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Article(id=1276616079644430609, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276616049617408127, articleNumber=null, orderNo=null, doi=10.3969/j.issn.1000-2561.2025.08.002, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1742832000000, receivedDateStr=2025-03-25, revisedDate=null, revisedDateStr=null, acceptedDate=1744819200000, acceptedDateStr=2025-04-17, onlineDate=1782298591766, onlineDateStr=2026-06-24, pubDate=1756051200000, pubDateStr=2025-08-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1782298591766, onlineIssueDateStr=2026-06-24, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1782298591766, creator=13701087609, updateTime=1782298591766, updator=13701087609, issue=Issue{id=1276616049617408127, tenantId=1146029695717560320, journalId=1235980609244409860, year='2025', volume='46', issue='8', pageStart='1785', pageEnd='2029', issueExtLink='null', onlineDate='null', pubDate='1756051200000', pubDateStr='2025-08-25', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1782298584608, creator='13701087609', updateTime=1782298660748, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1276616369089147039, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276616049617408127, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1276616369089147040, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276616049617408127, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=1795, endPage=1805, ext={EN=ArticleExt(id=1276616081733194003, articleId=1276616079644430609, tenantId=1146029695717560320, journalId=1235980609244409860, language=EN, title=Identification of Cassava C3H Genes and Analysis of the Expression Pattern in Response to Xanthomonas phaseoli pv. manihotis, columnId=1236256430337085821, journalTitle=Chinese Journal of Tropical Crops, columnName=Omics & Biotechnology, runingTitle=null, highlight=null, articleAbstract=

Cassava is one of the three major tuber crops in the world, and is also an important food crop in tropical and subtropical regions. Bacterial blight (CBB) is an important disease for cassava. CCCH (C3H)-type zinc finger proteins are widely present in most of organisms, not only involved in plant growth and development, hormone regulation, but also in response to biotic and abiotic stresses. Although the function of some C3H genes in other plants has been reported, but that in cassava has not been reported. In this study, we analyzed the phylogenetic characteristics, chromosome location, gene structure, promoter cis-acting elements and collinearity of cassava C3H genes. A total of 89 C3H genes (MeC3H01-MeC3H89) were identified in cassava, which were distributed on 18 chromosomes. The phylogenetic tree showed that 89 MeC3H proteins were classified into 4 subfamilies. The C3H gene promoters contains multiple stress, hormones, plant growth and development response elements. 39 pairs of cassava C3H genes had homologous relationships, indicating that these genes may be amplified by replication during evolution. We also analyzed the response expression pattern of C3H genes after cassava was infected by Xanthomonas phaseoli pv. manihotis (Xpm) by transcriptome and real-time quantitative PCR (qPCR) experiments. The expression levels of MeC3H49, MeC3H42 and MeC3H68 were significantly up-regulated, and those of MeC3H35, MeC3H77, MeC3H36, MeC3H31 and MeC3H86 were significantly down-regulated. In summary, we identified the cassava C3H gene family members and their expression patterns in response to Xpm infection, which lays a foundation for further analysis of the function of MeC3Hs.

, authors=null, authorsList=Min LI, Bingzheng WEI, Suhang JIA, Yinhua CHEN, Chunxia LI, authorCompany=null, correspAuthors=Chunxia LI, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=null), CN=ArticleExt(id=1276616083591270693, articleId=1276616079644430609, tenantId=1146029695717560320, journalId=1235980609244409860, language=CN, title=木薯C3H基因鉴定及应答菜豆黄单胞菌侵染表达模式分析, columnId=1236256430517440904, journalTitle=热带作物学报, columnName=组学与生物技术, runingTitle=null, highlight=null, articleAbstract=

木薯是世界三大薯类作物之一，同时也是热带、亚热带地区的重要粮食作物。木薯在生长发育过程中易遭受病虫害影响，其中细菌性枯萎病（cassava bacterial blight，CBB）是为害木薯的重要病害。CCCH（C3H）型锌指蛋白广泛存在于各种生物体内，不仅参与植物生长发育、激素调控，同时也响应生物及非生物胁迫。在拟南芥、水稻、大豆、小麦、矮牵牛、棉花等植物中，部分C3H基因的功能已有报道，但木薯C3H基因功能还鲜有研究。本研究利用生物信息学工具分析木薯MeC3H基因家族成员的系统进化特征、染色体位置、基因结构、启动子顺式作用元件和共线性等，并利用转录组和实时荧光定量PCR（qPCR）技术研究木薯受病原菌（Xanthomonas phaseoli pv. manihotis，Xpm）侵染后，MeC3H基因的表达模式。结果表明：木薯共有89个MeC3H基因，命名为MeC3H01~MeC3H89，分布在18条不同的染色体上；系统进化树显示，这89个MeC3H蛋白分为4个亚家族；MeC3H基因启动子含有逆境、植物激素以及植物生长发育响应元件；共线性分析发现，有39对木薯C3H基因具有同源关系，表明在进化过程中可能通过复制扩增基因家族成员；通过转录组和实时荧光定量PCR分析发现，在Xpm处理后，部分C3H基因表达发生显著变化，其中MeC3H49、MeC3H42、MeC3H68的表达量显著上调，MeC3H35、MeC3H77、MeC3H36、MeC3H31、MeC3H86的表达量显著下调。本研究结果为进一步分析MeC3H基因的功能提供参考。

, authors=

李敏（1998—），女，硕士研究生，研究方向：植物抗病机制。

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* 李春霞（LI Chunxia），E-mail：。
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李敏（1998—），女，硕士研究生，研究方向：植物抗病机制。

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李敏（1998—），女，硕士研究生，研究方向：植物抗病机制。

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(in Chinese), articleTitle=Identification and expression analysis of the HSP70 gene family underabiotic stresses in Litchi chinensis, refAbstract=null)], funds=[Fund(id=1276616101924573536, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, awardId=Qhys2023-250, language=CN, fundingSource=海南省研究生创新课题(Qhys2023-250), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1276616083821957415, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, xref=null, ext=[AuthorCompanyExt(id=1276616083834540328, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, companyId=1276616083821957415, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=School of Tropical Agriculture and Forestry, Hainan University / Key Laboratory of Sustainable Utilization of Tropical Biological Resources, Haikou, Hainan 570228, China), AuthorCompanyExt(id=1276616083838734633, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, companyId=1276616083821957415, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=海南大学热带农林学院/海南省热带生物资源可持续利用重点实验室，海南海口　570228)])], figs=[ArticleFig(id=1276616095167549774, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=EN, label=Fig. 1, caption=Chromosomal localization of MeC3H genes, figureFileSmall=5DZeDi+RO+ulhJQz2gA8Cg==, figureFileBig=LiCPyPoCVkeZxYije5KX7Q==, tableContent=null), ArticleFig(id=1276616097017237839, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=CN, label=图1, caption=木薯MeC3H基因的染色体定位, figureFileSmall=5DZeDi+RO+ulhJQz2gA8Cg==, figureFileBig=LiCPyPoCVkeZxYije5KX7Q==, tableContent=null), ArticleFig(id=1276616097315033424, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=EN, label=Fig. 2, caption=Phylogenetic relationships of C3H proteins, figureFileSmall=Ajvphi5geBZ5zeajOjgwNQ==, figureFileBig=gdfP9txnZxhG4fAVqe9pUw==, tableContent=null), ArticleFig(id=1276616097608634705, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=CN, label=图2, caption=C3H蛋白的系统进化关系, figureFileSmall=Ajvphi5geBZ5zeajOjgwNQ==, figureFileBig=gdfP9txnZxhG4fAVqe9pUw==, tableContent=null), ArticleFig(id=1276616099311522130, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=EN, label=Fig. 3, caption=The structural characteristics of MeC3H proteins sequence, figureFileSmall=KpQXtqzb+1eRPNA5vZM0Xw==, figureFileBig=fj6V0EFn+fuvn75uXHyLpg==, tableContent=null), ArticleFig(id=1276616099709981011, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=CN, label=图3, caption=MeC3H蛋白序列的结构特征

A：MeC3H基因家族进化树；B：MeC3H的15个保守基序分布，不同颜色代表不同的基序；C：木薯C3H基因结构。

, figureFileSmall=KpQXtqzb+1eRPNA5vZM0Xw==, figureFileBig=fj6V0EFn+fuvn75uXHyLpg==, tableContent=null), ArticleFig(id=1276616099810644308, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=EN, label=Fig. 4, caption=Intra-species collinearity analysis of the MeC3H family genes in cassava, figureFileSmall=WHAqHjcCmoms7ZOoGDcqHA==, figureFileBig=Jpx+ipSnLllyvQUnqxHszw==, tableContent=null), ArticleFig(id=1276616100116828501, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=CN, label=图4, caption=MeC3H家族基因的物种内共线性分析, figureFileSmall=WHAqHjcCmoms7ZOoGDcqHA==, figureFileBig=Jpx+ipSnLllyvQUnqxHszw==, tableContent=null), ArticleFig(id=1276616100188131670, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=EN, label=Fig. 5, caption=Syntenic relationship of C3H genes in Arabidopsis thaliana, Oryza sativa and Manihot esculenta, figureFileSmall=AvpJYfkrX9llzrphWVi60Q==, figureFileBig=akphnI5xvQKOcr+jlvLEbQ==, tableContent=null), ArticleFig(id=1276616100259434839, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=CN, label=图5, caption=木薯和拟南芥、水稻间的C3H基因共线性分析, figureFileSmall=AvpJYfkrX9llzrphWVi60Q==, figureFileBig=akphnI5xvQKOcr+jlvLEbQ==, tableContent=null), ArticleFig(id=1276616100569813336, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=EN, label=Fig. 6, caption=The homeopathic elements in the promoters of MeC3H genes, figureFileSmall=EYaHPRQPI1+mvtNjd7Nr2A==, figureFileBig=frbgQOu5kFpAbhgdHB1qkA==, tableContent=null), ArticleFig(id=1276616100636922201, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=CN, label=图6, caption=MeC3Hs基因启动子的顺式作用元件, figureFileSmall=EYaHPRQPI1+mvtNjd7Nr2A==, figureFileBig=frbgQOu5kFpAbhgdHB1qkA==, tableContent=null), ArticleFig(id=1276616100695642458, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=EN, label=Fig. 7, caption=Expression levels of MeC3H genes upon Xpm infection, figureFileSmall=3sgfYm9g/uT7KKjiZO3+pg==, figureFileBig=1c64Bdh2yTIRIEiZtdmVWQ==, tableContent=null), ArticleFig(id=1276616101018603867, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=CN, label=图7, caption=MeC3H基因在Xpm感染后的表达模式, figureFileSmall=3sgfYm9g/uT7KKjiZO3+pg==, figureFileBig=1c64Bdh2yTIRIEiZtdmVWQ==, tableContent=null), ArticleFig(id=1276616101085712732, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=EN, label=Fig. 8, caption=The expression patterns of cassava C3H genes upon Xpm infection, figureFileSmall=TJKcsfgDurNPmSVGVYWqYQ==, figureFileBig=UCm/bdXQMO74yep7rJy2BQ==, tableContent=null), ArticleFig(id=1276616101169598813, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=CN, label=图8, caption=MeC3H基因在Xpm侵染后的表达模式

*表示差异显著（P<0.05）；**表示差异极显著（P<0.01）。

, figureFileSmall=TJKcsfgDurNPmSVGVYWqYQ==, figureFileBig=UCm/bdXQMO74yep7rJy2BQ==, tableContent=null), ArticleFig(id=1276616101375119710, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=EN, label=Tab. 1, caption=

Primer sequences for qRT-PCR

, figureFileSmall=null, figureFileBig=null, tableContent=
引物名称Primer name正向引物（5′-3′）Forward primer (5′-3′)反向引物（5′-3′）Reverse primer (5′-3)
MeTubulinATGCGGTTCTTGATGTTGTTCTCGGTGAAGGGAATACAGAGA
MeC3H01CCAAATCTTTTTGAACCGGCCACAGCAACAACACCCCTAATCG
MeC3H03TGGCAGTCTTGTGCTTTGTGGCTACATATGTGCCATCCGCA
MeC3H06CTCAGGAGCTACAGCCAGAATAAACCTTCTTGGAAGCCAGCA
MeC3H07GAGTCCTTACTCGGCACCTGTGAGAGGCCAGCAACAGAAG
MeC3H31TGCCAGTCTGGTCAGTGTTCTGCCTTAACAACATTCGGCAC
MeC3H36ACCTTCAGCTCACACCTTCATACCACTGCACATACCCTAATTCTAT
MeC3H39TTGTTACGGTTTCGCGCTTCGTTGATAAGGAGTCCATTTCTTGCT
MeC3H42TGGTAGTCTTTCTCAAGGCGATCAGACCTAACCCCAATCCACA
MeC3H46ATCAGCGTGCCAAAGTAGCATAGCTCACGGCCACTCATTC
MeC3H49CAGCAGCAAGACAAAACGACACGATACTCACATTCACTTCCCTTTT
MeC3H58GGAGTTCTAGGTCTCTGGCGTTTTCCATGCGCAAACTGGC
MeC3H68FGCAAATCGTCTTCTCTGATTTGTGTCTGCCTCAACTGTACCATTTGTAT
MeC3H71GCAAGTTCAATCACCCTGCTACACCGGAGTTTTGGCTCTTGA
MeC3H77CGAGGCTGCTCTGAATGGAAAGGGCACCCAGAAGTACTGA
MeC3H86CTTTGTCAGGTAAGGCTGTTCCTGGTTGGCAGAAAGACCTTCA
), ArticleFig(id=1276616101459005791, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276616079644430609, language=CN, label=表1, caption=

qRT-PCR所用引物序列

, figureFileSmall=null, figureFileBig=null, tableContent=
引物名称Primer name正向引物（5′-3′）Forward primer (5′-3′)反向引物（5′-3′）Reverse primer (5′-3)
MeTubulinATGCGGTTCTTGATGTTGTTCTCGGTGAAGGGAATACAGAGA
MeC3H01CCAAATCTTTTTGAACCGGCCACAGCAACAACACCCCTAATCG
MeC3H03TGGCAGTCTTGTGCTTTGTGGCTACATATGTGCCATCCGCA
MeC3H06CTCAGGAGCTACAGCCAGAATAAACCTTCTTGGAAGCCAGCA
MeC3H07GAGTCCTTACTCGGCACCTGTGAGAGGCCAGCAACAGAAG
MeC3H31TGCCAGTCTGGTCAGTGTTCTGCCTTAACAACATTCGGCAC
MeC3H36ACCTTCAGCTCACACCTTCATACCACTGCACATACCCTAATTCTAT
MeC3H39TTGTTACGGTTTCGCGCTTCGTTGATAAGGAGTCCATTTCTTGCT
MeC3H42TGGTAGTCTTTCTCAAGGCGATCAGACCTAACCCCAATCCACA
MeC3H46ATCAGCGTGCCAAAGTAGCATAGCTCACGGCCACTCATTC
MeC3H49CAGCAGCAAGACAAAACGACACGATACTCACATTCACTTCCCTTTT
MeC3H58GGAGTTCTAGGTCTCTGGCGTTTTCCATGCGCAAACTGGC
MeC3H68FGCAAATCGTCTTCTCTGATTTGTGTCTGCCTCAACTGTACCATTTGTAT
MeC3H71GCAAGTTCAATCACCCTGCTACACCGGAGTTTTGGCTCTTGA
MeC3H77CGAGGCTGCTCTGAATGGAAAGGGCACCCAGAAGTACTGA
MeC3H86CTTTGTCAGGTAAGGCTGTTCCTGGTTGGCAGAAAGACCTTCA
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木薯C3H基因鉴定及应答菜豆黄单胞菌侵染表达模式分析
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李敏 , 魏炳峥 , 贾素行 , 陈银华 , 李春霞 *
热带作物学报 | 组学与生物技术 2025,46(8): 1795-1805
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热带作物学报 |组学与生物技术 2025 , 46 (8) : 1795 -1805
木薯C3H基因鉴定及应答菜豆黄单胞菌侵染表达模式分析
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李敏, 魏炳峥, 贾素行, 陈银华, 李春霞*
作者信息
  • 海南大学热带农林学院/海南省热带生物资源可持续利用重点实验室，海南海口　570228
通讯作者:
* 李春霞（LI Chunxia），E-mail：。
Identification of Cassava C3H Genes and Analysis of the Expression Pattern in Response to Xanthomonas phaseoli pv. manihotis
Min LI, Bingzheng WEI, Suhang JIA, Yinhua CHEN, Chunxia LI*
Affiliations
  • School of Tropical Agriculture and Forestry, Hainan University / Key Laboratory of Sustainable Utilization of Tropical Biological Resources, Haikou, Hainan 570228, China
出版时间: 2025-08-25 doi: 10.3969/j.issn.1000-2561.2025.08.002
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摘要
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木薯是世界三大薯类作物之一，同时也是热带、亚热带地区的重要粮食作物。木薯在生长发育过程中易遭受病虫害影响，其中细菌性枯萎病（cassava bacterial blight，CBB）是为害木薯的重要病害。CCCH（C3H）型锌指蛋白广泛存在于各种生物体内，不仅参与植物生长发育、激素调控，同时也响应生物及非生物胁迫。在拟南芥、水稻、大豆、小麦、矮牵牛、棉花等植物中，部分C3H基因的功能已有报道，但木薯C3H基因功能还鲜有研究。本研究利用生物信息学工具分析木薯MeC3H基因家族成员的系统进化特征、染色体位置、基因结构、启动子顺式作用元件和共线性等，并利用转录组和实时荧光定量PCR（qPCR）技术研究木薯受病原菌（Xanthomonas phaseoli pv. manihotis，Xpm）侵染后，MeC3H基因的表达模式。结果表明：木薯共有89个MeC3H基因，命名为MeC3H01~MeC3H89，分布在18条不同的染色体上；系统进化树显示，这89个MeC3H蛋白分为4个亚家族；MeC3H基因启动子含有逆境、植物激素以及植物生长发育响应元件；共线性分析发现，有39对木薯C3H基因具有同源关系，表明在进化过程中可能通过复制扩增基因家族成员；通过转录组和实时荧光定量PCR分析发现，在Xpm处理后，部分C3H基因表达发生显著变化，其中MeC3H49、MeC3H42、MeC3H68的表达量显著上调，MeC3H35、MeC3H77、MeC3H36、MeC3H31、MeC3H86的表达量显著下调。本研究结果为进一步分析MeC3H基因的功能提供参考。

木薯  /  C3H基因家族  /  基因表达  /  抗病响应  /  细菌性枯萎病

Cassava is one of the three major tuber crops in the world, and is also an important food crop in tropical and subtropical regions. Bacterial blight (CBB) is an important disease for cassava. CCCH (C3H)-type zinc finger proteins are widely present in most of organisms, not only involved in plant growth and development, hormone regulation, but also in response to biotic and abiotic stresses. Although the function of some C3H genes in other plants has been reported, but that in cassava has not been reported. In this study, we analyzed the phylogenetic characteristics, chromosome location, gene structure, promoter cis-acting elements and collinearity of cassava C3H genes. A total of 89 C3H genes (MeC3H01-MeC3H89) were identified in cassava, which were distributed on 18 chromosomes. The phylogenetic tree showed that 89 MeC3H proteins were classified into 4 subfamilies. The C3H gene promoters contains multiple stress, hormones, plant growth and development response elements. 39 pairs of cassava C3H genes had homologous relationships, indicating that these genes may be amplified by replication during evolution. We also analyzed the response expression pattern of C3H genes after cassava was infected by Xanthomonas phaseoli pv. manihotis (Xpm) by transcriptome and real-time quantitative PCR (qPCR) experiments. The expression levels of MeC3H49, MeC3H42 and MeC3H68 were significantly up-regulated, and those of MeC3H35, MeC3H77, MeC3H36, MeC3H31 and MeC3H86 were significantly down-regulated. In summary, we identified the cassava C3H gene family members and their expression patterns in response to Xpm infection, which lays a foundation for further analysis of the function of MeC3Hs.

cassava  /  C3H gene family  /  gene expression  /  disease resistance response  /  bacterial blight
李敏, 魏炳峥, 贾素行, 陈银华, 李春霞. 木薯C3H基因鉴定及应答菜豆黄单胞菌侵染表达模式分析. 热带作物学报, 2025 , 46 (8) : 1795 -1805 . DOI: 10.3969/j.issn.1000-2561.2025.08.002
Min LI, Bingzheng WEI, Suhang JIA, Yinhua CHEN, Chunxia LI. Identification of Cassava C3H Genes and Analysis of the Expression Pattern in Response to Xanthomonas phaseoli pv. manihotis[J]. Chinese Journal of Tropical Crops, 2025 , 46 (8) : 1795 -1805 . DOI: 10.3969/j.issn.1000-2561.2025.08.002
正文
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锌指蛋白家族是真核生物中最大的转录因子家族之一，其中CCCH（C3H）型锌指蛋白是一类在生物体中广泛存在的超家族蛋白，通过与DNA、RNA或蛋白质相互作用，调控植物的生长发育、激素响应及生物胁迫[1]。C3H锌指转录因子（zinc finger, Znf）由1~6个CCCH类型的锌指基序组成，该基序由3个半胱氨酸和1个组氨酸组成[1]。拟南芥（Arabidopsis thaliana）中有68个C3H家族成员，主要参与植物对盐、干旱等胁迫以及激素、光的响应，同时也调控花的发育[2]。例如，拟南芥HUA1是一种与花发育相关的RNA结合蛋白[3]，而AtTZF1可以在细胞核和细胞质间穿梭，与DNA、RNA均有结合活性，参与盐、激素等响应[4]。谷子（Setaria italica）SiC3H49是与干旱有关的锌指蛋白，可能通过调控气孔开放，在耐旱性中发挥关键作用[5]；棉花（Gossypium hirsutum）GhZFP1与GZIRD21A、GZIPR5蛋白相互作用介导植物的抗逆性[6]。在烟草（Nicotiana tabacum）中过表达GhZFP1可显著增加其耐盐性和抗病性[6]。水稻（Oryza sativa）C3H蛋白OsDOS可调控茉莉酸信号转导，过表达可延缓叶片衰老[7]；OsLOL2通过调节赤霉素（gibberellins, GAs）生物合成、调控水稻生长[8]。此外，C3H还参与植物的抗病响应。例如，水稻C3H基因OsBIRF转入烟草后可增强烟草对花叶病毒、野火病毒和烟粉虱的抗性[9]；拟南芥AtC3H18正调控水杨酸（salicylic acid, SA）和茉莉酸（jasmonic acid, JA）信号通路，从而增强对丁香假单胞菌（Pseudomonas syringae）的抗性[10]；甘蔗（Saccharum officinarum）C3H基因Sc-zf受黑穗病菌诱导表达，可能调控植物黑穗病抗性[11]。目前已在拟南芥、大豆（Glycine max）、水稻、小麦（Triticum aestivum）、棉花等植物中分析和鉴定出C3H基因家族成员[6]，但在木薯中还未有详细报道。
木薯（Manihot esculenta）是世界三大薯类作物之一，也是热带、亚热带地区重要的粮食和经济作物，有“地下粮仓”的誉称[12]。木薯起源于热带美洲，迄今已有4000多年的栽培历史，具有耐干旱、耐贫瘠、生长快、产量高、用途广等特点[12]。木薯的块根是其主要的营养器官，块根中富含大量的淀粉，是理想的工业原料，常用于生物燃料、造纸以及纺织业等。自19世纪20年代引种以来，木薯已成为我国第六大热带作物[13]。华南地区木薯已经有200多年的栽培历史，随着我国经济发展以及对生物能源、粮食生产等方面的需求，木薯已成为我国华南地区一种重要的旱地经济作物[12]，同时我国是世界上最大的木薯进口国。
近年来，病害一直是制约木薯产量的重要因素。由菜豆黄单胞菌木薯萎蔫致病变种（Xanthomonas phaseoli pv. manihotis，Xpm）引起的细菌性枯萎病（cassava bacterial blight, CBB）是木薯的重要病害，病原菌侵染后通过堵塞维管束、诱发叶片坏死和系统性萎蔫[14]，会造成叶片光合效率降低50%以上，严重时可使木薯减产20%~70%[15]。当前抗CBB木薯品种极度匮乏，抗病种质资源也很少，传统育种已很难满足产业需求，急需获得新的抗病种质或通过现代遗传技术创制抗病种质。了解木薯响应病原菌的防御机制，对抗病品种培育有重要作用。本研究首先运用生物信息学方法分析鉴定木薯MeC3H基因家族，从系统进化、染色体分布、Motif分析、启动子顺式作用元件、蛋白质理化性质等方面分析木薯MeC3H蛋白家族成员的特性，利用转录组和实时荧光定量PCR等技术分析C3H基因响应病原菌侵染下的表达模式，为木薯C3H基因的功能解析及木薯抗病育种提供理论依据。
1　材料与方法
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1.1　材料
供试材料为华南8号（South China 8，SC8）木薯，菜豆黄单胞菌木薯萎蔫致病变种（Xanthomonas phaseoli pv. manihotis，Xpm）CHN11由本实验室分离和保存。
1.2　方法
1.2.1　MeC3H家族成员筛选与鉴定
从Phytozome（https://phytozome.jgi.doe.gov/）数据库获取木薯SC8参考基因组以及拟南芥TAIR10基因组数据；从Pfam（http://pfam.xfam.org/）数据库下载C3H蛋白的结构域模型PF00642，使用HMMER软件从木薯数据库中搜索C3H蛋白（E-value<10-5），初步筛选出木薯MeC3H家族成员，再通过NCBI-CDD（https://www.ncbi.nlm.nih.gov/cdd/）、SMART（https:smart.embl.de/）等数据库验证候选蛋白。
1.2.2　MeC3H蛋白亚细胞定位预测
通过Wolf Post Ⅱ（https://www.genscript.com）在线软件预测木薯C3H蛋白的亚细胞定位[2]。
1.2.3　MeC3H家族系统进化分析
通过MEGA 11.0软件[16]（Pennsylvania State University，美国）中的Muscle功能对木薯和拟南芥C3H蛋白进行序列比对分析，并且采用临近-连接法（Neighbor-Joining algorithm，NJ）构建系统进化树，Bootstrap重复值为1000，用p-distance方法进行计算。对MeC3H家族成员进行亚家族分类，使用Evolview（https://www.evolgenius.info//evolview-v2）在线网站美化。
1.2.4　 MeC3H基因结构及蛋白保守基序分析
根据木薯基因组注释文件中MeC3H家族成员结构信息，使用TB tools v2.152[17]软件进行结构可视化分析。通过MEME软件预测C3H蛋白的保守基序（motif数为15，其他参数使用默认值），使用TB tools v2.152[17]软件可视化motif结果。
1.2.5　 MeC3H基因启动子顺式作用元件预测
通过TB tools从木薯基因组序列中提取MeC3H家族成员转录起始位点上游2000 bp的序列，利用PlantCARE（http://bioinformatics.psb.ugent.be/webtools/plantcare/html/）在线软件预测启动子区域的顺式作用元件[18]。
1.2.6　 MeC3H基因的染色体定位及共线性分析
从木薯基因组注释文件中获取MeC3H基因家族成员信息，使用TB tools v 2.152[17]软件将木薯C3H基因定位在染色体上。采用TB tools软件的MCScanX工具分析木薯与拟南芥、木薯与烟草C3H基因的共线性。
1.2.7　 MeC3H基因的表达模式分析
为了解MeC3H家族基因受Xpm侵染的诱导表达模式，利用课题组前期获得的RNA-seq数据，获得病原菌侵染不同时间点MeC3H家族成员的转录本FPKM（fragments per kilobase of exon per million reads mapped）值，并利用TBtools v2.152软件对MeC3H基因表达水平进行聚类分析以及热图绘制，挑选16个表达趋势显著的MeC3H基因进行qPCR验证。将生长30 d左右的SC8木薯叶片接种Xpm CHN11菌株[19]，对照组（CK）用灭菌ddH2O替代病原菌悬浮液，其他步骤与试验组相同，在不同时间点（8、24、50 h）取样，使用SteadyPure植物RNA快速提取试剂盒（艾科瑞，AG21040）提取总RNA。使用NanoDrop 2000分光光度计以及琼脂糖凝胶电泳检测RNA样品的完整性、浓度。利用EvoM-MLV反转录试剂盒（艾科瑞，AG11728）合成cDNA。在qTower3实时荧光定量PCR仪（耶拿，美国）上进行PCR。所用引物序列见表1，反应程序为预变性（95 ℃，30 s）；变性（95 ℃，10 s）、退火延伸（60 ℃，30 s），40个循环。采用2-ΔΔCt法计算基因相对表达水平[8]。利用GraphPad Prism 9软件（GraphPad Software，美国）绘图，并使用t检验比较数据间的差异。本试验包含3个生物学重复和3个技术重复。
2　结果与分析
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2.1　MeC3H蛋白亚细胞定位预测
通过全基因组分析发现，在木薯中共有89个MeC3H家族成员，根据其在染色体上的位置将其命名为MeC3H01~MeC3H89。据预测，MeC3H30、MeC3H43、MeC3H60、MeC3H72等4个蛋白定位于细胞质；MeC3H01、MeC3H08、MeC3H18、MeC3H20、MeC3H31、MeC3H39、MeC3H44、MeC3H50、MeC3H58等9个蛋白定位于叶绿体；其余76个蛋白定位于细胞核。亚细胞定位预测结果表明，大部分MeC3H家族成员可能在细胞核中发挥作用。
2.2　 MeC3H基因的染色体定位分析
89个MeC3H基因分布在18条染色体和1条Scaffold上，主要分布于Chr2（8个）、Chr1（7个）、Chr6（7个）、Chr8（7个）、Chr14（7个）、Chr3（6个）、Chr10（6个）、Chr11（6个）、Chr12（5个）、Chr9（4个）、Chr15（4个）、Chr16（4个），其余6条染色体上C3H基因数量为2~3个（图1）。除Chr1、Chr2、Chr6、Chr8、Chr14染色体上的C3H基因分布比较集中外，其余染色体上C3H基因分布比较分散。
2.3　MeC3H家族基因系统进化分析
根据拟南芥（AtC3Hs, 68个）和木薯（MeC3Hs,89个）C3H蛋白序列构建系统进化树（图2）。2个物种C3H蛋白可归为Ⅰ、Ⅱ、Ⅲ、Ⅳ 4个组。Ⅰ组包含34个MeC3H，28个AtC3H；Ⅱ组包含12个MeC3H，12个AtC3H；Ⅲ组包含17个MeC3H，12个AtC3H；Ⅳ组包含26个MeC3H，16个AtC3H；其中Ⅰ组还可划分为Ⅰa、Ⅰb、Ⅰc 3个亚类，分别包含16、16、2个MeC3H家族成员。大多数MeC3H分布在Ⅰ组和Ⅳ组。
2.4　 MeC3H基因结构及蛋白保守基序分析
为进一步预测MeC3H蛋白的功能，运用MEME软件进行保守基序预测。结果显示，15个保守基序的长度范围为15（第9个基序）~50个（第1、4、7、8、13、15个基序）氨基酸（图3）。木薯MeC3H基因的内含子数为0~27，其中有12个基因（MeC3H35、MeC3H52、MeC3H40、MeC3H44、MeC3H68、MeC3H04、MeC3H13、MeC3H36、MeC3H48、MeC3H28、MeC3H02、MeC3H09）含有1个内含子，7个基因不含内含子（MeC3H63、MeC3H30、MeC3H17、MeC3H81、MeC3H19、MeC3H80、MeC3H43）。
2.5　 MeC3H基因共线性分析
39对MeC3H基因具有共线性关系，如MeC-3H02与MeC3H09、MeC3H03与MeC3H10、Me-C3H04与MeC3H13、MeC3H05与MeC3H15、MeC-3H07与MeC3H22，其余成员间不存在共线性关系（图4）。因此，MeC3H基因家族存在基因复制的现象，在进化过程中可能复制扩增C3H家族成员。
为进一步探索MeC3H基因家族成员之间的进化关系，研究木薯与拟南芥、水稻的C3H家族之间的共线性，构建了两两物种之间的关系图（图5）。木薯与拟南芥、水稻中存在同源基因对，其中木薯与拟南芥之间的同源基因对数量远多于木薯与水稻之间的同源基因对。因此，相较于木薯与水稻，木薯与拟南芥的C3H基因家族具有更相近的同源进化关系。
2.6　 MeC3H基因启动子顺式作用元件预测
通过对木薯C3H家族成员转录起始位点上游对2000 bp左右启动子区域的分析发现，其启动子区域包含多种类型的顺式作用元件，除常见的TATA-box、CAAT等主要的顺式作用元件外，还包含了具有响应功能的顺式作用元件。按照功能分为光响应元件（G-Box、Box 4、GT1-motif、TCT-motif、GATA-motif）、逆境响应元件（ABRE、ARE、MBS、LTR、CGTCA-motif、TCA-element、TGACG-motif）和生长发育元件（CAT-box、TGA-element）（图6）。绝大多数MeC3H基因的启动子含有与光响应相关的顺式作用元件，光响应元件在全部89个MeC3H基因中存在数量最多，有1052个；其次是非生物胁迫响应元件有784个，其中脱落酸响应元件269个，占非生物胁迫响应元件的34%，抗氧化反应元件有259个，占非生物胁迫响应元件的33%；数量最少的顺式作用元件是植物生长发育响应元件（157个）。这表明MeC3H可能与光响应有关，同时在植物生长发育、生物胁迫以及非生物胁迫中也发挥重要作用。
2.7　 MeC3H基因响应Xpm的表达模式分析
为了解MeC3H基因受Xpm侵染后的表达模式，将生长30 d左右的SC8木薯苗接种Xpm CHN11菌株。本研究根据本实验室转录组数据分析结果（图7），挑选16个表达趋势显著的基因（MeC3H01、MeC3H03、MeC3H06、MeC3H07、MeC3H31、MeC3H36、MeC3H39、MeC3H42、MeC3H46、MeC3H49、MeC3H58、MeC3H68、MeC3H71、MeC3H77、MeC3H86）作为检测对象，进一步利用qPCR技术分析MeC3Hs基因的表达模式。结果表明，MeC3H49、MeC3H03、MeC3H42、MeC3H68、MeC3H01、MeC3H35、MeC3H36、MeC3H71、MeC3H31、MeC3H86基因的表达量均与RNA-seq结果一致，MeC3H58、MeC3H39、MeC3H77、MeC3H07、MeC3H06、MeC3H46这6个基因的qPCR与RNA-seq表达模式不一致。可能是2种技术检测范围和灵敏度不一致造成的。MeC3H49、MeC3H42、MeC3H68的表达量显著上调，MeC3H35、MeC3H77、MeC3H36、MeC3H31、MeC3H86的表达量显著下调（图8）。以上结果表明部分MeC3H基因可能在木薯抗病响应中有比较重要的作用。
3　讨论
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C3H基因广泛存在于多种真核生物中，如酵母、植物和哺乳动物等[20]，在植物的生长发育、生物胁迫以及非生物胁迫等方面具有重要的调控作用。尽管C3H基因家族在拟南芥、水稻、杨树、番茄等植物中已有较深入的研究，但关于木薯C3H基因家族的研究还鲜有报道。
本研究从木薯基因组中成功鉴定到89个MeC3H基因家族成员，它们定位在18条染色体上和1条Scaffold上。除Chr1、Chr2、Chr6、Chr8、Chr14染色体上的C3H基因分布比较集中外，其余染色体上C3H基因分布比较分散。基于89个木薯C3H蛋白和68个拟南芥C3H蛋白的系统发育进化树结果，可将C3H蛋白划分为Ⅰ、Ⅱ、Ⅲ、Ⅳ4个亚族，同一个亚族内MeC3H基因家族成员关系较近，有较高的同源性，同源基因发生复制时可能来自同一个祖先，推测其具有相似的功能。
基因启动子区域的顺式作用元件可以为后续该基因功能研究提供重要的方向指引。本研究鉴定到MeC3H基因启动子区域有大量光响应元件、逆境响应元件、激素响应元件，推测其成员广泛参与木薯抗逆和生长发育等重要途径[21]。内含子是高等生物基因组成的重要特征，在调控基因转录、表达和可变剪接等方面有重要作用。如在拟南芥部分SAPs基因中内含子缺失可增强转录效率[22]。基因在进化过程中会出现内含子增加或缺失的现象[23]。在MeC3H基因家族有7个成员不含内含子，可能与更快合成应激相关蛋白，调控应激响应有关。通过对MeC3H家族成员的基因结构进行分析发现，同一亚家族成员的外显子-内含子数量、保守基序特征相似，可能与同一亚族成员的生物学功能相似有关。木薯共有39对MeC3H同源基因之间存在共线性关系，表明在进化过程中基因可能通过复制方式扩增C3H家族成员。
通过SC8木薯在病原菌Xpm侵染下转录组数据以及实时荧光定量qPCR分析表明，部分MeC3H基因如MeC3H07、MeC3H68、MeC3H77等在Xpm侵染后快速表达，而MeC3H06、MeC3H31、MeC3H36、MeC3H71、MeC3H86等的表达则被Xpm抑制。说明这些转录调节因子可能在木薯抵御细菌性枯萎病菌侵染中发挥重要的作用。已有研究发现C3H基因参与调控植物的抗病响应，如OsBIRF、OsLOL2、OsDOS等调控水稻的抗病力，AtC3H18调控拟南芥抗细菌病害的能力以及Sc-zf调控甘蔗抗病力等[8-11]。综上推测，C3H基因可能参广泛参与木薯的抗病响应，调控木薯与病原菌的互作。
本研究通过对MeC3H基因家族进行生物信息学分析和对部分MeC3H基因在病原菌侵染下的调控机制分析，为进一步研究C3H基因在木薯抗细菌性枯萎病方面的功能奠定基础。
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  • 海南省研究生创新课题(Qhys2023-250)
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参考文献 引证文献
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2025年第46卷第8期
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doi: 10.3969/j.issn.1000-2561.2025.08.002
  • 接收时间：2025-03-25
  • 首发时间：2026-06-24
  • 出版时间：2025-08-25
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  • 收稿日期：2025-03-25
  • 录用日期：2025-04-17
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    海南大学热带农林学院/海南省热带生物资源可持续利用重点实验室，海南海口　570228

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* 李春霞（LI Chunxia），E-mail：。
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
species		 占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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