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Article(id=1208489275478360869, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1208489266397692345, articleNumber=null, orderNo=null, doi=10.16438/j.0513-4870.2021-0899, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1624204800000, receivedDateStr=2021-06-21, revisedDate=1628438400000, revisedDateStr=2021-08-09, acceptedDate=null, acceptedDateStr=null, onlineDate=1766055895919, onlineDateStr=2025-12-18, pubDate=1639238400000, pubDateStr=2021-12-12, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1766055895919, onlineIssueDateStr=2025-12-18, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1766055895919, creator=13701087609, updateTime=1766055895919, updator=13701087609, issue=Issue{id=1208489266397692345, tenantId=1146029695717560320, journalId=1189982191388893191, year='2021', volume='56', issue='12', pageStart='3203', pageEnd='3554', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1766055893754, creator=13701087609, updateTime=1766136983434, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1208829381217227030, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1208489266397692345, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1208829381217227031, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1208489266397692345, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=3325, endPage=3330, ext={EN=ArticleExt(id=1208489276250112861, articleId=1208489275478360869, tenantId=1146029695717560320, journalId=1189982191388893191, language=EN, title=Identification and expression analysis of the DaNAC transcription factor family in Dipsacus asper, columnId=1208489267291079103, journalTitle=Acta Pharmaceutica Sinica, columnName=Special Reports Ⅱ: Biosynthesis of Plant Natural Products and Synthetic Biology for Their Production, runingTitle=null, highlight=null, articleAbstract=

NAC transcription factor genes play an important role in regulating plant adversity stress tolerance and secondary metabolism. To explore DaNAC transcription factor participation in the synthesis of asperosaponin Ⅵ in Dipsacus asper, we analyzed the expression of DaNAC genes based on full-length transcriptome data from different tissues (root, stem, leaf, flower, seed) to provide a theoretical foundation for regulating the metabolism of D. asper. RNA-seq data was used to identify open reading frames. Bioinformatic methods were used to identify the conserved domain motifs and construct an evolutionary tree. qRT-PCR was carried out to analyze tissue-specific and adversity-stressed expression. Twenty-nine DaNAC sequences were identified, all of which contain the conserved NAM domain and conserved motif 1 and motif 2 at the N terminal. Five DaNAC genes are closely related to the NAC genes in Arabidopsis thaliana and rice that are involved in adversity stress and are clustered in the Group Ⅰ subfamily. qRT-PCR revealed that DaNAC genes are differentially expressed between tissues. The expression levels were highest in leaves, followed by roots, stems and petioles, and the lowest in flowers and seeds. Compared with normal growth conditions, the expression of four NAC genes was up-regulated by treatment with low temperature (15 ℃). The expression of three genes (34564NAC2, 33883NAC48, 6727NAC14) was up-regulated and one gene (34480NAC22) was down-regulated by 150 μmol·L-1 MeJA. The results illustrate that the expression of NAC genes is induced by adversity stress, which provides a foundation for further study on the role of NAC family members in adversity stress in D. asper.

, correspAuthors=Tao ZHOU, authorNote=null, correspAuthorsNote=null, copyrightStatement=Copyright ©2021 Acta Pharmaceutica Sinica. All rights reserved., copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, authorCompany=null, fund=null, authors=null, authorsList=Jiao XU, Zheng-ping HU, Hua HE, Juan GUO, Wei-ke JIANG, Cheng-hong XIAO, Liang-yuan LI, Tao ZHOU), CN=ArticleExt(id=1208489279790104703, articleId=1208489275478360869, tenantId=1146029695717560320, journalId=1189982191388893191, language=CN, title=川续断DaNAC基因家族成员的鉴定及表达分析, columnId=1208489268524204487, journalTitle=药学学报, columnName=专题报道Ⅱ：植物天然产物生物合成及合成生物学, runingTitle=null, highlight=null, articleAbstract=

NAC基因在植物的逆境胁迫及次生代谢等过程具有重要的调控作用。为挖掘参与川续断皂苷Ⅵ合成的NAC基因家族成员, 本研究利用川续断根、茎、叶、花、种子等不同组织构建的全长转录组数据库筛选包含完整开放阅读框的NAC家族成员, 对其进行保守结构域及motif分析、进化树比对分析、组织表达差异分析和逆境胁迫诱导表达分析。本文共筛选得到29个NAC家族成员, 在N端都具有NAC基因家族特有的NAM结构域以及保守的motif 1和motif 2; 其中5个DaNAC蛋白和拟南芥及水稻中与逆境胁迫有关的NAC亲缘关系较近, 共同聚在Group Ⅰ亚家族; qRT-PCR表达分析发现, DaNAC基因在叶片中表达量较高, 其次是根、茎、叶柄, 在花和种子中表达量最低; 15 ℃低温处理川续断后, 川续断4个DaNAC基因均受低温诱导上调表达, 但表达模式存在一定的差异; 另外, 34564NAC2、33883NAC48、6727NAC14基因均受150 μmol·L-1茉莉酸甲酯的诱导上调表达, 34480NAC22基因受茉莉酸甲酯的诱导下调表达。结果表明川续断NAC基因受逆境胁迫诱导表达, 为进一步揭示NAC家族成员在川续断逆境胁迫中的作用奠定了基础。

, correspAuthors=周涛, authorNote=null, correspAuthorsNote=
*周涛，Tel: 13985175215, E-mail:
, copyrightStatement=版权所有©《药学学报》编辑部2021, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=zI5YkNSw/Mfc1pQMv3dI2Q==, magXml=08F2Eobx9DVY4il4beiUEg==, pdfUrl=null, pdf=UsZTeOz6uE8fKnbBIIVtIA==, pdfFileSize=2338647, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=OB6QYwu8gQAJVkU7jy7jXQ==, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=3sMudeRoynp5/ykYpB2Cuw==, mapNumber=null, authorCompany=null, fund=null, authors=null, authorsList=徐娇, 胡正平, 何华, 郭娟, 江维克, 肖承鸿, 李良远, 周涛)}, authors=[Author(id=1208489280469582047, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1208489275478360869, 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=1208489280587022577, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1208489275478360869, authorId=1208489280469582047, language=EN, stringName=Jiao XU, firstName=Jiao, middleName=null, lastName=XU, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=1, address=1. 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Gene name Forward primer sequence (5'-3') Reverse primer sequence (5'-3') Product of PCR/bp
34564NAC2 GTCTTGCCGATGTTGACCGCT CTGTGCCTTCTCCACCGTTCC 110
33883NAC48 CTTGAGGTTGGACGATTGGGTAT CAGTTGGGTTAGGATTTGTGGC 136
6727NAC14 GAGCAAAGATGAAACCCAACACG CACCACAAACAACCCTAAAATCAT 169
34480NAC22 AAGACCTAACCGGACCACTGAAC TACCATATTCTTGGAGGAGGGGA 208
DaACT103 ACAGCCGAGCGGGAAATTGT TCTGGGCAGCGGAATCTCTCT 170
), ArticleFig(id=1208489288950465528, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1208489275478360869, language=CN, label=Table 1, caption=

Primer information of qRT-PCR

, figureFileSmall=null, figureFileBig=null, tableContent=
Gene name Forward primer sequence (5'-3') Reverse primer sequence (5'-3') Product of PCR/bp
34564NAC2 GTCTTGCCGATGTTGACCGCT CTGTGCCTTCTCCACCGTTCC 110
33883NAC48 CTTGAGGTTGGACGATTGGGTAT CAGTTGGGTTAGGATTTGTGGC 136
6727NAC14 GAGCAAAGATGAAACCCAACACG CACCACAAACAACCCTAAAATCAT 169
34480NAC22 AAGACCTAACCGGACCACTGAAC TACCATATTCTTGGAGGAGGGGA 208
DaACT103 ACAGCCGAGCGGGAAATTGT TCTGGGCAGCGGAATCTCTCT 170
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川续断DaNAC基因家族成员的鉴定及表达分析
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徐娇 1 , 胡正平 1 , 何华 1 , 郭娟 2 , 江维克 1 , 肖承鸿 1 , 李良远 1 , 周涛 1, *
药学学报 | 专题报道Ⅱ：植物天然产物生物合成及合成生物学 2021,56(12): 3325-3330
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药学学报 | 专题报道Ⅱ：植物天然产物生物合成及合成生物学 2021, 56(12): 3325-3330
川续断DaNAC基因家族成员的鉴定及表达分析
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徐娇1, 胡正平1, 何华1, 郭娟2, 江维克1, 肖承鸿1, 李良远1, 周涛1, *
作者信息
  • 1.贵州中医药大学, 贵州 贵阳 550025
  • 2.中国中医科学院中药资源中心, 北京 100700

通讯作者:

*周涛，Tel: 13985175215, E-mail:
Identification and expression analysis of the DaNAC transcription factor family in Dipsacus asper
Jiao XU1, Zheng-ping HU1, Hua HE1, Juan GUO2, Wei-ke JIANG1, Cheng-hong XIAO1, Liang-yuan LI1, Tao ZHOU1, *
Affiliations
  • 1. Guizhou University of Traditional Chinese Medicine, Guiyang 550025, China
  • 2. National Resource Center for Chinese Materia Medica, China Academy of Chinese Medical Sciences, Beijing 100700, China
出版时间: 2021-12-12 doi: 10.16438/j.0513-4870.2021-0899
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摘要
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NAC基因在植物的逆境胁迫及次生代谢等过程具有重要的调控作用。为挖掘参与川续断皂苷Ⅵ合成的NAC基因家族成员, 本研究利用川续断根、茎、叶、花、种子等不同组织构建的全长转录组数据库筛选包含完整开放阅读框的NAC家族成员, 对其进行保守结构域及motif分析、进化树比对分析、组织表达差异分析和逆境胁迫诱导表达分析。本文共筛选得到29个NAC家族成员, 在N端都具有NAC基因家族特有的NAM结构域以及保守的motif 1和motif 2; 其中5个DaNAC蛋白和拟南芥及水稻中与逆境胁迫有关的NAC亲缘关系较近, 共同聚在Group Ⅰ亚家族; qRT-PCR表达分析发现, DaNAC基因在叶片中表达量较高, 其次是根、茎、叶柄, 在花和种子中表达量最低; 15 ℃低温处理川续断后, 川续断4个DaNAC基因均受低温诱导上调表达, 但表达模式存在一定的差异; 另外, 34564NAC2、33883NAC48、6727NAC14基因均受150 μmol·L-1茉莉酸甲酯的诱导上调表达, 34480NAC22基因受茉莉酸甲酯的诱导下调表达。结果表明川续断NAC基因受逆境胁迫诱导表达, 为进一步揭示NAC家族成员在川续断逆境胁迫中的作用奠定了基础。

川续断  /  NAC转录因子  /  表达分析  /  川续断皂苷Ⅵ

NAC transcription factor genes play an important role in regulating plant adversity stress tolerance and secondary metabolism. To explore DaNAC transcription factor participation in the synthesis of asperosaponin Ⅵ in Dipsacus asper, we analyzed the expression of DaNAC genes based on full-length transcriptome data from different tissues (root, stem, leaf, flower, seed) to provide a theoretical foundation for regulating the metabolism of D. asper. RNA-seq data was used to identify open reading frames. Bioinformatic methods were used to identify the conserved domain motifs and construct an evolutionary tree. qRT-PCR was carried out to analyze tissue-specific and adversity-stressed expression. Twenty-nine DaNAC sequences were identified, all of which contain the conserved NAM domain and conserved motif 1 and motif 2 at the N terminal. Five DaNAC genes are closely related to the NAC genes in Arabidopsis thaliana and rice that are involved in adversity stress and are clustered in the Group Ⅰ subfamily. qRT-PCR revealed that DaNAC genes are differentially expressed between tissues. The expression levels were highest in leaves, followed by roots, stems and petioles, and the lowest in flowers and seeds. Compared with normal growth conditions, the expression of four NAC genes was up-regulated by treatment with low temperature (15 ℃). The expression of three genes (34564NAC2, 33883NAC48, 6727NAC14) was up-regulated and one gene (34480NAC22) was down-regulated by 150 μmol·L-1 MeJA. The results illustrate that the expression of NAC genes is induced by adversity stress, which provides a foundation for further study on the role of NAC family members in adversity stress in D. asper.

Dipsacus asper  /  NAC transcription factor  /  expression analysis  /  asperosaponin Ⅵ
徐娇, 胡正平, 何华, 郭娟, 江维克, 肖承鸿, 李良远, 周涛. 川续断DaNAC基因家族成员的鉴定及表达分析. 药学学报, 2021 , 56 (12) : 3325 -3330 . DOI: 10.16438/j.0513-4870.2021-0899
Jiao XU, Zheng-ping HU, Hua HE, Juan GUO, Wei-ke JIANG, Cheng-hong XIAO, Liang-yuan LI, Tao ZHOU. Identification and expression analysis of the DaNAC transcription factor family in Dipsacus asper[J]. Acta Pharmaceutica Sinica, 2021 , 56 (12) : 3325 -3330 . DOI: 10.16438/j.0513-4870.2021-0899
正文
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中药续断为川续断科植物川续断Dipsacus asper Wall. ex Henry的干燥根, 具有补肝肾、强筋骨、安胎的功效[1], 现代临床常用于骨科疾病的治疗。川续断皂苷Ⅵ是续断药材的主要活性成分, 属于齐墩果烷三萜皂苷类化合物。研究发现川续断皂苷Ⅵ可诱导成骨细胞成熟分化, 进而促进骨形成[2], 而且对小鼠急性肝损伤[3]及缺氧诱导的心肌细胞凋亡具有一定的保护作用[4]。
已有研究认为, 三萜皂苷类化学成分主要由甲羟戊酸(MVA) 途径合成[5], 主要包括三萜皂苷上游前体的形成、三萜皂苷骨架的构建、三萜皂苷骨架下游的修饰。三萜皂苷的生物合成受干旱、低温[6, 7]等调控, 植物合成三萜皂苷等次生代谢产物, 可抵御多种逆境胁迫环境。研究发现, 不同产地、不同海拔生长的川续断受逆境胁迫的影响各不相同, 其主要活性成分川续断皂苷Ⅵ在根部的积累也存在较大的差异[8, 9], 川续断皂苷Ⅵ的合成和积累与环境因子密切相关。作者前期研究发现, 茉莉酸甲酯可显著促进川续断中川续断皂苷Ⅵ的合成和积累, 有利于植物抵抗逆境环境的胁迫[10]。但是逆境胁迫如何调控三萜皂苷合成机制尚不明确。NAC基因是响应植物逆境胁迫应答反应中非常重要的一类转录因子[11]。已有研究发现, NAC-MYB-GRN基因调控网络调控木质素的生物合成, 促进植物对逆境环境的适应[12]。同时, NAC转录因子参与植物的耐旱、耐盐、耐低温等多种非生物逆境胁迫, 调节植物次生代谢产物的合成, 提高植物对多种逆境胁迫的抗性[13, 14]。因此, 推测NAC转录因子调控川续断皂苷Ⅵ的合成, 提高川续断对逆境环境的适应性。
本研究以川续断为研究材料, 在前期川续断全长转录组测序的基础上, 利用生物信息学分析技术挖掘川续断中逆境相关的DaNAC转录因子, 并对其氨基酸序列进行生物信息学分析, 采用qRT-PCR技术分析其在川续断不同组织中的表达模式, 探索DaNAC基因对低温胁迫及茉莉酸甲酯的应答, 为解析逆境胁迫调控川续断皂苷Ⅵ合成的分子机制奠定理论基础。
材料与方法
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材料  川续断(Dipsacus asperoides) 采集于贵州省毕节市威宁县梅花山, 栽培于贵州中医药大学试验田及温室中。试验田中新鲜根、茎、叶、叶柄、花、果实, 采集后在液氮中速冻。温室中苗期川续断一部分于15 ℃低温处理和23 ℃常温培养, 分别处理6 h、1 d、2 d; 一部分用MS培养液进行水培, 添加茉莉酸甲酯(MeJA) 进行处理, 对照处理加等量的乙醇, 分别处理6 h、1 d、3 d、5 d。处理后采集川续断根, 速冻于液氮中, 所有样品均于-80 ℃保存, 用于总RNA提取。
试剂  植物总RNA提取试剂盒购自普洛麦格(北京) 生物技术有限公司, 焦碳酸二乙酯(DEPC) 购自于西格玛奥德里奇(上海) 贸易有限公司, PrimeScript RT Reagent Kit和TB GreenTM Premix Ex TaqTM Ⅱ购自宝生物工程(大连) 有限公司, 所有引物均由生工生物工程(上海) 股份有限公司合成。
仪器  5810R型低温冷冻离心机(Eppendorf, 德国), Nano‐Drop2000分光光度计(Thermo Fisher Scientific, 美国), CFX96 PCR仪(Bio-Rad, 美国), 高灵敏发光成像系统(Bio-Rad, 美国)。
NAC蛋白序列的获得  从川续断全长转录组数据库中下载川续断NAC转录因子序列, 根据功能注释并利用SMART (http://smart.embl-heidelberg.de/) 分析蛋白结构域, 筛选获得具有NAC保守结构域的候选基因; 在TAIR和NCBI上下载拟南芥(Arabidopsis thaliana)和水稻(Oryza sativa) 的NAC蛋白质序列(AT1G52890, ANAC019; AT3G15500, ANAC055; AT4G27410, ANAC72; AT5G63790, ANAC102; DQ394702.1, OsNAC1; AB028185.1, OsNAC6; AB028183.1, OsNAC4)。
NAC蛋白家族系统进化树分析  利用MEGA-X软件对获得的NAC蛋白序列进行比对, 根据多重序列比对的结果, 在MEGA-X软件中采用邻位相连法(neighbor-joining, NJ) 构建川续断、拟南芥、水稻NAC蛋白家族进化树。
NAC蛋白序列保守motif分析  利用MEME (http://meme-suite.org/) 分析NAC蛋白序列保守的motif, motif数量小于10个, 最大宽度值为200, 最小宽度值为6, 其余参数为默认设置。
川续断DaNAC基因的表达分析  通过川续断全长转录组数据库获得基因序列, 利用Primer5设计引物(表 1) 用于实时荧光定量分析。将-80 ℃保存的川续断样品在研钵中用液氮磨碎, 依据植物总RNA提取试剂盒的方法提取川续断RNA, 根据PrimeScript RT Reagent Kit试剂盒说明书的方法将川续断RNA反转录成cDNA, 利用DaACT103作为内参基因进行荧光定量PCR[15]。反应体系: cDNA 1 μL, TB GreenTM Premix Ex TaqTM Ⅱ 5 μL, Primer-L 0.3 μL, Primer-R 0.3 μL, ddH2O 3.4 μL, PCR反应条件依据TB GreenTM Premix Ex TaqTMⅡ试剂盒说明书, 每组设置3次生物学重复。
结果与分析
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1 DaNAC蛋白进化树分析
从川续断全长转录组数据库中下载川续断NAC转录因子具有完整ORF序列的有29个, 根据功能注释及SMART进行氨基酸结构域分析, 均含有保守NAM结构域。根据进化树结果分析(图 1), 可将川续断NAC蛋白家族分为6个亚家族(Ⅰ~Ⅵ)。其中, Group Ⅰ主要包含部分川续断蛋白及水稻、拟南芥逆境相关NAC蛋白, Group Ⅱ含有4个川续断NAC蛋白, Group Ⅲ含有3个川续断NAC蛋白, Group Ⅳ含有7个川续断NAC蛋白, Group Ⅴ仅含有1个川续断NAC蛋白, Group Ⅵ含有9个川续断NAC蛋白。在川续断中有5个DaNAC蛋白与拟南芥和水稻逆境胁迫相关的NAC蛋白聚在同一分支, 分别是34564NAC2、33883NAC48、32393NAC81、6187NAC14、6727NAC14。
2 NAC蛋白序列保守motif分析
用MEME软件分析川续断DaNAC蛋白序列的保守motif, 根据E-value对川续断DaNAC家族成员蛋白序列的保守motif进行排序, motif 1和motif 2的保守性最强(图 2)。从川续断DaNAC家族成员保守motif结果分析可以看出, 保守motif (motif 1和motif 2) 主要集中分布在DaNAC蛋白序列的N端, 与NAC蛋白保守结构域NAM的位置一致。
3 川续断不同组织中DaNACs基因的表达分析
本文选取4个DaNAC基因, 分析其在川续断不同组织中的表达模式(图 3), 包括与拟南芥和水稻逆境胁迫相关NAC蛋白聚在同一分支, 且较为同源的34564NAC2、33883NAC48、6727NAC14, 以及与其聚在不同分支, 同源性较低的34480NAC22。上述4个DaNAC在川续断不同组织中均有表达, 在叶片中表达量最高, 在花和种子中表达量较低。其中, 6727NAC14在叶片、叶柄、根、茎中表达量较高, 在花、种子中有较低的表达; 34564NAC2、33883NAC48、34480NAC22主要在川续断的叶片中表达, 而在川续断其他组织中的表达量均较低。
4 低温胁迫诱导DaNACs基因的表达分析
进一步分析发现, 4个DaNAC基因均受低温诱导表达(图 4)。与25 ℃正常温度培养的川续断相比, 在15 ℃低温胁迫6 h后, 4个基因的表达量均有不同程度升高。34564NAC2、33883NAC48、6727NAC14主要在15 ℃低温处理前期(6 h) 表达量有明显的升高, 而在15 ℃低温处理1 d后, 表达量升高得不明显; 在低温处理6 h到2 d, 34480NAC22均有一定的上调表达。因此, 结果说明川续断4个DaNAC基因均受低温的诱导上调表达, 但表达模式存在一定的差异。
5 茉莉酸甲酯诱导DaNACs基因的表达分析
150 μmol·L-1 MeJA处理后, 4个DaNAC基因的表达量均有一定的差异(图 5)。与对照相比, MeJA处理6 h至1 d后, 34564NAC2、33883NAC48基因上调表达; 在MeJA处理6 h时, 6727NAC14基因的表达量升高; MeJA处理过程中, 34480NAC22基因均受茉莉酸甲酯诱导下调表达。因此, 结果说明川续断4个NAC基因均受茉莉酸甲酯的诱导表达, 但是不同的基因诱导表达的模式存在差异。
讨论
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川续断皂苷Ⅵ主要在川续断的根中积累。Wang等[16]通过转录组等研究分析其生物合成相关酶的候选基因及川续断三萜皂苷合成途径。但是, 植物次生代谢产物的合成途径往往是一个复杂的调控网络, 转录因子发挥重要的调控作用。本研究通过全长转录组数据库筛选得到29个具有完整ORF的DaNAC转录因子序列, 其中, 有5个DaNAC与拟南芥和水稻中与逆境胁迫相关的NAC较为同源。已有研究证明, 拟南芥ANAC019、ANAC055和ANAC072较为同源, 作为转录因子调控茉莉酸信号并诱导防御基因的表达, 增强植物对逆境胁迫的防御反应[17]; 水稻SNAC1、OsNAC6等基因受茉莉酸、低温、干旱等逆境胁迫的诱导表达, 参与多种生物逆境和非生物逆境胁迫的响应, 过表达SNAC1显著提高水稻对干旱和盐碱的抗性[11]。川续断中这5个与逆境胁迫密切相关的NAC转录因子, 在川续断抵御低温、干旱等逆境胁迫中同样发挥着重要功能。
植物逆境应答是一个复杂的调控网络, 逆境胁迫能促进植物次生代谢产物的积累。研究发现, 在新疆紫草悬浮细胞中, 低温胁迫可促进紫草酸B、紫草呋喃A的积累[18]。在番茄中, 茉莉酸诱导番茄萜烯合成酶基因的表达, 促进番茄中萜烯的生物合成, 提高番茄对逆境胁迫的抗性[19]。本课题组前期研究发现15 ℃低温胁迫[20]、150 μmol·L-1茉莉酸甲酯[10]处理可增加川续断根中川续断皂苷Ⅵ的积累, DaNAC基因参与川续断响应低温及茉莉酸甲酯的逆境胁迫应答。组织表达分析显示, DaNAC基因在川续断根茎叶等组织中均有表达, 但其表达模式不同, 说明川续断对逆境胁迫应答存在一定的组织特异性。进一步对DaNAC基因进行不同非生物逆境诱导模式表达分析, 发现4个DaNAC基因均受低温诱导表达; 34564NAC2、33883NAC48基因受茉莉酸甲酯诱导上调表达, 34480NAC22基因受茉莉酸甲酯诱导下调表达。因此, 推测DaNAC基因参与调控川续断中川续断皂苷Ⅵ等次生代谢产物的合成, 促进川续断对非生物逆境胁迫的抵抗。后续将针对DaNAC基因如何调控川续断皂苷Ⅵ等次生代谢产物的合成开展进一步的研究。
作者贡献: 徐娇设计了本实验, 完成了数据分析、论文初稿的写作; 徐娇、胡正平、李良远和肖承鸿完成了川续断的种植、处理、qRT-PCR等相关实验; 何华、周涛、江维克及郭娟共同指导论文的写作与修改。
利益冲突: 本文的研究内容无任何利益冲突。
基金
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  • 国家自然科学基金资助项目(81860675)
  • 中央本级重大增减支项目(2060302)
  • 贵州省科技计划(黔科合基础[2020] 1Y370)
  • 贵州省高层次创新人才项目(黔科合平台人才[2018] 5638-2)
  • 贵州省科技创新人才团队项目(黔科合平台人才[2019] 5611)
  • 贵州省教育厅重大研究项目(黔教合KY字[2018] 022)
文献
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2021年第56卷第12期
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doi: 10.16438/j.0513-4870.2021-0899
  • 接收时间：2021-06-21
  • 首发时间：2025-12-18
  • 出版时间：2021-12-12
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  • 收稿日期：2021-06-21
  • 修回日期：2021-08-09
基金
国家自然科学基金资助项目(81860675)
中央本级重大增减支项目(2060302)
贵州省科技计划(黔科合基础[2020] 1Y370)
贵州省高层次创新人才项目(黔科合平台人才[2018] 5638-2)
贵州省科技创新人才团队项目(黔科合平台人才[2019] 5611)
贵州省教育厅重大研究项目(黔教合KY字[2018] 022)
作者信息
    1.贵州中医药大学, 贵州 贵阳 550025
    2.中国中医科学院中药资源中心, 北京 100700

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*周涛，Tel: 13985175215, 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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