX, YU CZ, MU WH, GU ZX, FENG Y, ZHANG Y, journalName=Nucleic Acids Research, refType=null, unstructuredReference=WU XX, YU CZ, MU WH, GU ZX, FENG Y, ZHANG Y. The structural mechanism for transcription activation by Caulobacter crescentus GcrA[J]. Nucleic Acids Research, 2023, 51(4): 1960-1970., articleTitle=The structural mechanism for transcription activation by Caulobacter crescentus GcrA, refAbstract=null), Reference(id=1226964062660116614, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2018, volume=45, issue=10, pageStart=1026, pageEnd=1038, url=null, language=null, rfNumber=[53], rfOrder=53, authorNames=张文婷, 姚玉峰, journalName=生物化学与生物物理进展, refType=null, unstructuredReference=张文婷, 姚玉峰. 细菌DNA甲基化研究进展[J]. 生物化学与生物物理进展, 2018, 45(10): 1026-1038., articleTitle=细菌DNA甲基化研究进展, refAbstract=null), Reference(id=1226964064081985679, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2018, volume=45, issue=10, pageStart=1026, pageEnd=1038, url=null, language=null, rfNumber=[53], rfOrder=54, authorNames=ZHANG WT, YAO YF, journalName=Progress in Biochemistry and Biophysics, refType=null, unstructuredReference=ZHANG WT, YAO YF. Research progress on bacterial DNA methylation[J]. Progress in Biochemistry and Biophysics, 2018, 45(10): 1026-1038 (in Chinese)., articleTitle=null, refAbstract=null), Reference(id=1226964064174260373, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=2012, volume=134, issue=21, pageStart=8896, pageEnd=8901, url=null, language=null, rfNumber=[54], rfOrder=55, authorNames=LI DY, DELANEY JC, PAGE CM, YANG XD, CHEN AS, WONG C, DRENNAN CL, ESSIGMANN JM, journalName=Journal of the American Chemical Society, refType=null, unstructuredReference=LI DY, DELANEY JC, PAGE CM, YANG XD, CHEN AS, WONG C, DRENNAN CL, ESSIGMANN JM. Exocyclic carbons adjacent to the N6 of adenine are targets for oxidation by the Escherichia coli adaptive response protein AlkB[J]. Journal of the American Chemical Society, 2012, 134(21): 8896-8901., articleTitle=Exocyclic carbons adjacent to the N6 of adenine are targets for oxidation by the Escherichia coli adaptive response protein AlkB, refAbstract=null), Reference(id=1226964064295895194, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, doi=null, pmid=null, pmcid=null, year=1997, volume=179, issue=10, pageStart=3139, pageEnd=3145, url=null, language=null, rfNumber=[55], rfOrder=56, authorNames=COLOMBI D, GOMES SL, journalName=Journal of Bacteriology, refType=null, unstructuredReference=COLOMBI D, GOMES SL. An alkB gene homolog is differentially transcribed during the Caulobacter crescentus cell cycle[J]. Journal of Bacteriology, 1997, 179(10): 3139-3145., articleTitle=An alkB gene homolog is differentially transcribed during the Caulobacter crescentus cell cycle, refAbstract=null)], funds=[Fund(id=1226964053319402201, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, awardId=32300151, language=EN, fundingSource=National Natural Science Foundation of China(32300151), fundOrder=null, country=null), Fund(id=1226964053411676897, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, awardId=32300151, language=CN, fundingSource=国家自然科学基金(32300151), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1226964048282042771, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, xref=null, ext=[AuthorCompanyExt(id=1226964048412066199, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, companyId=1226964048282042771, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=College of Bioscience and Biotechnology, Yangzhou University, Yangzhou, Jiangsu, China), AuthorCompanyExt(id=1226964048420454810, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, companyId=1226964048282042771, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=扬州大学 生物科学与技术学院，江苏 扬州)])], figs=[ArticleFig(id=1226964052308574836, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=EN, label=Figure 1, caption=Sequence recognition and catalytic mechanism of CcrM. A: Schematic diagram of CcrM catalyzing the formation of G(6mA)NTC from GANTC (The adenine residue is presented in structural formula, and “dR” represents phosphodeoxyribose); B: Structure of the CcrM monomer (PDB: 6PBD); C: Mechanism of CcrM binding to and catalyzing DNA (PDB: 6PBD)[17]. The dsDNA is shown in red. The blue and green colors indicate the two monomers of the CcrM dimer, respectively. The blue monomer mainly functions in recognition and methyl-transfer of target site, while the green monomer is mainly involved in non-specific binding to the complementary strand of the target site. The content within the dashed box on the right side shows a schematic diagram of the mechanism by which CcrM recognizes and catalyzes GANTC. Several motifs used for recognizing the GANTC motif are all marked in a modular form. The modules with a blue background are from the blue monomer in figure C, and the module with a green background is from the green monomer of the dimer in figure C. The green arcs represent the motifs used for non-specific recognition of the complementary strand of the target site., figureFileSmall=psr7GmLPBsQ+BjiebOo4Nw==, figureFileBig=tQZtAZpWKZIiRB32wgyl6w==, tableContent=null), ArticleFig(id=1226964052413432450, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=CN, label=图1, caption=CcrM的序列识别与催化机制。A：CcrM催化GANTC生成G(6mA)NTC的示意图(腺嘌呤核苷酸以结构式呈现，dR表示磷酸脱氧核糖)；B：CcrM单体结构(PDB: 6PBD)；C：CcrM结合和催化DNA的机制(PDB: 6PBD)[17]。红色表示双链DNA (double-stranded DNA, dsDNA)，蓝色和绿色分别指示CcrM二聚体的2个单体，其中蓝色单体主要负责靶位点的识别和催化甲基转移，绿色单体主要负责靶位点互补链的非特异性结合，右侧虚线框内容展示了CcrM识别和催化GANTC的机制，用于识别GANTC基序的几个模体均以模块形式标注，其中蓝色底色的模块来自图C左侧二聚体中的蓝色单体，绿色底色的模块来自图C左侧二聚体中的绿色单体，绿色弧线表示用于非特异性识别靶位点互补链的模体。, figureFileSmall=psr7GmLPBsQ+BjiebOo4Nw==, figureFileBig=tQZtAZpWKZIiRB32wgyl6w==, tableContent=null), ArticleFig(id=1226964052539261579, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=EN, label=Figure 2, caption=Regulatory mechanism of CcrM. The grey arrows represent the genes to be transcribed, the black arrow indicates transcription, the red arrows indicate activation, the blue “T”-shaped lines indicate inhibition, and “CH3” represents the methyl group transferred to the GANTC motif catalyzed by CcrM. Each rectangle contains the biochemical processes regulated by CcrM, and only representative genes and biochemical processes are included., figureFileSmall=YIIep/G3rwzmBrvj6wnYbQ==, figureFileBig=W+G2g59crKYgXJo6YpBRkA==, tableContent=null), ArticleFig(id=1226964052635730583, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=CN, label=图2, caption=CcrM的调控机制。灰色箭头表示待转录的基因，黑色箭头表示转录，红色箭头表示激活，蓝色“T”形线表示抑制，“CH3”表示由CcrM催化转移到GANTC基序上的甲基，各矩形包含受CcrM调控的生理过程，仅选取了代表性基因和生理过程。, figureFileSmall=YIIep/G3rwzmBrvj6wnYbQ==, figureFileBig=W+G2g59crKYgXJo6YpBRkA==, tableContent=null), ArticleFig(id=1226964052728005280, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=EN, label=Figure 3, caption=Mechanism of GcrA activating gene expression. The blue structure represents the schematic model of the DNA-binding domain (DBD) of GcrA. The purple structure represents the schematic model of the σ-factor interaction domain (SID) of GcrA. The two are connected by a grey linker (schematic model). The yellow structure is the schematic model of the σ70 factor, and the light-grey structure represents the schematic model of RNA polymerase. GcrA recognizes the G(6mA)NTC motif (rectangle with red background) through the DBD, and then interacts with the σ70 factor through the SID to activate transcription. The model was summarized and drawn based on the research findings of Gonzalez et al. and Wu et al.[50-52]., figureFileSmall=YQ+/s3ZVmVKX6DAVGMvejw==, figureFileBig=SuVr5P84UVZnu0plEFas/A==, tableContent=null), ArticleFig(id=1226964052832862890, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=CN, label=图3, caption=GcrA激活基因表达的机制。蓝色结构表示GcrA的DNA结合结构域(DBD)示意模型，紫色结构表示GcrA的σ因子互作结构域(SID)示意模型，两者由灰色的linker (示意模型)连接，黄色结构为σ70因子示意模型，浅灰色结构表示RNA聚合酶示意模型。GcrA通过DBD识别G(6mA)NTC (红底色矩形)，再通过SID与σ70因子互作，激活转录。模型基于Gonzalez等和Wu等[50-52]的研究结果总结绘制。, figureFileSmall=YQ+/s3ZVmVKX6DAVGMvejw==, figureFileBig=SuVr5P84UVZnu0plEFas/A==, tableContent=null), ArticleFig(id=1226964052962886328, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=EN, label=Table 1, caption=

Cell cycle-related genes regulated by CcrM

, figureFileSmall=null, figureFileBig=null, tableContent=
GeneFunctionActivated (A)/repressed (R) by CcrMActivated by GcrAValidated strainReferences
dnaADNA replicationControversialRepressed by GcrACv[8,33,45-47]
ligADNA replicationA-Cv[8]
gyrADNA replicationA-Cv, Bm[8,40,46]
nrdADNA replicationA-Cv[8]
recJDNA replicationA-Cv[8]
ssbDNA replicationA-Cv[8]
thyADNA replicationA-Cv[8]
repABCDNA replicationAYesAr[8]
nstAChromosome segregationAYesCv[41]
parEChromosome segregationA-Cv, Bm[8]
ftsZCell divisionAYesCv, Bm, Ar[8,38-40]
ftsNCell divisionAYesCv, Bm[4,8,38,40]
ftsWCell divisionR-Cv[8]
ftsECell divisionA-Cv[8]
mipZCell divisionAYesCv, Bm[4,38,40,45]
pleCPolarityAYesCv[4,43]
popZPolarityR-Cv[8]
podJPolarity regulatorAYesCv[4,38,45]
tipFPolarity regulatorAYesCv[4]
flaYMotilityAYesCv[4]
staRStalk biogenesisA-Cv[8,46]
creSCell shapeAYesCv[8,43]
ctrACell cycle regulatorRYesCv, Ar, Bm, Em[4,8,38,45,48]
mopJpleiotropic regulatorAYesCv[42]
), ArticleFig(id=1226964053097104072, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956549617267460, language=CN, label=表1, caption=

CcrM调控的细胞周期相关基因

, figureFileSmall=null, figureFileBig=null, tableContent=
GeneFunctionActivated (A)/repressed (R) by CcrMActivated by GcrAValidated strainReferences
dnaADNA replicationControversialRepressed by GcrACv[8,33,45-47]
ligADNA replicationA-Cv[8]
gyrADNA replicationA-Cv, Bm[8,40,46]
nrdADNA replicationA-Cv[8]
recJDNA replicationA-Cv[8]
ssbDNA replicationA-Cv[8]
thyADNA replicationA-Cv[8]
repABCDNA replicationAYesAr[8]
nstAChromosome segregationAYesCv[41]
parEChromosome segregationA-Cv, Bm[8]
ftsZCell divisionAYesCv, Bm, Ar[8,38-40]
ftsNCell divisionAYesCv, Bm[4,8,38,40]
ftsWCell divisionR-Cv[8]
ftsECell divisionA-Cv[8]
mipZCell divisionAYesCv, Bm[4,38,40,45]
pleCPolarityAYesCv[4,43]
popZPolarityR-Cv[8]
podJPolarity regulatorAYesCv[4,38,45]
tipFPolarity regulatorAYesCv[4]
flaYMotilityAYesCv[4]
staRStalk biogenesisA-Cv[8,46]
creSCell shapeAYesCv[8,43]
ctrACell cycle regulatorRYesCv, Ar, Bm, Em[4,8,38,45,48]
mopJpleiotropic regulatorAYesCv[42]
)], attaches=null, journal=Journal(id=1192105720683257860, delFlag=0, nameCn=微生物学报, nameEn=Acta Microbiologica Sinica, nameHistory1=null, nameHistory2=null, issn=0001-6209, eissn=null, cn=11-1995/Q, coden=null, periodic=0, language=CN, oaType=null, ccby=null, superviseOffice=null, ownerOffice=null, pubOffice=null, editorOffice=null, officeType=null, aims=null, clcCode=null, officeProv=null, officeCity=null, officeAddr=null, officeZip=null, officeEmail=null, officePhone=null, editDirector=null, officeDirector=null, officeDirectorPhone=null, officeStaffNum=null, officeEmpNum=null, coverPicUrl=tNA7JigLZj/rxynSmzKgDQ==, journalPrice=null, startedYear=null, abbrevIsoEn=null, journalRemark=null, publicationField=null, createdTime=1762149752067, updatedTime=1762150746905, createdBy=18614031015, updatedBy=13701087609, firstLetterCn=A, firstLetterEn=A, subjectCode=Life Sciences, subjectName=Life Sciences, subjectCodeEn=Life Sciences, subjectNameEn=null, picCn=tNA7JigLZj/rxynSmzKgDQ==, picEn=R/d5eSUu8/o5mAGWCF3M5Q==, jcr=null, cjcr=null, exts=[JournalExt(id=1192109893441171829, language=CN, name=微生物学报, nameHistory1=null, nameHistory2=null, managedBy=, sponsoredBy=, publishedBy=, editorOffice=, officeProv=null, officeCity=null, officeAddr=, officeZip=, editDirector=, officeDirector=null, officePhone=null, coverPicUrl=null, journalRemark=, submitArticleUrl=null, websiteUrl=, createdTime=1762150746928, updatedTime=1762150746928, createdBy=13701087609, updatedBy=13701087609, submissionGuidelinesUrl=, submissionAuthorUrl=https://actamicro.ijournals.cn/actamicrocn/author/login, submissionEditorUrl=https://actamicro.ijournals.cn/actamicrocn/editor/login, submissionReviewUrl=https://actamicro.ijournals.cn/actamicrocn/reviewer/login, submissionCeEditorUrl=, submissionAeEditorUrl=, option={"copyright":""}), JournalExt(id=1192109893512474998, language=EN, name=Acta Microbiologica Sinica, nameHistory1=null, nameHistory2=null, managedBy=, sponsoredBy=, publishedBy=, editorOffice=, officeProv=null, officeCity=null, officeAddr=, officeZip=, editDirector=, officeDirector=null, officePhone=null, coverPicUrl=null, journalRemark=, submitArticleUrl=null, websiteUrl=, createdTime=1762150746944, updatedTime=1762150746944, createdBy=13701087609, updatedBy=13701087609, submissionGuidelinesUrl=, submissionAuthorUrl=https://actamicro.ijournals.cn/actamicrocn/author/login, submissionEditorUrl=https://actamicro.ijournals.cn/actamicrocn/editor/login, submissionReviewUrl=https://actamicro.ijournals.cn/actamicrocn/reviewer/login, submissionCeEditorUrl=, submissionAeEditorUrl=, option={"copyright":""})], databaseList=null, tenantJournalId=1192105938417971205, websiteList=[Website(id=1192106105867223981, webName=null, webTitle=null, webDomain=null, webCopyrigh=null, webIpcNo=null, seoTitle=null, seoKeywords=null, seoDescription=null, tenantJournalId=null, journalId=1192105938417971205, journalNameCn=null, journalNameEn=null, grayFlag=null, tenantId=1146029695717560320, platformId=null, journalGroupId=null, journalGroupNameCn=null, journalGroupNameEn=null, type=1, domain=https://castjournals.cast.org.cn/joweb/wswxb/CN, language=CN, createTime=1762149843899, createBy=18614031015, updateTime=1762149888800, updateBy=18614031015, name=微生物学报-中文, tplId=1146099689490845704, title=微生物学报, delFlag=0, indexPage=/home, props=[WebsiteProps(id=1192107120863626198, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106105867223981, code=articleTextType, value=kx, createTime=1762150085893, updateTime=1762150085893, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107120834266067, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106105867223981, code=banner, value=null, createTime=1762150085886, updateTime=1762150085886, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107120892986329, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106105867223981, code=grayFlag, value=0, createTime=1762150085900, updateTime=1762150085900, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107120825877458, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106105867223981, code=logo, value=https://castjournals.cast.org.cn/joweb/wswxb/CN/file/pic?fileId=FOz4Ks7dC79FYnCEBIlMdw==, createTime=1762150085884, updateTime=1762150085884, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107120905569243, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106105867223981, code=minRunFlag, value=0, createTime=1762150085903, updateTime=1762150085903, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107120846848981, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106105867223981, code=picServerUrl, value=https://castjournals.cast.org.cn/joweb/wswxb/CN/file/pic, createTime=1762150085889, updateTime=1762150085889, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107120897180634, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106105867223981, code=silenceFlag, value=0, createTime=1762150085901, updateTime=1762150085901, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107120842654676, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106105867223981, code=staticResourcePath, value=https://castjournals.cast.org.cn/joweb/cast_kjdb_cn_619/, createTime=1762150085888, updateTime=1762150085888, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107120872014807, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106105867223981, code=themeColor, value=null, createTime=1762150085895, updateTime=1762150085895, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107120880403416, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106105867223981, code=themeStyle, value=null, createTime=1762150085897, updateTime=1762150085897, creator=18614031015, updator=18614031015)]), Website(id=1192106106018218929, webName=null, webTitle=null, webDomain=null, webCopyrigh=null, webIpcNo=null, seoTitle=null, seoKeywords=null, seoDescription=null, tenantJournalId=null, journalId=1192105938417971205, journalNameCn=null, journalNameEn=null, grayFlag=null, tenantId=1146029695717560320, platformId=null, journalGroupId=null, journalGroupNameCn=null, journalGroupNameEn=null, type=1, domain=https://castjournals.cast.org.cn/joweb/wswxb/EN, language=EN, createTime=1762149843935, createBy=18614031015, updateTime=1762149925242, updateBy=18614031015, name=微生物学报-英文, tplId=1146101810881728533, title=Acta Microbiologica Sinica, delFlag=0, indexPage=/home, props=[WebsiteProps(id=1192107140455220192, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106106018218929, code=articleTextType, value=kx, createTime=1762150090564, updateTime=1762150090564, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107140434248669, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106106018218929, code=banner, value=null, createTime=1762150090559, updateTime=1762150090559, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107140476191715, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106106018218929, code=grayFlag, value=0, createTime=1762150090569, updateTime=1762150090569, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107140425860060, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106106018218929, code=logo, value=https://castjournals.cast.org.cn/joweb/wswxb/EN/file/pic?fileId=FOz4Ks7dC79FYnCEBIlMdw==, createTime=1762150090557, updateTime=1762150090557, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107140484580325, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106106018218929, code=minRunFlag, value=0, createTime=1762150090571, updateTime=1762150090571, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107140451025887, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106106018218929, code=picServerUrl, value=https://castjournals.cast.org.cn/joweb/wswxb/EN/file/pic, createTime=1762150090563, updateTime=1762150090563, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107140480386020, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106106018218929, code=silenceFlag, value=0, createTime=1762150090570, updateTime=1762150090570, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107140442637278, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106106018218929, code=staticResourcePath, value=https://castjournals.cast.org.cn/joweb/cast_kjdb_en_623/, createTime=1762150090561, updateTime=1762150090561, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107140463608801, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106106018218929, code=themeColor, value=null, createTime=1762150090566, updateTime=1762150090566, creator=18614031015, updator=18614031015), WebsiteProps(id=1192107140467803106, tenantId=1146029695717560320, journalId=null, journalGroupId=null, siteId=1192106106018218929, code=themeStyle, value=null, createTime=1762150090567, updateTime=1762150090567, creator=18614031015, updator=18614031015)])], journalTitle=微生物学报, weixinUrl=null, journalUrl=https://actamicro.ijournals.cn, iacademicId=null, status=1, seqNo=null, journalTitleEn=Acta Microbiologica Sinica, journalPhotoCn=tNA7JigLZj/rxynSmzKgDQ==, journalPhotoEn=R/d5eSUu8/o5mAGWCF3M5Q==, journalFirstLetter=A, journalRecommend=null, journalNew=null, journalCollection=null, jcrJf=null, cjcrJf=null, jcrJfStr=null, cjcrJfStr=null, submissionFirstDecision=null, sciSubjectClassification=null, casSubjectClassification=null, citeScore=null, totalCitationFrequency=null, icpCode=null, psCode=null, advertisingLicenseCode=null, copyrightInformation=null, country=null, option=, provinceCode=null, provinceName=null, collectFlag=false), detailUrlCn=https://castjournals.cast.org.cn/joweb/wswxb/CN/10.13343/j.cnki.wsxb.20250131, detailUrlEn=https://castjournals.cast.org.cn/joweb/wswxb/EN/10.13343/j.cnki.wsxb.20250131, pdfUrlCn=https://castjournals.cast.org.cn/joweb/wswxb/CN/PDF/10.13343/j.cnki.wsxb.20250131, pdfUrlEn=https://castjournals.cast.org.cn/joweb/wswxb/EN/PDF/10.13343/j.cnki.wsxb.20250131, aliStartDate=null, aliEndDate=null, collectionFlag=false, citedCount=null, citedUrl=null, reference=null)
收藏切换
参与表观遗传调控的细菌DNA甲基转移酶CcrM的研究进展
收藏切换
PDF下载
王浩 , 王尔雅 , 郭敏亮
微生物学报 | 综述 2025,65(9): 3848-3858
收起
收藏切换
微生物学报 | 综述 2025, 65(9): 3848-3858
参与表观遗传调控的细菌DNA甲基转移酶CcrM的研究进展
全屏
王浩, 王尔雅, 郭敏亮
作者信息
  • 扬州大学 生物科学与技术学院，江苏 扬州
Research progress of bacterial DNA methyltransferase CcrM involved in epigenetic regulation
Hao WANG, Erya WANG, Minliang GUO
Affiliations
  • College of Bioscience and Biotechnology, Yangzhou University, Yangzhou, Jiangsu, China
出版时间: 2025-09-04 doi: 10.13343/j.cnki.wsxb.20250131
文章导航
收藏切换
摘要
收起

DNA甲基化是细菌进行表观遗传调控的重要方式，α变形菌利用受细胞周期调控的DNA甲基转移酶(cell cycle-regulated DNA methyltransferase, CcrM)对DNA进行甲基化。CcrM不含限制性内切酶功能单元，属于孤儿甲基转移酶。CcrM通过对序列中腺嘌呤进行甲基化影响DNA与蛋白质的相互作用，从而调节大量基因的表达，对α变形菌细胞周期等过程的调控至关重要。本文综述了CcrM的功能、结构及其表观调控机制，阐明了CcrM对DNA的识别、催化及活性调控的机理，总结了细胞周期全局性调控因子(global cell-cycle regulator, GcrA)利用CcrM的甲基化信号调节基因表达的机制，并展望了CcrM未来的潜在研究方向，为进一步深入研究细菌表观遗传调控机制提供了参考。

受细胞周期调控的DNA甲基转移酶(CcrM)  /  细菌DNA甲基化  /  表观遗传调控  /  细胞周期全局性调控因子(GcrA)

DNA methylation is an important way of epigenetic regulation in bacteria. Alphaproteobacteria methylate DNA by using the cell cycle-regulated DNA methyltransferase (CcrM). CcrM does not contain a functional unit of restriction endonuclease, thus belonging to an orphan methyltransferase. By methylating adenine in DNA sequences, CcrM influences the interaction between DNA and proteins, regulates the expression of numerous genes, and is crucial for the regulation of processes such as the cell cycle of Alphaproteobacteria. We reviewed the function, structure, and epigenetic regulation of CcrM, clarified the mechanisms of CcrM in DNA recognition, catalysis, and activity regulation, summarized the mechanism by which the global cell-cycle regulator (GcrA) utilizes the methylation signals produced by CcrM to regulate gene expression, and provided an outlook on the potential future research directions of CcrM, providing a reference for further in-depth study of the epigenetic regulation mechanisms in bacteria.

CcrM  /  bacterial DNA methylation  /  epigenetic regulation  /  GcrA
王浩, 王尔雅, 郭敏亮. 参与表观遗传调控的细菌DNA甲基转移酶CcrM的研究进展. 微生物学报, 2025 , 65 (9) : 3848 -3858 . DOI: 10.13343/j.cnki.wsxb.20250131
Hao WANG, Erya WANG, Minliang GUO. Research progress of bacterial DNA methyltransferase CcrM involved in epigenetic regulation[J]. Acta Microbiologica Sinica, 2025 , 65 (9) : 3848 -3858 . DOI: 10.13343/j.cnki.wsxb.20250131
正文
收起
在细菌中DNA甲基化广泛存在，调控着复杂的生命进程。细菌DNA甲基化系统分为2类，一类是含有限制性内切酶的限制与修饰系统(restriction and modification systems，R-M系统)，R-M系统包含2种能够识别同一特定序列的酶，即限制性内切酶和甲基转移酶(methyltransferase, MTase)；另一类是不含限制性内切酶的孤儿甲基转移酶[1]。这2类甲基化系统的功能完全不同。R-M系统的主要功能是防御外源DNA入侵，该系统通过甲基转移酶对内源DNA的特定位点进行甲基化修饰，从而使相应的限制性内切酶无法识别和切割内源DNA；而外源DNA由于缺乏对应位点的甲基化修饰，会被限制性内切酶水解[2]。孤儿甲基转移酶不含限制性内切酶，不能像R-M系统那样发挥防御外源DNA的作用，通常通过甲基化修饰改变蛋白质与特定序列的结合，从而调控细菌的生理过程[3]。
在α变形菌中研究最为深入的孤儿甲基转移酶是受细胞周期调控的DNA甲基转移酶(cell cycle-regulated DNA methyltransferase, CcrM)。α变形菌中主要的DNA甲基化位点为腺嘌呤6号位、胞嘧啶4号位以及胞嘧啶5号位[3]。CcrM催化的甲基化位点与I型R-M系统相同，暗示CcrM可能源于I型R-M系统丢失限制性内切酶后进化而来。功能上，CcrM完全独立于R-M系统，其催化的腺嘌呤6号位甲基化是α变形菌中最主要的表观遗传调控信号[3]，细胞周期全局性调控因子(global cell-cycle regulator, GcrA)正是通过识别该信号来控制基因表达[4]。此外，α变形菌中还存在另一种孤儿甲基转移酶，即胞嘧啶甲基转移酶。目前此类酶的功能尚不清楚，与CcrM不同，该酶对于细菌生存多为非必需[5-6]。
CcrM在弧形柄杆菌(Caulobacter vibrioides)中首次被发现[7]，在α变形菌中高度保守[5]。目前仅在立克次氏体目(Rickettsiales)和磁球菌目(Magnetococcales)中未发现其同源基因[8-10]。CcrM对于α变形菌大多是必需的，缺失CcrM会导致细菌死亡[9,11-12]。CcrM对α变形菌的细胞周期调控至关重要。CcrM通过甲基化调控多个关键的细胞周期因子的表达，进而调控细胞周期的有序进行[5]。在马耳他布鲁氏菌(Brucella melitensis)和放射形农杆菌(Agrobacterium radiobacter)中，CcrM还影响细菌的毒力[12-13]。
细菌的表观遗传调控是细菌基因表达调控的重要方式，CcrM所催化的腺嘌呤甲基化是α变形菌的主要表观遗传调控形式，其对细菌细胞周期等过程的调控十分关键。因此本文对CcrM在细菌表观遗传调控中的作用机制进行综述，阐述CcrM的功能、结构及调控机制，以期为进一步深入研究细菌表观遗传调控机制提供参考。
1 CcrM催化特定位点的甲基化
收起
CcrM属于DNA甲基转移酶β家族，与流感嗜血杆菌(Haemophilus influenzae)的Hinf Ⅰ甲基转移酶具有同源性。其催化单元负责识别并结合底物S-腺苷甲硫氨酸(S-adenosyl-methionine, SAM)，将甲基从SAM转移到GANTC (其中N代表任意核苷酸残基)基序的腺嘌呤核苷酸残基的6号位上，形成N6-甲基腺嘌呤(N6-methyl-adenine, 6mA) (图1A)[7]。CcrM对GANTC基序的识别具有特异性，单个碱基替换(如5′-AA NTC-3′)，会导致CcrM对其催化活性降低至原有的1/107[14]。CcrM对单链DNA和RNA表现出相似的特异性识别[14]。GANTC基序的侧翼序列也会影响CcrM的酶活性，侧翼序列的差异导致CcrM催化速度在0.6-8.0 min-1之间波动[15]。相较于未甲基化的DNA双链底物，CcrM对半甲基化的DNA双链具有更高的催化活性[16]。这与CcrM在细胞中的功能场景有关，在C. vibrioides中CcrM在复制后期的前分裂细胞中表达活跃；GANTC基序由于DNA复制的半保留特性处于半甲基化状态(只有亲本链处于甲基化状态)，此时CcrM开始表达并对新生链进行甲基化，最终形成全甲基化的DNA双链[7]。
2 CcrM的结构及催化机制
收起
CcrM主要由N端的甲基转移酶结构域和C端的非特异性DNA结合结构域组成。N端的MTase结构域主要由一个开放的“αβα三明治”结构构成，C端的结构域则由多种二级结构元件堆积形成，两者由一段灵活的短肽链连接在一起(图1B)；CcrM以二聚体形式发挥功能，二聚体界面主要由2个单体MTase结构域的部分氨基酸残基构成(图1C)[17]。
靶链和互补链之间的分离是实现甲基化反应的前提之一[18-19]。CcrM二聚体与DNA结合时形成2条碱性通道，分别将靶链和互补链扭曲，使靶链识别区和互补链维持在分离状态(图1C)[17]。R350残基对于链分离过程是必需的，SAM能够稳定双链的分离状态[20]。链分离过程与CcrM催化甲基转移时的构象变化是相互独立的[19]。
CcrM二聚体中2个单体对DNA结合和催化的贡献并不均匀[17]。其中一个单体的MTase结构域负责特异性识别靶序列并催化甲基化反应，而另一个单体的C端结构域则与靶链的互补链的磷酸和糖骨架相互作用，实现对DNA的非特异性结合(图1C)[17-18]。MTase结构域中β-折叠片的连接环提供了绝大部分用于识别GANTC的氨基酸残基。其中，R44识别G；D31/P32/P33/Y34识别A；第3位的N是可变的，结合在L38-L42之间的疏水残基处；H94/R129等残基识别T，L42/P123/F125/K126识别C (图1C)[17]。第2位的A是甲基受体，被DPPY 4个氨基酸残基包围，DPPY是MTase中保守的催化活性模体[21]。DPPY分别与腺嘌呤环的极性基团形成氢键，将腺嘌呤6号位的N原子与SAM的甲基、硫原子紧密排列，从而构成MTase催化甲基转移反应的必要环境[21]。
3 CcrM的活性调控
收起
CcrM活性受细胞周期的动态调控。在C. vibrioides中，CcrM蛋白仅在前分裂细胞中可检测到，并在DNA复制完成后才发挥作用[22]。如果在整个细胞周期中持续合成CcrM，细胞会出现分裂异常并过度启动DNA复制的现象[9,22]，表明CcrM活性的时间控制对细胞周期至关重要。CcrM的活性调控基于严格的转录和转录后调控机制。α变形菌中保守的细胞周期调节因子CtrA能够结合ccrM启动子，激活ccrM的转录(图2)[23-25]。CtrA在C. vibrioides前分裂细胞和游动细胞中含量丰富且处于激活状态，其转录激活活性受自身磷酸化状态的影响[23,26]。CtrA对ccrM的激活还受到small CtrA inhibitory protein (SciP)因子的调控(图2)，SciP通过与CtrA互作阻止其招募RNA聚合酶，从而抑制CtrA对ccrM的转录激活(图2)[27]。值得注意的是，CcrM也能够甲基化自身启动子的GANTC基序，这一修饰最终导致ccrM转录的抑制(图2)[28]，但具体的抑制机制目前尚不清楚。除了转录调控，CcrM还受到转录后调控。CcrM是一种不稳定的蛋白质，能够被Lon蛋白酶降解[22,29-30]，因此Lon蛋白酶可以通过控制CcrM在细胞中的含量实现对CcrM活性的调控(图2)。Lon蛋白酶使得CcrM的蛋白质水平能够随着细胞周期中ccrM转录的波动而变化[31]。此外，MucR作为一种保守的转录调节因子也能够调控CcrM的活性(图2)，MucR通过覆盖在GANTC基序上阻止CcrM与GANTC的结合，从而使CcrM处于无效活性状态[32]。
4 CcrM的调控功能
收起
CcrM在α变形菌中高度保守，对于A. radiobacter、B. melitensis和草木栖剑菌(Ensifer meliloti)等细菌的生存是必需的[9,11-12]，表明CcrM的生理功能对α变形菌来说十分重要。过表达或持续表达ccrM基因会导致GANTC位点在整个细胞周期中始终保持完全甲基化状态，此时细胞通常会积累2条以上的染色体，呈现出异常形态，并丧失适应能力[7,22,33]。CcrM还显著影响细菌的运动性、生物膜形成和毒力[12,34]，在根瘤菌中CcrM与共生固氮有关[35]。
在C. vibrioides中大约10%的基因表达会因ccrM的缺失而发生变化，其中包含大量参与细胞周期调控的基因，这些基因在其启动子区域通常含有一个或多个GANTC基序[8]。C. vibrioides的细胞周期涉及柄细胞和游动细胞的分化，CtrA调控因子在其中发挥重要作用。ctrA启动子的上游调控区域包含2个GANTC基序，该启动子仅在半甲基化时才能激活ctrA转录[33]。由于CcrM在DNA复制阶段不表达，而亲代DNA处于全甲基化状态，因此复制会产生2个半甲基化的子代DNA，CcrM正是在此时被激活表达，控制半甲基化DNA向全甲基化DNA的过渡，进而控制CtrA等调控因子的表达[7,33]。CtrA能够与复制原点结合以保护其半甲基化状态；细胞分裂完成后，在游动细胞中CtrA仍然与复制原点结合，确保复制原点处于半甲基化状态，不会进一步启动复制；然而在柄细胞中，CtrA被蛋白酶水解或失活，复制原点完全甲基化后可在DnaA的作用下再次启动复制[36]。除了在C. vibrioides中，CcrM在A. radiobacter[11]、B. melitensis[24]、R. meliloti[9,37]等α变形菌中通过CtrA调控DNA复制的机制具有保守性。目前来看，CcrM对细胞分化的控制是Caulobacter特异性的。
除了通过CtrA调控DNA的复制和细胞分化，CcrM还影响一系列细胞周期相关基因的表达(图2)[31]。在C. vibrioides中参与DNA复制的ligA、gyrA、nrdA等基因的表达受CcrM水平的影响[8]；FtsZ是细胞分裂调节蛋白，在C. vibrioides、B. melitensis和A. radiobacter中CcrM协同GcrA调控FtsZ的表达[29,38-41]；在C. vibrioides中PopZ和PopJ等细胞极性调节因子受CcrM/GcrA的调控[8,42]，但在B. melitensis中两者的表达并不受CcrM/GcrA调控[40]。在A. radiobacter中CcrM通过GcrA促进repABCCh2 操纵子中基因的表达，调控复制起始和染色体分离[34]；在C. vibrioides中细菌运动性相关因子FlaY、细胞形态相关因子CreS等均受到CcrM的表达调控[8,43]。研究表明CcrM介导的甲基化控制着dnaA的转录[33]。然而之后的研究显示dnaA的表达与ccrM无关[44]。除了调节细胞周期，A. radiobacter的CcrM还通过GcrA调控Ti质粒拷贝和毒力基因表达[13]。受CcrM调控的细胞周期相关基因的具体信息详见表1。
在不同α变形菌中CcrM调控的基因大多集中在DNA复制、细胞形态等方面，可见CcrM在调控细胞周期过程中的保守作用。然而有些受调控的基因也体现了菌种特异性，如popZ和popJ在C. vibrioides中受CcrM调控，而在B. melitensis中则不受其影响。CcrM调控部分菌株特有的致病或共生固氮等过程也体现了CcrM功能的菌种特异性。尽管在调控功能上有差异，但CcrM调控基因表达的机制看起来高度保守。在已验证的受调控基因中大部分都是通过GcrA实现了表达调控(表1)。
5 CcrM/GcrA表观调控系统
收起
GcrA是一种受细胞周期调控的转录调节蛋白，控制数百个基因的表达[41,46,49]。转录组分析表明，CcrM调控的基因与GcrA调控的基因存在显著重叠[8,40]。在α变形菌中，GcrA通常与CcrM共同保守存在[10,38]。ΔgcrA和ΔccrM突变体的表型具有高度相似性，在快速生长条件下均无法进行细胞分裂[38-39,50]。在缺乏CcrM的细胞中，GcrA对染色体区域的亲和力较低[4,49]。染色质免疫共沉淀测序(chromatin immunoprecipitation sequencing, ChIP-Seq)实验证实，GcrA在体内优先与含有GANTC基序的σ70因子依赖型启动子结合[49]。GANTC的甲基化对GcrA与启动子的结合至关重要，GcrA对G(6mA)NTC的亲和力显著高于GANTC[4,13]。在一些测试的启动子(包括mipZ启动子)上，GcrA对半甲基化GANTC基序的亲和力取决于甲基化的DNA链，因此GcrA可能通过结合甲基化的GANTC链不对称地调控某些基因表达[4]。这些研究表明GcrA是一种CcrM相关的表观遗传调节因子，其转录调节活性受其靶启动子中GANTC甲基化状态的影响。
最近的几项研究揭示了GcrA的结构及其调节基因转录的机制。GcrA包含2个主要的结构域：位于N端的DNA结合结构域(DNA-binding domain, DBD)和位于C端的σ因子互作结构域(σ factor-interacting domain, SID) (图3)，DBD负责识别并结合甲基化的GANTC基序，SID则负责与σ70因子结合，调节转录水平[51]。
与DNA结合时，DBD的螺旋-转角-螺旋结构镶嵌到双链DNA的大沟中，并通过氢键、堆积力等相互作用来识别序列中的GANTC基序[51-52]。DBD表面有2个“口袋”，它们参与结合DNA 2条链上的甲基基团。第一个深口袋容纳正链上的甲基基团，第二个浅口袋容纳负链上的甲基基团；DBD也可以结合非甲基化的DNA，其与非甲基化DNA的结合方式与结合甲基化DNA基本相同，只是用于结合甲基基团的2个口袋是空的，这解释了为什么GcrA与甲基化DNA的亲和力高于非甲基化DNA[51]。
GcrA能够通过SID与σ70互作，从而组成一个GcrA-σ⁷⁰-RNAP全酶(图3)，共同在基因组DNA上滑动扫描[50-52]。DBD识别并结合G(6mA)NTC后，便阻止GcrA-σ⁷⁰-RNAP在DNA上继续滑动，保证RNAP处于启动转录的合适位置，从而开启下游基因的转录(图3)[50]。此外，DBD与启动子的结合还能够在一定程度上提高启动子的解链速率，加速转录[52]。
6 总结与展望
收起
CcrM作为α变形菌中保守的孤儿甲基转移酶，对细胞周期等过程起到至关重要的调控作用。CcrM能够识别DNA中的GANTC基序，并对其中腺嘌呤的6号位进行甲基化。通过这种甲基化实现对众多基因的表观遗传调控。CcrM的活性呈周期性变化，转录和转录后调控的多种组合实现了细菌对CcrM活性的时间控制。GcrA是CcrM实现基因表达调控的重要因子，其DNA结合结构域能够特异性识别甲基化的GANTC基序，σ因子互作结构域则通过与σ70因子互作，激活下游基因的表达。
在γ变形菌中存在一种与CcrM相对应的孤儿甲基转移酶，即DNA腺嘌呤甲基化酶(DNA adenine methylase, Dam)。Dam与CcrM一样催化腺嘌呤6号位的甲基化，调控细胞周期等过程。然而Dam在识别位点、时空表达方面与CcrM存在显著不同。Dam的识别序列为GATC，且其在整个细胞周期均有表达[3]。CcrM敲除在α变形菌中通常是致死的，而Dam敲除大多并不致死[53]。CcrM和Dam在表观遗传调控机制上也完全不同。CcrM协同转录激活因子GcrA促进基因表达，而Dam则利用甲基化抑制leucine-responsive regulatory protein (Lrp)、redox-sensitive regulator (OxyR)等转录阻遏蛋白与启动子的结合，从而起到基因表达开关的作用[3]。此外，CcrM和Dam在细胞周期调控上也有不同的功能倾向。Dam在复制起始、错配修复、鞭毛相变等方面起到直接的调控作用，而CcrM则在调控复制、细胞形态、染色体分离等方面发挥重要作用。CcrM和Dam的差异反映了α变形菌和γ变形菌在表观遗传调控上的长期分化。
表观遗传调控对细菌毒力影响深远，CcrM在人类、动物和植物病原细菌中广泛存在，它是否参与调控毒力基因表达呢？在B. melitensis等细菌的研究中尚未发现CcrM直接调控毒力基因的证据。然而王尔雅在研究A. radiobacter毒力基因表达时发现，CcrM介导的甲基化能够通过GcrA调控毒力基因所在质粒的复制，进而调控一系列毒力基因的表达[13]。这一发现证明了CcrM在A. radiobacter毒力基因表达调控中的重要作用，相信在其他重要的病原细菌中也存在CcrM调控毒力基因表达的机制。
尽管CcrM通过GcrA调控相关基因表达的途径已经较为清楚，但仍然存在相当数量的基因受到CcrM调控，而与GcrA似乎无关，不同物种受CcrM/GcrA调控的基因差异也很大[40]，有些α变形菌中甚至缺乏GcrA同源蛋白[6]。因此，GcrA可能并非CcrM实现其表观遗传调控的唯一途径，寻找和研究其他相关调节因子将成为CcrM表观遗传调控研究的重要方向之一。值得注意的是，有些基因是受到CcrM抑制的(表1)，而GcrA主要通过与σ因子互作激活下游基因表达，那么这些基因受到CcrM抑制的机制是什么？是否存在与GcrA对应的抑制因子？为了寻找新的CcrM依赖性表观遗传调控因子，利用染色质免疫沉淀(chromatin immunoprecipitation, ChIP)技术富集与GANTC基序结合的蛋白质，再通过质谱技术在富集得到的蛋白质中寻找相关调控因子是一种可行的方法。
在真核生物DNA甲基化表观调控系统中还存在一个重要的调控因子，即DNA去甲基化酶。DNA甲基化酶和去甲基化酶共同调控着基因组的甲基化状态。原核生物的去甲基化酶还有待进一步发掘。研究表明大肠杆菌AlkB在体外能够介导6mA去甲基化[54]，C. vibrioidesalkB基因的表达和ccrM一样受到细胞周期的影响[55]。有推测认为α变形菌的AlkB能够像真核生物去甲基化酶一样参与表观遗传的调控。然而，数据显示在正常生长的C. vibrioides中AlkB对基因组DNA的甲基化并无全局性的改变，尽管有部分CcrM调控的基因也受到了alkB突变的影响[29]。因此，α变形菌中是否存在CcrM对应的去甲基化酶还有待进一步探究。挖掘α变形菌中的去甲基化酶需要更多蛋白结构的支持。随着AI对蛋白质结构预测越来越准确，相信在α变形菌的基因组中能够找到更多潜在的去甲基化酶。此外，细菌突变体库的构建和新一代测序技术的应用也将有助于在α变形菌中寻找CcrM对应的去甲基化酶。
基金
收起
  • 国家自然科学基金(32300151)
文献
收起
参考文献 引证文献
排序方式：
[1]
OLIVEIRA PH, FANG G. Conserved DNA methyltransferases: a window into fundamental mechanisms of epigenetic regulation in bacteria[J]. Trends in Microbiology, 2021, 29(1): 28-40.
[2]
GORRELL R, KWOK T. The Helicobacter pylori methylome: roles in gene regulation and virulence[J]. Current Topics in Microbiology and Immunology, 2017, 400: 105-127.
[3]
WION D, CASADESÚS J. N6-methyl-adenine: an epigenetic signal for DNA-protein interactions[J]. Nature Reviews Microbiology, 2006, 4(3): 183-192.
[4]
FIORAVANTI A, FUMEAUX C, MOHAPATRA SS, BOMPARD C, BRILLI M, FRANDI A, CASTRIC V, VILLERET V, VIOLLIER PH, BIONDI EG. DNA binding of the cell cycle transcriptional regulator GcrA depends on N6-adenosine methylation in Caulobacter crescentus and other Alphaproteobacteria [J]. PLoS Genetics, 2013, 9(5): e1003541.
[5]
CHRISTEN B, ABELIUK E, COLLIER JM, KALOGERAKI VS, PASSARELLI B, COLLER JA, FERO MJ, McADAMS HH, SHAPIRO L. The essential genome of a bacterium[J]. Molecular Systems Biology, 2011, 7: 528.
[6]
CURTIS PD, BRUN YV. Identification of essential alphaproteobacterial genes reveals operational variability in conserved developmental and cell cycle systems[J]. Molecular Microbiology, 2014, 93(4): 713-735.
[7]
ZWEIGER G, MARCZYNSKI G, SHAPIRO L. A Caulobacter DNA methyltransferase that functions only in the predivisional cell[J]. Journal of Molecular Biology, 1994, 235(2): 472-485.
[8]
GONZALEZ D, KOZDON JB, McADAMS HH, SHAPIRO L, COLLIER J. The functions of DNA methylation by CcrM in Caulobacter crescentus: a global approach[J]. Nucleic Acids Research, 2014, 42(6): 3720-3735.
[9]
WRIGHT R, STEPHENS C, SHAPIRO L. The CcrM DNA methyltransferase is widespread in the alpha subdivision of proteobacteria, and its essential functions are conserved in Rhizobium meliloti and Caulobacter crescentus [J]. Journal of Bacteriology, 1997, 179(18): 5869-5877.
[10]
BRILLI M, FONDI M, FANI R, MENGONI A, FERRI L, BAZZICALUPO M, BIONDI EG. The diversity and evolution of cell cycle regulation in alpha-proteobacteria: a comparative genomic analysis[J]. BMC Systems Biology, 2010, 4: 52.
[11]
KAHNG LS, SHAPIRO L. The CcrM DNA methyltransferase of Agrobacterium tumefaciens is essential, and its activity is cell cycle regulated[J]. Journal of Bacteriology, 2001, 183(10): 3065-3075.
[12]
ROBERTSON GT, REISENAUER A, WRIGHT R, JENSEN RB, JENSEN A, SHAPIRO L, ROOP RM 2nd. The Brucella abortus CcrM DNA methyltransferase is essential for viability, and its overexpression attenuates intracellular replication in murine macrophages[J]. Journal of Bacteriology, 2000, 182(12): 3482-3489.
[13]
王尔雅. 根癌农杆菌细胞周期调节因子GcrA调节vir基因表达的机制研究[D]. 扬州: 扬州大学硕士学位论文, 2024.
WANG EY. Mechanism of vir gene expression regulated by Agrobacterium tumefaciens cell cycle regulator GcrA[D]. Yangzhou: Master’s Thesis of Yangzhou University, 2024 (in Chinese).
[14]
REICH NO, DANG E, KURNIK M, PATHURI S, WOODCOCK CB. The highly specific, cell cycle-regulated methyltransferase from Caulobacter crescentus relies on a novel DNA recognition mechanism[J]. Journal of Biological Chemistry, 2018, 293(49): 19038-19046.
[15]
WOODCOCK CB, YAKUBOV AB, REICH NO. Caulobacter crescentus cell cycle-regulated DNA methyltransferase uses a novel mechanism for substrate recognition[J]. Biochemistry, 2017, 56(30): 3913-3922.
[16]
BERDIS AJ, LEE I, COWARD JK, STEPHENS C, WRIGHT R, SHAPIRO L, BENKOVIC SJ. A cell cycle-regulated adenine DNA methyltransferase from Caulobacter crescentus processively methylates GANTC sites on hemimethylated DNA[J]. Proceedings of the National Academy of Sciences of the United States of America, 1998, 95(6): 2874-2879.
[17]
HORTON JR, WOODCOCK CB, OPOT SB, REICH NO, ZHANG X, CHENG XD. The cell cycle-regulated DNA adenine methyltransferase CcrM opens a bubble at its DNA recognition site[J]. Nature Communications, 2019, 10: 4600.
[18]
KONTTINEN O, CARMODY J, PATHURI S, ANDERSON K, ZHOU XF, REICH N. Cell cycle regulated DNA methyltransferase: fluorescent tracking of a DNA strand-separation mechanism and identification of the responsible protein motif[J]. Nucleic Acids Research, 2020, 48(20): 11589-11601.
[19]
KONTTINEN O, CARMODY J, KURNIK M, JOHNSON KA, REICH N. High fidelity DNA strand-separation is the major specificity determinant in DNA methyltransferase CcrM’s catalytic mechanism[J]. Nucleic Acids Research, 2023, 51(13): 6883-6898.
[20]
MALONE T, BLUMENTHAL RM, CHENG X. Structure-guided analysis reveals nine sequence motifs conserved among DNA amino-methyltransferases, and suggests a catalytic mechanism for these enzymes[J]. Journal of Molecular Biology, 1995, 253(4): 618-632.
[21]
SCHUBERT HL, BLUMENTHAL RM, CHENG XD. Many paths to methyltransfer: a chronicle of convergence[J]. Trends in Biochemical Sciences, 2003, 28(6): 329-335.
[22]
WRIGHT R, STEPHENS C, ZWEIGER G, SHAPIRO L, ALLEY MR. Caulobacter Lon protease has a critical role in cell-cycle control of DNA methylation[J]. Genes & Development, 1996, 10(12): 1532-1542.
[23]
REISENAUER A, QUON K, SHAPIRO L. The CtrA response regulator mediates temporal control of gene expression during the Caulobacter cell cycle[J]. Journal of Bacteriology, 1999, 181(8): 2430-2439.
[24]
BELLEFONTAINE AF, PIERREUX CE, MERTENS P, VANDENHAUTE J, LETESSON JJ, de BOLLE X. Plasticity of a transcriptional regulation network among alpha-proteobacteria is supported by the identification of CtrA targets in Brucella abortus [J]. Molecular Microbiology, 2002, 43(4): 945-960.
[25]
FRANCIS N, PONCIN K, FIORAVANTI A, VASSEN V, WILLEMART K, ONG TAP, RAPPEZ L, LETESSON JJ, BIONDI EG, de BOLLE X. CtrA controls cell division and outer membrane composition of the pathogen Brucella abortus [J]. Molecular Microbiology, 2017, 103(5): 780-797.
[26]
COLLIER J. Cell cycle control in Alphaproteobacteria [J]. Current Opinion in Microbiology, 2016, 30: 107-113.
[27]
GORA KG, TSOKOS CG, CHEN YE, SRINIVASAN BS, PERCHUK BS, LAUB MT. A cell-type-specific protein-protein interaction modulates transcriptional activity of a master regulator in Caulobacter crescentus [J]. Molecular Cell, 2010, 39(3): 455-467.
[28]
STEPHENS CM, ZWEIGER G, SHAPIRO L. Coordinate cell cycle control of a Caulobacter DNA methyltransferase and the flagellar genetic hierarchy[J]. Journal of Bacteriology, 1995, 177(7): 1662-1669.
[29]
CAMPBELL M, BARTON IS, ROOP RM 2nd, CHIEN P. Comparison of CcrM-dependent methylation in Caulobacter crescentus and Brucella abortus by nanopore sequencing[J]. Journal of Bacteriology, 2024, 206: e00083-24.
[30]
ZHOU XF, WANG JR, HERRMANN J, MOERNER WE, SHAPIRO L. Asymmetric division yields progeny cells with distinct modes of regulating cell cycle-dependent chromosome methylation[J]. Proceedings of the National Academy of Sciences of the United States of America, 2019, 116(31): 15661-15670.
[31]
MOUAMMINE A, COLLIER J. The impact of DNA methylation in Alphaproteobacteria [J]. Molecular Microbiology, 2018, 110(1): 1-10.
[32]
ARDISSONE S, REDDER P, RUSSO G, FRANDI A, FUMEAUX C, PATRIGNANI A, SCHLAPBACH R, FALQUET L, VIOLLIER PH. Cell cycle constraints and environmental control of local DNA hypomethylation in α-proteobacteria [J]. PLoS Genetics, 2016, 12(12): e1006499.
[33]
COLLIER J, McADAMS HH, SHAPIRO L. A DNA methylation ratchet governs progression through a bacterial cell cycle[J]. Proceedings of the National Academy of Sciences of the United States of America, 2007, 104(43): 17111-17116.
[34]
MARTIN S, FOURNES F, AMBROSINI G, ISELI C, BOJKOWSKA K, MARQUIS J, GUEX N, COLLIER J. DNA methylation by CcrM contributes to genome maintenance in the Agrobacterium tumefaciens plant pathogen[J]. Nucleic Acids Research, 2024, 52(19): 11519-11535.
[35]
DICENZO GC, CANGIOLI L, NICOUD Q, CHENG JHT, BLOW MJ, SHAPIRO N, WOYKE T, BIONDI EG, ALUNNI B, MENGONI A, MERGAERT P. DNA methylation in Ensifer species during free-living growth and during nitrogen-fixing symbiosis with Medicago spp.[J]. mSystems, 2022, 7(1): e0109221.
[36]
MARCZYNSKI GT, SHAPIRO L. Control of chromosome replication in Caulobacter crescentus [J]. Annual Review of Microbiology, 2002, 56: 625-656.
[37]
PINI F, de NISCO NJ, FERRI L, PENTERMAN J, FIORAVANTI A, BRILLI M, MENGONI A, BAZZICALUPO M, VIOLLIER PH, WALKER GC, BIONDI EG. Cell cycle control by the master regulator CtrA in Sinorhizobium meliloti [J]. PLoS Genetics, 2015, 11(5): e1005232.
[38]
MURRAY SM, PANIS G, FUMEAUX C, VIOLLIER PH, HOWARD M. Computational and genetic reduction of a cell cycle to its simplest, primordial components[J]. PLoS Biology, 2013, 11(12): e1001749.
[39]
GONZALEZ D, COLLIER J. DNA methylation by CcrM activates the transcription of two genes required for the division of Caulobacter crescentus [J]. Molecular Microbiology, 2013, 88(1): 203-218.
[40]
ADHIKARI S, ERILL I, CURTIS PD. Transcriptional rewiring of the GcrA/CcrM bacterial epigenetic regulatory system in closely related bacteria[J]. PLoS Genetics, 2021, 17(3): e1009433.
[41]
NARAYANAN S, JANAKIRAMAN B, KUMAR L, RADHAKRISHNAN SK. A cell cycle-controlled redox switch regulates the topoisomerase Ⅳ activity[J]. Genes & Development, 2015, 29(11): 1175-1187.
[42]
SANSELICIO S, BERGÉ M, THÉRAULAZ L, RADHAKRISHNAN SK, VIOLLIER PH. Topological control of the Caulobacter cell cycle circuitry by a polarized single-domain PAS protein[J]. Nature Communications, 2015, 6: 7005.
[43]
MOHAPATRA SS, FIORAVANTI A, VANDAME P, SPRIET C, PINI F, BOMPARD C, BLOSSEY R, VALETTE O, BIONDI EG. Methylation-dependent transcriptional regulation of crescentin gene (creS) by GcrA in Caulobacter crescentus [J]. Molecular Microbiology, 2020, 114(1): 127-139.
[44]
JONAS K, CHEN YE, LAUB MT. Modularity of the bacterial cell cycle enables independent spatial and temporal control of DNA replication[J]. Current Biology, 2011, 21(13): 1092-1101.
[45]
HOLTZENDORFF J, HUNG D, BRENDE P, REISENAUER A, VIOLLIER PH, McADAMS HH, SHAPIRO L. Oscillating global regulators control the genetic circuit driving a bacterial cell cycle[J]. Science, 2004, 304(5673): 983-987.
[46]
KOZDON JB, MELFI MD, LUONG K, CLARK TA, BOITANO M, WANG S, ZHOU B, GONZALEZ D, COLLIER J, TURNER SW, KORLACH J, SHAPIRO L, McADAMS HH. Global methylation state at base-pair resolution of the Caulobacter genome throughout the cell cycle[J]. Proceedings of the National Academy of Sciences of the United States of America, 2013, 110(48): E4658-E4667.
[47]
XU CR, HOLLIS H, DAI M, YAO XY, WATSON LT, CAO Y, CHEN MH. Modeling the temporal dynamics of master regulators and CtrA proteolysis in Caulobacter crescentus cell cycle[J]. PLoS Computational Biology, 2022, 18(1): e1009847.
[48]
REISENAUER A, SHAPIRO L. DNA methylation affects the cell cycle transcription of the CtrA global regulator in Caulobacter [J]. EMBO Journal, 2002, 21(18): 4969-4977.
[49]
HAAKONSEN DL, YUAN AH, LAUB MT. The bacterial cell cycle regulator GcrA is a σ70 cofactor that drives gene expression from a subset of methylated promoters[J]. Genes & Development, 2015, 29(21): 2272-2286.
[50]
GONZALEZ D, COLLIER J. Genomic adaptations to the loss of a conserved bacterial DNA methyltransferase[J]. mBio, 2015, 6(4): e00952.
[51]
WU XX, HAAKONSEN DL, SANDERLIN AG, LIU YJ, SHEN LQ, ZHUANG NN, LAUB MT, ZHANG Y. Structural insights into the unique mechanism of transcription activation by Caulobacter crescentus GcrA[J]. Nucleic Acids Research, 2018, 46(6): 3245-3256.
[52]
WU XX, YU CZ, MU WH, GU ZX, FENG Y, ZHANG Y. The structural mechanism for transcription activation by Caulobacter crescentus GcrA[J]. Nucleic Acids Research, 2023, 51(4): 1960-1970.
[53]
张文婷, 姚玉峰. 细菌DNA甲基化研究进展[J]. 生物化学与生物物理进展, 2018, 45(10): 1026-1038.
ZHANG WT, YAO YF. Research progress on bacterial DNA methylation[J]. Progress in Biochemistry and Biophysics, 2018, 45(10): 1026-1038 (in Chinese).
[54]
LI DY, DELANEY JC, PAGE CM, YANG XD, CHEN AS, WONG C, DRENNAN CL, ESSIGMANN JM. Exocyclic carbons adjacent to the N6 of adenine are targets for oxidation by the Escherichia coli adaptive response protein AlkB[J]. Journal of the American Chemical Society, 2012, 134(21): 8896-8901.
[55]
COLOMBI D, GOMES SL. An alkB gene homolog is differentially transcribed during the Caulobacter crescentus cell cycle[J]. Journal of Bacteriology, 1997, 179(10): 3139-3145.
2025年第65卷第9期
PDF下载
154
57
引用本文
BibTeX
文章信息
doi: 10.13343/j.cnki.wsxb.20250131
  • 接收时间：2025-02-22
  • 首发时间：2026-02-07
  • 出版时间：2025-09-04
补充材料
相关文章
文章信息
作者
出版历史
  • 收稿日期：2025-02-22
  • 录用日期：2025-04-23
基金
National Natural Science Foundation of China(32300151)
国家自然科学基金(32300151)
作者信息
    扬州大学 生物科学与技术学院，江苏 扬州
参考文献
分享链接
https://castjournals.cast.org.cn/joweb/wswxb/CN/10.13343/j.cnki.wsxb.20250131
分享至
全文二维码

扫描看全文

引用本文
BibTeX
本文的引用情况
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
关闭全屏
相关链接
论文
最新文章
热读文章
热点专题
内参
蓝皮书
产业发展报告
传播力报告
期刊分级目录
资讯
学术新闻
行业新闻
学术会议
行业会议
关于
关于我们
联系我们
北京市海淀区学院南路86号
100081
010-62199257
qkjq@cast.org.cn

中国科学技术协会版权所有 京ICP 备10216604号-4 海淀分局备案 1101084647
科技导报社主办 中国科协信息中心技术支持