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Article(id=1276618344086238010, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276618341674520821, articleNumber=null, orderNo=null, doi=10.3969/j.issn.1000-2561.2025.12.002, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1754496000000, receivedDateStr=2025-08-07, revisedDate=null, revisedDateStr=null, acceptedDate=1757433600000, acceptedDateStr=2025-09-10, onlineDate=1782299131651, onlineDateStr=2026-06-24, pubDate=1766592000000, pubDateStr=2025-12-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1782299131651, onlineIssueDateStr=2026-06-24, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1782299131651, creator=13701087609, updateTime=1782299131651, updator=13701087609, issue=Issue{id=1276618341674520821, tenantId=1146029695717560320, journalId=1235980609244409860, year='2025', volume='46', issue='12', pageStart='2817', pageEnd='3084', issueExtLink='null', onlineDate='null', pubDate='1766592000000', pubDateStr='2025-12-25', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1782299131077, creator='13701087609', updateTime=1782299208862, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1276618668385637092, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276618341674520821, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1276618668385637093, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276618341674520821, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=2831, endPage=2839, ext={EN=ArticleExt(id=1276618344581165884, articleId=1276618344086238010, tenantId=1146029695717560320, journalId=1235980609244409860, language=EN, title=Functional Characterization of the Key Cold Resistance Gene MeCAD2 in cassava, columnId=1236256430337085821, journalTitle=Chinese Journal of Tropical Crops, columnName=Omics & Biotechnology, runingTitle=null, highlight=null, articleAbstract=

Cassava (Manihot esculenta Crantz), a vital tropical food crop, exhibits significant sensitivity to low temperature, which severely restricts its cultivation range. Lignin is a key stress-resistant component in cell walls, playing a crucial role in plant adaptation to low temperature stress. However, the function of cinnamyl alcohol dehydrogenase (CAD), a pivotal enzyme in lignin biosynthesis, during low-temperature stress responses remains unclear. This study analyzed physiological phenotypes, CAD enzyme activity and lignin accumulation in cassava under low temperature stress. CAD enzyme activity was significantly increased by 66.7% and lignin content was also increased by 14.8% at the low temperature of 10 ℃. Transcriptome profiling identified MeCAD2 as the key gene responsive to low temperature stress. Heterologous expression of MeCAD2 confirmed that its encoded MeCAD2 protein possessed CAD enzymatic activity. Virus-induced gene silencing (VIGS) of MeCAD2 in cassava resulted in a 18.9% reduction in CAD enzyme activity and a 12.3%–22.4% decrease in lignin content. Furthermore, under low temperature stress, MeCAD2-silenced plants exhibited exacerbated leaf damage, a 172.5% increase in malondialdehyde (MDA) accumulation, and intensified reactive oxygen species (ROS) staining. The results indicate that MeCAD2 enhances cassava low-temperature tolerance by regulating lignin deposition. This study elucidated the critical role of MeCAD2 in cassava’s response to low-temperature stress, providing a novel candidate gene for molecular breeding.

, authors=null, authorsList=Zheng CAI, Fan ZHANG, Kunhang LIU, Shiting DENG, Youzhi LI, Pingchuan ZHU, Xianwei FAN, authorCompany=null, correspAuthors=Pingchuan ZHU, Xianwei FAN, 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=1276618350767764298, articleId=1276618344086238010, tenantId=1146029695717560320, journalId=1235980609244409860, language=CN, title=木薯抗低温关键基因MeCAD2的功能鉴定, columnId=1236256430517440904, journalTitle=热带作物学报, columnName=组学与生物技术, runingTitle=null, highlight=null, articleAbstract=

木薯（Manihot esculenta Crantz）属于热带粮食作物，对低温的敏感性严重制约了种植区域。木质素是细胞壁中的组分，在抗低温胁迫中发挥关键作用，但木质素合成关键酶肉桂酸脱氢酶（CAD）在低温响应中的作用尚不明确。本研究通过分析低温胁迫下木薯的生理表型、CAD酶活及木质素变化，发现10 ℃低温处理显著诱导木薯中CAD酶活上升66.7%，且木质素增加14.8%。同时表达谱分析筛选出低温响应的核心基因MeCAD2。经原核表达证实其编码蛋白具有CAD催化活性，VIGS沉默该基因后，植株CAD酶活降低18.9%，木质素含量下降12.3%~22.4%。在低温胁迫下，MeCAD2沉默株系叶片损伤加剧，丙二醛积累量升高172.5%，活性氧染色加深，表明MeCAD2基因通过调控木质素沉积增强了木薯抗低温胁迫的能力。本研究揭示MeCAD2在木薯低温胁迫中的关键作用，为抗低温分子育种提供新靶点。

, authors=

蔡政（1996—），男，博士研究生，研究方向：木薯分子育种。

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* 朱平川（ZHU Pingchuan），E-mail：
樊宪伟（FAN Xianwei），E-mail：。
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蔡政（1996—），男，博士研究生，研究方向：木薯分子育种。

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蔡政（1996—），男，博士研究生，研究方向：木薯分子育种。

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Plant Physiology and Biochemistry, 2023, 194: 394-405., articleTitle=MeSWEET15a/b genes play a role in the resistance of cassava (Manihot esculenta Crantz) to water and salt stress by modulating sugar distribution, refAbstract=null), Reference(id=1276618370770400162, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, doi=null, pmid=null, pmcid=null, year=2024, volume=14, issue=1, pageStart=31213, pageEnd=null, url=null, language=null, rfNumber=[20], rfOrder=23, authorNames=HE Y X, ZHANG W J, ZUO X H, LI J N, XING M, ZHANG Y J, YOU J, ZHAO W, CHEN X, journalName=Scientific Reports, refType=null, unstructuredReference=HE Y X, ZHANG W J, ZUO X H, LI J N, XING M, ZHANG Y J, YOU J, ZHAO W, CHEN X. Dynamic transcriptomics unveils parallel transcriptional regulation in artemisinin and phenylpropanoid biosynthesis pathways under cold stress in Artemisia annua[J]. 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International Journal of Biological Macromolecules, 2025, 290: 139979., articleTitle=The CaCAD1-CaPOA1 module positively regulates pepper resistance to cold stress by increasing lignin accumulation, refAbstract=null), Reference(id=1276618370929783716, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, doi=null, pmid=null, pmcid=null, year=2020, volume=180, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[22], rfOrder=25, authorNames=MEHROTRA S, VERMA S, KUMAR S, KUMARI S, MISHRA B N, journalName=Environmental and Experimental Botany, refType=null, unstructuredReference=MEHROTRA S, VERMA S, KUMAR S, KUMARI S, MISHRA B N. Transcriptional regulation and signalling of cold stress response in plants: an overview of current understanding[J]. 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BMC Plant Biology, 2014, 14: 208., articleTitle=Increased expression of native cytosolic Cu/Zn superoxide dismutase and ascorbate peroxidase improves tolerance to oxidative and chilling stresses in cassava (Manihot esculenta Crantz), refAbstract=null), Reference(id=1276618371336631208, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, doi=null, pmid=null, pmcid=null, year=2019, volume=17, issue=4, pageStart=1121, pageEnd=1129, url=null, language=null, rfNumber=[25], rfOrder=29, authorNames=沈婕, 李淑霞, 彭明, journalName=分子植物育种, refType=null, unstructuredReference=沈婕, 李淑霞, 彭明. 木薯响应低温胁迫差异蛋白质的功能分析及鉴定[J]. 分子植物育种, 2019, 17(4): 1121-1129., articleTitle=木薯响应低温胁迫差异蛋白质的功能分析及鉴定, refAbstract=null), Reference(id=1276618371454071721, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, doi=null, pmid=null, pmcid=null, year=2019, volume=17, issue=4, pageStart=1121, pageEnd=1129, url=null, language=null, rfNumber=[25], rfOrder=30, authorNames=SHEN J, LI S X, PENG M, journalName=Molecular Plant Breeding, refType=null, unstructuredReference=SHEN J, LI S X, PENG M. Analysis and identification of differentially expressed proteins in response to low temperature stress in cassava[J]. Molecular Plant Breeding, 2019, 17(4): 1121-1129. (in Chinese), articleTitle=Analysis and identification of differentially expressed proteins in response to low temperature stress in cassava, refAbstract=null)], funds=[Fund(id=1276618365015815048, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, awardId=32160429, language=CN, fundingSource=国家自然科学基金项目(32160429), fundOrder=null, country=null), Fund(id=1276618365099701129, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, awardId=2021GXNSFDA196009, language=CN, fundingSource=广西自然科学基金项目(2021GXNSFDA196009), fundOrder=null, country=null), Fund(id=1276618365162615690, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, awardId=YCBZ2025017, language=CN, fundingSource=广西研究生教育创新计划资助项目(YCBZ2025017), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1276618351027811148, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, xref=null, ext=[AuthorCompanyExt(id=1276618351040394061, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, companyId=1276618351027811148, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=State Key Laboratory for Conservation and Utilization of Subtropical Agro-bioresources / College of Life Science and Technology, Guangxi University, Nanning, Guangxi 530004, China), AuthorCompanyExt(id=1276618351052976974, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, companyId=1276618351027811148, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=亚热带农业生物资源保护与利用国家重点实验室/广西大学生命科学与技术学院，广西南宁　530004)])], figs=[ArticleFig(id=1276618362268545916, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, language=EN, label=Fig. 1, caption=Impact of low temperature stress on cassava growth and development phenotype, CAD enzyme activity and lignin content, figureFileSmall=1HifIhXvSeynbPHwHBDL9Q==, figureFileBig=Yx1N0jLJ/0QgU08OldCIxw==, tableContent=null), ArticleFig(id=1276618362331460477, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, language=CN, label=图1, caption=低温对木薯生长发育表型、CAD酶活力及木质素含量的影响

A：木薯表型；B：木薯叶片的NBT和DAB染色；C：木薯叶片的CAD酶活；D：木薯叶片的木质素含量。*表示差异显著（P<0.05）；**表示差异极显著（P<0.01）；ns表示无显著差异。

, figureFileSmall=1HifIhXvSeynbPHwHBDL9Q==, figureFileBig=Yx1N0jLJ/0QgU08OldCIxw==, tableContent=null), ArticleFig(id=1276618364248257406, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, language=EN, label=Fig. 2, caption=Expression levels of MeCADs gene family under low temperature (6 ℃) stress, figureFileSmall=MwYXpzCMZNeUm3DBENPCuA==, figureFileBig=XL9oR7Nv6nCKs/zdOkpCrQ==, tableContent=null), ArticleFig(id=1276618364340532095, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, language=CN, label=图2, caption=MeCADs基因家族在低温（6 ℃）胁迫下的表达量

*表示差异显著（P<0.05）；**表示差异极显著（P<0.01）；***表示差异极显著（P<0.001）；ns表示无显著差异。

, figureFileSmall=MwYXpzCMZNeUm3DBENPCuA==, figureFileBig=XL9oR7Nv6nCKs/zdOkpCrQ==, tableContent=null), ArticleFig(id=1276618364399252352, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, language=EN, label=Fig. 3, caption=Purification and enzyme activity detection of GST-MeCAD2 protein, figureFileSmall=AROOEcBEdEjzfX0Fu+svfQ==, figureFileBig=C+/nPyHIe6sXXKhomyaBKA==, tableContent=null), ArticleFig(id=1276618364487332737, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, language=CN, label=图3, caption=GST-MeCAD2的蛋白纯化及酶活分析

A：GST-MeCAD2融合蛋白的SDS-PAGE图；B：GST蛋白与GST-MeCAD2重组蛋白的CAD酶活性。**表示差异极显著（P<0.01）。

, figureFileSmall=AROOEcBEdEjzfX0Fu+svfQ==, figureFileBig=C+/nPyHIe6sXXKhomyaBKA==, tableContent=null), ArticleFig(id=1276618364558635906, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, language=EN, label=Fig. 4, caption=Gene silencing efficiency of MeCAD2 and CAD enzyme activity and lignin content of silenced plants, figureFileSmall=emVqcgNUt+LB/vHKtIVkbQ==, figureFileBig=1/hys0VqX/z7nE5fVi2jWg==, tableContent=null), ArticleFig(id=1276618364617356163, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, language=CN, label=图4, caption=MeCAD2的沉默效率及沉默植株的叶片的CAD酶活性和木质素含量

*表示差异显著（P<0.05）；**表示差异极显著（P<0.01）。

, figureFileSmall=emVqcgNUt+LB/vHKtIVkbQ==, figureFileBig=1/hys0VqX/z7nE5fVi2jWg==, tableContent=null), ArticleFig(id=1276618364692853636, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, language=EN, label=Fig. 5, caption=Impact of low temperature (10 ℃) stress on MeCAD2 gene silencing lines, figureFileSmall=rhYeK78Ym/TXPW6zQVmrQA==, figureFileBig=L1SPLpqlRJpvSutVJ4JhLA==, tableContent=null), ArticleFig(id=1276618364759962501, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, language=CN, label=图5, caption=低温胁迫（10 ℃）对MeCAD2基因沉默株系的影响

A、B：植株的表型；C、D：植株的DAB和NBT染色；E、F、G、H：植株叶片的MDA含量、脯氨酸含量、CAD酶活力和木质素含量变化。*表示差异显著（P<0.05）；**表示差异极显著（P<0.01）；***表示差异极显著（P<0.001）；ns表示无显著差异。

, figureFileSmall=rhYeK78Ym/TXPW6zQVmrQA==, figureFileBig=L1SPLpqlRJpvSutVJ4JhLA==, tableContent=null), ArticleFig(id=1276618364848042886, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, language=EN, label=Tab. 1, caption=

Primers

, figureFileSmall=null, figureFileBig=null, tableContent=
引物名称Primer name引物序列（5′-3′）Primer sequence（5′-3′）
pGEX-4T-1-CAD2FgatctggttccgcgtggatccATGGTAGCCAAATTGCCAGAAG
pGEX-4T-1-CAD2RacccgggaattccggggatccATGTCAATAACAAATCGGTATCTAACATC
pCAMBIA2300-CAD2FggagaggacagggtacccgggATGGTAGCCAAATTGCCAGAAG
pCAMBIA2300-CAD2RcgactctagaggatccccgggGCTGCTGCATTGGATTGTATTG
pTRV2CAD2FagaaggcctccatggggatccGATAATTGCACAAACAATCTTGAAAA
pTRV2CAD2RcgtgagctcggtaccggatccCATCAAGAGGCATGTTGTCTGG
qMeCAD1FTTGTAATGATAGGGAAGAAG
qMeCAD1RACAGTAATTCCAGCACATA
qMeCAD2FGCTGATGTTAGATACCGAT
qMeCAD2RCTCATAATAATAGACTGCAAA
qMeCAD3FGTATTGGAGGAATGAAAGAG
qMeCAD3RCAAATCGGTATCTAACATCA
qMeCAD4FACAGAGTTTCCAAAGAGCA
qMeCAD4RAAGGGAGAAAGATGACCAGAT
qMeCAD5FTCAGTCCTCTTCATCGTTT
qMeCAD5RAAATCCCACAGTACAAAACC
qMeCAD6FCAGAGGTCGGGAGCAAAGT
qMeCAD6RAGAGTAGCCGCCGTAAGTG
qMeCAD7FTCAACATTTTCTGCTTTCA
qMeCAD7RGATGGGATGCTCTTCTTCT
qMeCAD8FGCAGGGATTGTGGGTTTAG
qMeCAD9FTACTGTTAAGGCCCGAAGC
qMeCAD9RGCAGCCCATCCAAATGACT
qMeCAD10FGTAGAGGTTGTCCCGATTG
qMeCAD10RCGATTTTATTATAGTAGGCT
qMeCAD11FGAAGCTCCTTATCCCAGTT
qMeCAD11RCTAAGATTCAAGGCAAACAA
qMeCAD12FCAACCACGCAACCAACCAC
qMeCAD12RGAGCTGCCCAGCCAGAAAC
qMeCAD13FACCGCTTCTTAGTCTGCTC
qMeCAD13RATCCTATGTTGCTTCCTCC
qMeCAD14FCTTGATGCTACTGCTCCTC
qMeCAD14RTAGCCTCCTGCTTCTTGTT
qMeCAD16FGATGGGATGTCACCAGAAC
qMeCAD16RCTCCTCTTAGCCCACTCTT
qMeEF1aFTGAACCACCCTGGTCAGATTGGAA
qMeEF1aRAACTTGGGCTCCTTCTCAAGCTCT
), ArticleFig(id=1276618364927734663, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276618344086238010, language=CN, label=表1, caption=

引物

, figureFileSmall=null, figureFileBig=null, tableContent=
引物名称Primer name引物序列（5′-3′）Primer sequence（5′-3′）
pGEX-4T-1-CAD2FgatctggttccgcgtggatccATGGTAGCCAAATTGCCAGAAG
pGEX-4T-1-CAD2RacccgggaattccggggatccATGTCAATAACAAATCGGTATCTAACATC
pCAMBIA2300-CAD2FggagaggacagggtacccgggATGGTAGCCAAATTGCCAGAAG
pCAMBIA2300-CAD2RcgactctagaggatccccgggGCTGCTGCATTGGATTGTATTG
pTRV2CAD2FagaaggcctccatggggatccGATAATTGCACAAACAATCTTGAAAA
pTRV2CAD2RcgtgagctcggtaccggatccCATCAAGAGGCATGTTGTCTGG
qMeCAD1FTTGTAATGATAGGGAAGAAG
qMeCAD1RACAGTAATTCCAGCACATA
qMeCAD2FGCTGATGTTAGATACCGAT
qMeCAD2RCTCATAATAATAGACTGCAAA
qMeCAD3FGTATTGGAGGAATGAAAGAG
qMeCAD3RCAAATCGGTATCTAACATCA
qMeCAD4FACAGAGTTTCCAAAGAGCA
qMeCAD4RAAGGGAGAAAGATGACCAGAT
qMeCAD5FTCAGTCCTCTTCATCGTTT
qMeCAD5RAAATCCCACAGTACAAAACC
qMeCAD6FCAGAGGTCGGGAGCAAAGT
qMeCAD6RAGAGTAGCCGCCGTAAGTG
qMeCAD7FTCAACATTTTCTGCTTTCA
qMeCAD7RGATGGGATGCTCTTCTTCT
qMeCAD8FGCAGGGATTGTGGGTTTAG
qMeCAD9FTACTGTTAAGGCCCGAAGC
qMeCAD9RGCAGCCCATCCAAATGACT
qMeCAD10FGTAGAGGTTGTCCCGATTG
qMeCAD10RCGATTTTATTATAGTAGGCT
qMeCAD11FGAAGCTCCTTATCCCAGTT
qMeCAD11RCTAAGATTCAAGGCAAACAA
qMeCAD12FCAACCACGCAACCAACCAC
qMeCAD12RGAGCTGCCCAGCCAGAAAC
qMeCAD13FACCGCTTCTTAGTCTGCTC
qMeCAD13RATCCTATGTTGCTTCCTCC
qMeCAD14FCTTGATGCTACTGCTCCTC
qMeCAD14RTAGCCTCCTGCTTCTTGTT
qMeCAD16FGATGGGATGTCACCAGAAC
qMeCAD16RCTCCTCTTAGCCCACTCTT
qMeEF1aFTGAACCACCCTGGTCAGATTGGAA
qMeEF1aRAACTTGGGCTCCTTCTCAAGCTCT
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木薯抗低温关键基因MeCAD2的功能鉴定
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蔡政 , 张凡 , 刘坤行 , 邓诗婷 , 李有志 , 朱平川 * , 樊宪伟 *
热带作物学报 | 组学与生物技术 2025,46(12): 2831-2839
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热带作物学报 |组学与生物技术 2025 , 46 (12) : 2831 -2839
木薯抗低温关键基因MeCAD2的功能鉴定
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蔡政, 张凡, 刘坤行, 邓诗婷, 李有志, 朱平川* , 樊宪伟*
作者信息
  • 亚热带农业生物资源保护与利用国家重点实验室/广西大学生命科学与技术学院，广西南宁　530004
通讯作者:
* 朱平川（ZHU Pingchuan），E-mail：
樊宪伟（FAN Xianwei），E-mail：。
Functional Characterization of the Key Cold Resistance Gene MeCAD2 in cassava
Zheng CAI, Fan ZHANG, Kunhang LIU, Shiting DENG, Youzhi LI, Pingchuan ZHU* , Xianwei FAN*
Affiliations
  • State Key Laboratory for Conservation and Utilization of Subtropical Agro-bioresources / College of Life Science and Technology, Guangxi University, Nanning, Guangxi 530004, China
出版时间: 2025-12-25 doi: 10.3969/j.issn.1000-2561.2025.12.002
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摘要
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木薯（Manihot esculenta Crantz）属于热带粮食作物，对低温的敏感性严重制约了种植区域。木质素是细胞壁中的组分，在抗低温胁迫中发挥关键作用，但木质素合成关键酶肉桂酸脱氢酶（CAD）在低温响应中的作用尚不明确。本研究通过分析低温胁迫下木薯的生理表型、CAD酶活及木质素变化，发现10 ℃低温处理显著诱导木薯中CAD酶活上升66.7%，且木质素增加14.8%。同时表达谱分析筛选出低温响应的核心基因MeCAD2。经原核表达证实其编码蛋白具有CAD催化活性，VIGS沉默该基因后，植株CAD酶活降低18.9%，木质素含量下降12.3%~22.4%。在低温胁迫下，MeCAD2沉默株系叶片损伤加剧，丙二醛积累量升高172.5%，活性氧染色加深，表明MeCAD2基因通过调控木质素沉积增强了木薯抗低温胁迫的能力。本研究揭示MeCAD2在木薯低温胁迫中的关键作用，为抗低温分子育种提供新靶点。

木薯  /  低温胁迫  /  肉桂酸脱氢酶  /  MeCAD2  /  木质素

Cassava (Manihot esculenta Crantz), a vital tropical food crop, exhibits significant sensitivity to low temperature, which severely restricts its cultivation range. Lignin is a key stress-resistant component in cell walls, playing a crucial role in plant adaptation to low temperature stress. However, the function of cinnamyl alcohol dehydrogenase (CAD), a pivotal enzyme in lignin biosynthesis, during low-temperature stress responses remains unclear. This study analyzed physiological phenotypes, CAD enzyme activity and lignin accumulation in cassava under low temperature stress. CAD enzyme activity was significantly increased by 66.7% and lignin content was also increased by 14.8% at the low temperature of 10 ℃. Transcriptome profiling identified MeCAD2 as the key gene responsive to low temperature stress. Heterologous expression of MeCAD2 confirmed that its encoded MeCAD2 protein possessed CAD enzymatic activity. Virus-induced gene silencing (VIGS) of MeCAD2 in cassava resulted in a 18.9% reduction in CAD enzyme activity and a 12.3%–22.4% decrease in lignin content. Furthermore, under low temperature stress, MeCAD2-silenced plants exhibited exacerbated leaf damage, a 172.5% increase in malondialdehyde (MDA) accumulation, and intensified reactive oxygen species (ROS) staining. The results indicate that MeCAD2 enhances cassava low-temperature tolerance by regulating lignin deposition. This study elucidated the critical role of MeCAD2 in cassava’s response to low-temperature stress, providing a novel candidate gene for molecular breeding.

cassava  /  low temperature stress  /  cinnamyl alcohol dehydrogenase  /  MeCAD2  /  lignin
蔡政, 张凡, 刘坤行, 邓诗婷, 李有志, 朱平川, 樊宪伟. 木薯抗低温关键基因MeCAD2的功能鉴定. 热带作物学报, 2025 , 46 (12) : 2831 -2839 . DOI: 10.3969/j.issn.1000-2561.2025.12.002
Zheng CAI, Fan ZHANG, Kunhang LIU, Shiting DENG, Youzhi LI, Pingchuan ZHU, Xianwei FAN. Functional Characterization of the Key Cold Resistance Gene MeCAD2 in cassava[J]. Chinese Journal of Tropical Crops, 2025 , 46 (12) : 2831 -2839 . DOI: 10.3969/j.issn.1000-2561.2025.12.002
正文
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木薯（Manihot esculenta Crantz）作为重要的热带粮食作物，是全球近8亿人口的主要热量来源[1]。然而，对低温的敏感性严重制约木薯向亚热带及温带地区的推广种植[2-3]。研究表明，10~15 ℃的低温即可导致木薯叶片光合效率下降、活性氧（ROS）积累及细胞膜损伤，最终影响木薯生产[4-5]。3个月龄的木薯苗在7 ℃的低温下处理4 h，茎尖部位和叶片即出现明显的失水和萎蔫现象[6]。低温不仅导致木薯种茎萌发率和存活率显著下降，而且还促进木薯块根的木质化[7]，这将增加从块根中提取淀粉的成本，降低块根的加工性能，影响鲜食木薯的风味和口感。
在逆境胁迫下，通过细胞壁修饰增强机械强度维持细胞完整性有助于植物抗逆[8]。肉桂酸脱氢酶（cinnamyl alcohol dehydrogenase, CAD）是木质素合成途径中的关键酶，催化肉桂醛还原为肉桂醇。辣椒中CAD活性受低温诱导上调，并通过木质素沉积增强植株抗冻性[9]。过表达IbCAD1增强了甘薯块茎中木质素积累和耐低温贮藏能力[10]。低温条件下，EjNAC3通过调控EjCAD转录诱导枇杷果实木质化[11]。芒草中低温通过诱导CAD活性增强木质素前体合成，进而促进细胞壁交联与机械强度提升，是其维持细胞完整性、提高抗冻性的关键机制[12]。但木薯中CAD基因家族的功能分化及其在低温胁迫中的作用尚未明确。
CAD基因家族的构成及对逆境响应存在较大差异。水稻基因组中包含12个CAD同源基因，其中OsCAD2、OsCAD3、OsCAD6和OsCAD8A参与不同生物和非生物胁迫的防御反应[13]；桑树基因组中的5个CAD基因表现出在木质素生物合成和胁迫反应中的功能分歧[14]；大豆GsCAD1被证实通过调控木质素增强了对大豆花病毒的抗性[15]。由此可见，不同物种中CAD基因可能通过时空特异性表达模式参与特定胁迫响应。然而，木薯基因组中共鉴定出16个CAD家族成员[16]，其表达模式与功能研究仍较少。前期转录组数据显示，低温胁迫可能激活木薯中部分CAD基因表达[17]，但其分子机制及生理效应亟待验证。
基于上述背景，本研究聚焦以下科学问题：（1）低温胁迫是否通过调控CAD活性诱导木薯木质素积累？（2）木薯CAD基因家族中是否存在低温响应的关键成员？（3）特定CAD基因的功能缺失是否削弱木薯抗寒性？为此，本研究系统分析了低温处理下木薯的生理表型、CAD酶活性及木质素含量变化，结合基因表达谱筛选出关键候选基因MeCAD2，并通过原核表达、病毒诱导基因沉默（VIGS）及抗性表型分析揭示其功能。本研究首次阐明MeCAD2在木薯低温响应中的调控作用，为解析木质素介导的抗寒机制提供新视角，也为分子育种提供潜在靶点。
1　材料与方法
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1.1　材料
1.1.1　植物材料及生长条件
供试木薯品种为南植199（NZ199），采自广西大学亚热带农科新城种植基地的成熟茎段，于28 ℃、光暗时间比为14∶10条件下温室培养，生长5周后用于低温处理或VIGS试验；供试烟草（Nicotiana benthamiana）于光照培养箱25 ℃培养，光暗时间比为14∶10。
1.1.2　菌株及载体
大肠杆菌DH5α及BL21（DE3）感受态细胞，农杆菌GV3101（pSoup）感受态细胞，均购自生工生物工程（上海）股份有限公司。病毒载体pTRV1、pTRV2由本实验室保存；载体pCAMBIA2300-eGFP、pGEX-4T-1由中国科学院遗传与发育研究所馈赠。
1.1.3　主要试剂
2×Taq PCR Master Mix、ClonExpress® Ⅱ One Step Cloning Kit、FastPure Universal Plant Total RNA Isolation Kit均购自南京诺唯赞生物科技有限公司；PerfectStartTM Green qPCR SuperMix、TransScript® One-Step gDNA Removal and cDNA Synthesis SuperMix均购自北京全式金生物技术有限公司；DNA纯化回收试剂盒、质粒小提试剂盒均购自江苏康为世纪生物科技股份有限公司；木质素含量检测试剂盒、肉桂醇脱氢酶（cinnamyl alcoholdehydrogenase, CAD）活性检测试剂盒、谷胱甘肽-琼脂糖凝胶4B（GST标签纯化树脂）、卡那霉素、氨苄青霉素、利福平均购自索莱宝生物科技有限公司。
1.2　方法
1.2.1　表达载体构建
从木薯基因组中获得MeCAD2（Manes. 13G102200）的cDNA序列，利用在线软件CE Design（http://www.vazyme.com）设计其无终止密码子的ORF框无缝克隆引物（表1）。将木薯RNA反转录为cDNA，将其作为模板，以特异性引物pGEX-4T-1-CAD2F/R和pCAMBIA2300-CAD2F/R进行PCR扩增得到目的基因片段。以限制性内切酶BamH酶切pGEX-Ⅰ4T-1载体、以限制性内切酶SamⅠ酶切pCAMBIA2300-eGFP载体，将目的基因片段和对应的线性化载体用无缝克隆连接，转化至大肠杆菌DH5α，提质粒后送生工生物工程（上海）股份有限公司测序。
1.2.2　实时荧光定量PCR
基于MeCOMTs序列设计特异性荧光定量PCR引物（表1）。以MeEF1a作为内参基因，使用荧光定量PCR仪（ABI 7000）进行检测。反应体系与PCR反应条件参照PerfectStartTM Green qPCR SuperMix试剂盒。每个样品检测重复3次，采用2–ΔΔCt法计算目的基因的相对表达量。
1.2.3　VIGS载体构建与木薯侵染试验
木薯VIGS试验参考ZENG等[18]的方法。利用SGN VIGS（https://vigs.solgenomics.net/）软件设计MeCAD2的VIGS区段，根据片段设计引物，以上述1.2.1中的阳性质粒为模板PCR扩增VIGS片段，经测序验证后，与酶切线性化载体pTRV2连接，转化至DH5α感受态细胞。提取质粒后送生工生物工程（上海）股份有限公司测序正确后，将重组质粒转化到GV3101（pSoup）感受态细胞。分别挑取pTRV1、pTRV2和pTRV2-CAD2已转化农杆菌GV3101的单克隆于抗性培养基中震荡培养24 h，离心收集菌体，加入重悬液：1 mol/L MES（吗啉乙磺酸）、1 mol/L MgCl2、200 mmol/L AS（乙酰丁香酮），重悬菌体并调整OD600为0.6，将pTRV2-CAD2和pTRV2分别与pTRV1等量混合，28 ℃黑暗放置4 h。每个处理组设置生物学重复3株，选择健康的叶片，用2 mL无针头注射器将悬浮菌液压渗注射进木薯叶片背面。以含有质粒pTRV2与pTRV1的混合侵染液为阴性对照（CK），含有质粒pTRV2-CAD2与pTRV1的混合侵染液为试验组。侵染14 d后，分别提取CK和试验组木薯新叶总RNA并反转录为cDNA检测目的基因表达量。
1.2.4　木薯低温胁迫后生理特性研究
选取MeCAD2基因沉默成功的试验组木薯苗（pTRV2-CAD2）和阴性对照木薯苗（CK），同时10 ℃低温处理5 d。参照北京索莱宝生物技术有限公司生理指标测定试剂盒的操作步骤，检测胁迫处理前后木薯株系叶片的木质素含量、肉桂酸脱氢酶（CAD）酶活、丙二醛（MDA）含量和脯氨酸（Pro）含量，用酶标仪测定吸光值。每个样品设3次重复。NBT及DAB染色参考FAN等[19]的方法，分别称取25 mg NBT和DAB充分溶解于50 mL磷酸缓冲盐溶液（phosphate buffered saline, PBS）pH 7.4，选取低温处理后CK和试验组木薯的叶片，清洗后浸泡在NBT和DAB染色液中，避光过夜染色后用95%乙醇沸水浴脱色，在显微镜下拍照观察。
1.2.5　融合蛋白的表达与纯化
参考大肠杆菌密码子偏好性，人工合成MeCAD2的编码序列并插入载体pGEX-4T-1。将重组质粒转入BL21（DE3）感受态细胞中，选取单克隆于LB培养基37 ℃培养至菌体OD600为0.4~0.6，加入终浓度为0.5 mmol/L的异丙基-D-硫代半乳糖苷（IPTG），16 ℃培养12 h。诱导产生的融合GST标签重组蛋白使用SDS-PAGE电泳检测可溶性，然后使用GST标签纯化树脂纯化重组蛋白。
2　结果与分析
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2.1　低温诱导木薯肉桂酸脱氢酶（CAD）活力升高及木质素积累
为了探究木质素及其关键合成酶CAD是否在木薯响应低温胁迫过程中发挥作用，本研究分析了低温处理下木薯的形态变化、叶片损伤程度、CAD酶活力和木质素含量（图1）。结果显示，与25 ℃相比，15 ℃处理木薯叶片无明显变化，10 ℃处理叶片下垂并发生一定程度卷曲（图1A)。叶片用二氨基联苯胺（DAB）和氮蓝四唑（NBT）染色显示（图1B），随着低温胁迫程度的加剧，染色逐渐加深，说明叶片受损伤程度逐渐加深。同时，低温处理显著提高了木薯的CAD酶活力和木质素含量（图1C、图1D），特别是在10 ℃处理下CAD酶活力和木质素含量分别提高66.7%和14.8%。由此推测，木薯中CAD可能通过调节木质素的生物合成参与低温胁迫响应。
2.2　木薯MeCADs基因家族在低温胁迫下的表达分析
为探究木薯在低温胁迫下调控CAD酶活力的关键基因，本研究对木薯低温胁迫下CAD家族成员的表达量进行分析（图2），其中共有12个基因在叶片中检测出表达，MeCAD3、MeCAD7、MeCAD8、MeCAD13表达量过低，无法检测。与CK相比，MeCAD9、MeCAD10、MeCAD12、MeCAD14、MeCAD15、MeCAD16的表达量在低温处理下无明显提高，MeCAD2、MeCAD5、MeCAD11基因在低温处理下12~36 h内表达量持续上升（图2），其中MeCAD2基因表达量在低温处理条件下36 h上升4倍，因此选择MeCAD2作为候选基因。
2.3　 MeCAD2的原核表达及酶活分析
为了确定MeCAD2基因的功能，首先在大肠杆菌中表达MeCAD2蛋白，将粗蛋白用GST标签蛋白纯化试剂盒（上海碧云天生物技术股份有限公司）纯化，SDS-PAGE电泳显示纯化后的GST-MeCAD2融合蛋白为60~70 kDa的单一条带（图3A），与预期大小一致。使用BCA法测定纯化后GST-MeCAD2蛋白浓度，使用CAD活性检测试剂盒检测CAD酶活性。结果表明纯化的GST-MeCAD2蛋白具有CAD酶活性（图3B）。
2.4　沉默MeCAD2降低木薯的CAD酶活性与木质素含量
利用VIGS技术沉默木薯MeCAD2的表达，进一步探究其在木薯中的功能。在接种14 d后，对接种植株新长出的叶片进行基因表达量检测，以注射pTRV2空载植株作为CK，注射pTRV2-MeCAD2的T1、T2、T3 3个株系中MeCAD2的表达量均下调70%以上（图4A），证明基因沉默成功。测定CK与MeCAD2基因沉默植株中CAD酶活性，结果显示MeCAD2沉默植株的CAD酶活性比CK降低了18.9%（图4B）。与此同时，对木质素含量的测定结果显示，与CK相比，T1、T2、T3 3个株系的木质素含量分别降低了15.0%、12.3%和22.4%（图4C）。结果表明，MeCAD2基因沉默不仅有效降低CAD酶活性，同时也导致木质素生物合成受阻，最终显著降低植株的木质素积累水平。
2.5　 MeCAD2介导的木质素积累改变木薯抗寒性
为了进一步分析木薯MeCAD2基因表达下调是否影响木薯对低温胁迫的抗性，本研究将MeCAD2沉默植株与CK进行低温胁迫处理。10 ℃处理5 d后，MeCAD2沉默植株的叶片损伤更为严重。叶片中的丙二醛和脯氨酸含量测定表明，MeCAD2沉默植株中的丙二醛含量比CK提高了172.5%，脯氨酸含量提高了98.4%。取CK和MeCAD2沉默植株相同部位叶片进行NBT和DAB染色，MeCAD2沉默植株叶片显示出更深的染色，说明MeCAD2沉默植株叶片受损伤更严重。结果表明沉默MeCAD2后木薯的抵御低温能力降低（图5）。
3　讨论
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本研究系统阐明肉桂酸脱氢酶基因MeCAD2在木薯响应低温胁迫中的关键作用，为解析木质素介导的植物耐低温提供了新证据，也为木薯耐低温分子育种奠定了重要基础。
前期转录组数据提示木薯CAD基因家族成员可能参与低温响应[6]，但缺乏功能验证。本研究通过时间动态表达谱分析，成功筛选出受低温显著诱导的核心成员MeCAD2，其表达模式与低温胁迫下CAD酶活力及木质素含量的显著升高高度吻合，暗示其可能是驱动木质素生物合成增强的关键因子。通过原核表达体系，本研究直接证实MeCAD2蛋白具备CAD酶活性，为其在木质素合成途径中的催化功能提供生化基础。更重要的是，利用VIGS技术特异性沉默MeCAD2，导致植株CAD总酶活力下降、木质素积累显著减少，并伴随严重的低温损伤表型。结果表明，MeCAD2是调控木薯低温下木质素合成通路的关键基因，其表达缺失直接削弱植株抗寒能力。
木质素作为细胞壁重要组分，其沉积可增强细胞壁机械强度，维持低温下细胞结构完整性[20-22]。本研究证实，低温胁迫能有效激活MeCAD2表达并提升CAD酶活力，进而促进木质素在木薯中的积累。沉默MeCAD2后木质素合成受阻，低温胁迫下植株表现出更严重的膜脂过氧化和活性氧（ROS）爆发，最终导致叶片萎蔫坏死。这一结果与辣椒[21]、甘薯[10]中CAD介导的木质素积累增强抗寒性/耐贮性的报道一致，表明CAD基因通过强化木质化防御屏障以减轻低温伤害，是植物中保守的胁迫适应策略。
值得注意的是，低温胁迫下脯氨酸在MeCAD2沉默植株中异常积累，可能反映细胞为补偿细胞壁强度不足而启动渗透保护。类似地，木薯耐寒品种在低温下可溶性糖含量增加，通过维持渗透平衡缓解脱水损伤[4]；籼稻品种（明恢86）通过维持高可溶性糖含量调节胞内水势，减少细胞失水，缓解膜损伤[23]。这表明木薯在抗低温方面是木质素沉积与多维度抗寒机制协同作用。在低温条件下，MeCAD2沉默株系中活性氧（和H2O2）的积累显著加剧，木质素沉积可能通过形成物理屏障限制ROS扩散，从而间接保护细胞膜系统。同时，抗氧化酶（如SOD、APX）的协同表达亦可增强抗寒性；超表达MeCu/ZnSOD与MeAPX2已被证明能显著提高转基因木薯的低温存活率[24-25]。这些发现提示，木质素加固细胞壁与ROS清除机制之间存在功能互补性，共同构成了抵御低温胁迫的复杂防御机制。
综上，本研究揭示MeCAD2作为木薯低温响应的核心调控因子，通过驱动木质素生物合成增强细胞壁机械屏障，从而提升植株抗寒能力。其功能缺失直接导致木质素沉积受阻、ROS清除失衡及膜系统稳定性下降，证实CAD介导的木质化途径在木薯低温防御中的必要性。本研究不仅为解析木质素介导的植物抗寒机制提供直接分子证据，也为木薯抗低温分子育种奠定了靶点。
基金
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  • 国家自然科学基金项目(32160429)
  • 广西自然科学基金项目(2021GXNSFDA196009)
  • 广西研究生教育创新计划资助项目(YCBZ2025017)
文献
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2025年第46卷第12期
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doi: 10.3969/j.issn.1000-2561.2025.12.002
  • 接收时间：2025-08-07
  • 首发时间：2026-06-24
  • 出版时间：2025-12-25
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  • 收稿日期：2025-08-07
  • 录用日期：2025-09-10
基金
国家自然科学基金项目(32160429)
广西自然科学基金项目(2021GXNSFDA196009)
广西研究生教育创新计划资助项目(YCBZ2025017)
作者信息
    亚热带农业生物资源保护与利用国家重点实验室/广西大学生命科学与技术学院，广西南宁　530004

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* 朱平川（ZHU Pingchuan），E-mail：
樊宪伟（FAN Xianwei），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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