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Article(id=1210518244239217288, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1210518228766421884, articleNumber=null, orderNo=null, doi=10.16438/j.0513-4870.2022-0666, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1654185600000, receivedDateStr=2022-06-03, revisedDate=1655049600000, revisedDateStr=2022-06-13, acceptedDate=null, acceptedDateStr=null, onlineDate=1766539639768, onlineDateStr=2025-12-24, pubDate=1670774400000, pubDateStr=2022-12-12, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1766539639768, onlineIssueDateStr=2025-12-24, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1766539639768, creator=13701087609, updateTime=1766539639768, updator=13701087609, issue=Issue{id=1210518228766421884, tenantId=1146029695717560320, journalId=1189982191388893191, year='2022', volume='57', issue='12', pageStart='0', pageEnd='3698', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1766539636078, creator=13701087609, updateTime=1766539730802, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1210518626109624560, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1210518228766421884, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1210518626109624561, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1210518228766421884, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=3686, endPage=3694, ext={EN=ArticleExt(id=1210518245900161721, articleId=1210518244239217288, tenantId=1146029695717560320, journalId=1189982191388893191, language=EN, title=Overexpression of (3S)-linalool synthase gene (LIS) regulates the glycyrrhizic acid biosynthesis in Glycyrrhiza uralensis hairy roots, columnId=1190335348761793317, journalTitle=Acta Pharmaceutica Sinica, columnName=Original Articles, runingTitle=null, highlight=null, articleAbstract=

(3S)-Linalool synthase (LIS) is a key enzyme involved in the monoterpene biosynthetic pathway. Based on our previous transcriptome study, the expression level of LIS gene was exceedingly related to glycyrrhizic acid (GA) biosynthesis. Therefore, we used hairy root culturing to further investigate the effect of LIS on the GA biosynthesis. A LIS gene (GenBank accession number: MZ169552) was cloned from Glycyrrhiza uralensis. The plant binary overexpression vector pCA-LIS was constructed by gene fusion. G. uralensis hairy roots overexpressing LIS were induced by the Agrobacterium rhizogenes ATCC15834. The expression levels of LIS were analyzed by real-time quantitative PCR (RT-qPCR) and the contents of GA in hairy root lines were determined by UPLC. It was found that in the hairy root lines overexpressing LIS, the expression levels of LIS were significantly higher than that in the wild type, while the contents of GA were remarkably lower than those in the wild type and negative control. These findings indicate that the expression level of LIS is negatively correlated with the accumulation of GA. In this study, LIS was cloned from G. uralensis for the first time and the negative regulatory effect of LIS on GA biosynthesis was confirmed by reverse genetics. This work provides support for further improvement of the molecular regulatory network of GA biosynthesis in G. uralensis.

, correspAuthors=Ping HE, Ying LIU, authorNote=null, correspAuthorsNote=null, copyrightStatement=Copyright ©2022 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=Ling-yu KE, Zi-yi CHEN, Wen-wen DING, Zhi-xin ZHANG, Ping HE, Ying LIU), CN=ArticleExt(id=1210518250388067125, articleId=1210518244239217288, tenantId=1146029695717560320, journalId=1189982191388893191, language=CN, title=过表达LIS基因对甘草毛状根中甘草酸生物合成的调控研究, columnId=1190335348896011050, journalTitle=药学学报, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=

芳樟醇合酶[(3S)-linalool synthase, LIS] 是单萜合成途径上的关键酶。本课题组前期在甘草转录组研究中发现LIS基因与甘草酸(glycyrrhizic acid, GA) 的生物合成密切相关, 因此本研究拟利用甘草毛状根培养体系, 进一步解析LIS对甘草酸生物合成的调控作用。本研究克隆了甘草LIS基因(GenBank注册号: MZ169552); 采用基因融合法构建过表达LIS基因的植物双元表达载体pCA-LIS; 利用发根农杆菌(Agrobacterium rhizogenes ATCC15834) 介导法诱导甘草毛状根系; 通过实时荧光定量PCR (RT-qPCR) 进行LIS基因的表达水平分析; 最终采用UPLC法测定各甘草毛状根系中的甘草酸含量。结果显示, 过表达LIS基因的甘草毛状根系中, LIS基因的表达水平显著高于野生型, 而甘草酸含量则显著低于野生型和阴性对照组, 表明LIS基因的表达水平与甘草酸含量呈负相关。本研究首次克隆得到甘草LIS基因, 通过反向遗传学策略证实了LIS基因对甘草酸生物合成的负调控作用, 为进一步完善甘草酸生物合成的分子调控网络提供了依据。

, correspAuthors=何平, 刘颖, authorNote=null, correspAuthorsNote=
*何平, Tel: 86-10-53912163, E-mail: ;
刘颖, E-mail:
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Bot Lett, 2020, 167: 219-231., articleTitle=Overexpressing chalcone synthase (CHS) gene enhanced flavonoids accumulation in Glycyrrhiza uralensis hairy roots, refAbstract=null)], funds=null, companyList=[AuthorCompany(id=1210518250606170948, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, xref=null, ext=[AuthorCompanyExt(id=1210518250614559557, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, companyId=1210518250606170948, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=School of Life Sciences, Beijing University of Chinese Medicine, Beijing 102488, China), AuthorCompanyExt(id=1210518250622948166, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, companyId=1210518250606170948, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=北京中医药大学生命科学学院, 北京 102488)])], figs=[ArticleFig(id=1210518255219904557, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=EN, label=null, caption=null, figureFileSmall=VpoOr49HloLaRCS0LjjZjA==, figureFileBig=FaJgSWc/JVmafEN7exF97A==, tableContent=null), ArticleFig(id=1210518255312179253, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=CN, label=Figure 1, caption= The biosynthetic pathway of triterpenoid and monoterpenoid in <i>G. uralensis</i>. MVA: Mevalonate; MEP: 2-<i>C</i>-methyl-<i>D</i>-erythritol 4-phosphate; IDI: Isopentenyl-diphosphate delta-isomerase; GGPS: Geranylgeranyl pyrophosphate synthase; LIS: (3<i>S</i>)-Linalool synthase , figureFileSmall=VpoOr49HloLaRCS0LjjZjA==, figureFileBig=FaJgSWc/JVmafEN7exF97A==, tableContent=null), ArticleFig(id=1210518255547060286, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=EN, label=null, caption=null, figureFileSmall=FBI8UfizyA46lEqgvbUIGA==, figureFileBig=21ZWOTIyISsK88KbAQ3a7g==, tableContent=null), ArticleFig(id=1210518255672889416, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=CN, label=Figure 2, caption= The cloning of <i>LIS</i> gene from <i>G. uralensis</i> and the bioinformatic analysis of LIS. a: The PCR verification result of <i>LIS</i> (M: DNA marker). b and c: The predicted secondary and 3D structures of LIS. d: The phylogenetic tree constructed based on 20 TPS orthologs , figureFileSmall=FBI8UfizyA46lEqgvbUIGA==, figureFileBig=21ZWOTIyISsK88KbAQ3a7g==, tableContent=null), ArticleFig(id=1210518255786135625, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=EN, label=null, caption=null, figureFileSmall=B7rLhLPcftlabSROE6Sapw==, figureFileBig=JBTE94IE2hCGGOBIS3QDPw==, tableContent=null), ArticleFig(id=1210518255882604622, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=CN, label=Figure 3, caption= Construction of the recombinant overexpression vector pCA-<i>LIS</i> , figureFileSmall=B7rLhLPcftlabSROE6Sapw==, figureFileBig=JBTE94IE2hCGGOBIS3QDPw==, tableContent=null), ArticleFig(id=1210518255983267922, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=EN, label=null, caption=null, figureFileSmall=QT2NZOs0ishXlcG+jAmTzA==, figureFileBig=Mpmwjj3hc5mFarshG/0hZQ==, tableContent=null), ArticleFig(id=1210518256079736917, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=CN, label=Figure 4, caption= The induction and identification of <i>G. uralensis</i> hairy roots. a: <i>LIS</i> amplified from vector pCA-<i>LIS</i>. b: The <i>G. uralensis</i> hairy roots after induction and culturing for 7 d and 28 d. WT: Wild type hairy roots, NC: Negative control hairy roots containing empty pCAMBIA1305.1. <i>LIS</i><sup>+</sup>: Hairy roots overexpressing <i>LIS</i>. c: <i>rolC</i> amplified from hairy roots (Lane 1: WT, 2: NC, and 3: <i>LIS</i><sup>+</sup>). d: The 500 bp-fragments (<i>GUS</i> + <i>LIS</i>) amplified from <i>LIS</i><sup>+</sup> hairy roots , figureFileSmall=QT2NZOs0ishXlcG+jAmTzA==, figureFileBig=Mpmwjj3hc5mFarshG/0hZQ==, tableContent=null), ArticleFig(id=1210518256188788827, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=EN, label=null, caption=null, figureFileSmall=RwCloV7FlLbOiucVDP7qPA==, figureFileBig=ZZ3LHpJY1IReh35AQnzRLA==, tableContent=null), ArticleFig(id=1210518256348172381, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=CN, label=Figure 5, caption= The relative expression levels of <i>LIS</i> in different <i>G. uralensis</i> hairy root lines (<i>n</i> = 3). <sup>*</sup><i>P</i> < 0.05, <sup>**</sup><i>P</i> < 0.01, <sup>***</sup> <i>P</i> < 0.001, <i>vs</i> WT , figureFileSmall=RwCloV7FlLbOiucVDP7qPA==, figureFileBig=ZZ3LHpJY1IReh35AQnzRLA==, tableContent=null), ArticleFig(id=1210518256436252769, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=EN, label=null, caption=null, figureFileSmall=p27SU+DIw6OkLazDpnOdgw==, figureFileBig=qru8i+YgeG7qNYjJFfXc6A==, tableContent=null), ArticleFig(id=1210518256528527463, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=CN, label=Figure 6, caption= GA content analyses in <i>G. uralensis</i> hairy roots. a: Hairy root samples culturing in liquid 6, 7-V media for 21 d. b: UPLC chromatograms. Lines 1-4 show the UPLC chromatograms of reference substance GA and the WT, NC, and <i>LIS</i><sup>+</sup> hairy root lines. c: GA content analyses in <i>G. uralensis</i> hairy root lines (<i>n</i> = 3). <sup><i>**</i></sup><i>P</i> < 0.01, <sup><i>***</i></sup><i>P</i> < 0.001 <i>vs</i> WT; <sup>△△△</sup><i>P</i> < 0.001 <i>vs</i> NC , figureFileSmall=p27SU+DIw6OkLazDpnOdgw==, figureFileBig=qru8i+YgeG7qNYjJFfXc6A==, tableContent=null), ArticleFig(id=1210518256608219239, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
No.ProjectPrimerSequence (5′-3′)
ⅠLIS cloningLIS-F1ATGTCTACTGCAAATATAATGGCTG
LIS-R1TCAATAGCTAGGAAGAGATGTGAAG
ⅡConstruction of LIS overexpression vectorLIS-F2CTCTTGACCATGGTAGATCTATGTCTACTGCAAATATAATGGCTG
LIS-R2TTGATCGGGTACAGACTAGTTCAATAGCTAGGAAGAGATGTGAAG
ⅢVerification of rolC in hairy rootsR-FCATATATGCCAAATTTACACTAG
R-RGTTAACAAACTAGGAAACAGG
ⅣIdentification of exogenous LIS in transgenic hairy rootsGL-FGCTTGCAGCATACAATGCCT
GL-RCCGAAGCGGAGCACGATAC
Ⅴβ-Actin expression detection by RT-qPCRqB-FCAAAAGGATGCCTATGTGGG
qB-RCAGGAGCAACACGCAATTC
ⅥLIS expression detection by RT-qPCRqLIS-FAGTTGGGACTTGTTGACA
qLIS-RTCATCCAATGTGCCATAAGTGT
), ArticleFig(id=1210518256717271151, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=CN, label=Table 1, caption=

Primer sequences used in this study

, figureFileSmall=null, figureFileBig=null, tableContent=
No.ProjectPrimerSequence (5′-3′)
ⅠLIS cloningLIS-F1ATGTCTACTGCAAATATAATGGCTG
LIS-R1TCAATAGCTAGGAAGAGATGTGAAG
ⅡConstruction of LIS overexpression vectorLIS-F2CTCTTGACCATGGTAGATCTATGTCTACTGCAAATATAATGGCTG
LIS-R2TTGATCGGGTACAGACTAGTTCAATAGCTAGGAAGAGATGTGAAG
ⅢVerification of rolC in hairy rootsR-FCATATATGCCAAATTTACACTAG
R-RGTTAACAAACTAGGAAACAGG
ⅣIdentification of exogenous LIS in transgenic hairy rootsGL-FGCTTGCAGCATACAATGCCT
GL-RCCGAAGCGGAGCACGATAC
Ⅴβ-Actin expression detection by RT-qPCRqB-FCAAAAGGATGCCTATGTGGG
qB-RCAGGAGCAACACGCAATTC
ⅥLIS expression detection by RT-qPCRqLIS-FAGTTGGGACTTGTTGACA
qLIS-RTCATCCAATGTGCCATAAGTGT
), ArticleFig(id=1210518256780185717, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
No.ProjectPCR program
1LIS cloning94 ℃ 5 min, 35 cycles of 94 ℃ 30 s, 50 ℃ 30 s, 72 ℃ 100 s, and 72 ℃ 5 min
2Construction of LIS overexpression vector94 ℃ 5 min, 35 cycles of 94 ℃ 30 s, 50 ℃ 30 s, 72 ℃ 100 s, and 72 ℃ 5 min
3Verification of rolC in hairy roots95 ℃ 5 min, 34 cycles of 95 ℃ 30 s, 56 ℃ 30 s, 72 ℃ 60 s, and 72 ℃ 5 min
4Identification of exogenous LIS in transgenic hairy roots95 ℃ 5 min, 34 cycles of 95 ℃ 30 s, 55 ℃ 30 s, 72 ℃ 60 s, and 72 ℃ 5 min
5β-Actin expression detection by RT-qPCR95 ℃ 1 min, 40 cycles of 95 ℃ 20 s, 60 ℃ 1 min
6LIS expression detection by RT-qPCR95 ℃ 1 min, 40 cycles of 95 ℃ 20 s, 60 ℃ 1 min
), ArticleFig(id=1210518256906014842, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=CN, label=Table 2, caption=

PCR programs used in this study

, figureFileSmall=null, figureFileBig=null, tableContent=
No.ProjectPCR program
1LIS cloning94 ℃ 5 min, 35 cycles of 94 ℃ 30 s, 50 ℃ 30 s, 72 ℃ 100 s, and 72 ℃ 5 min
2Construction of LIS overexpression vector94 ℃ 5 min, 35 cycles of 94 ℃ 30 s, 50 ℃ 30 s, 72 ℃ 100 s, and 72 ℃ 5 min
3Verification of rolC in hairy roots95 ℃ 5 min, 34 cycles of 95 ℃ 30 s, 56 ℃ 30 s, 72 ℃ 60 s, and 72 ℃ 5 min
4Identification of exogenous LIS in transgenic hairy roots95 ℃ 5 min, 34 cycles of 95 ℃ 30 s, 55 ℃ 30 s, 72 ℃ 60 s, and 72 ℃ 5 min
5β-Actin expression detection by RT-qPCR95 ℃ 1 min, 40 cycles of 95 ℃ 20 s, 60 ℃ 1 min
6LIS expression detection by RT-qPCR95 ℃ 1 min, 40 cycles of 95 ℃ 20 s, 60 ℃ 1 min
), ArticleFig(id=1210518256977318014, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
Time/minAcetonitrile (A)/%0.05% Phosphoric acid (B)/%
01486
1.362377
3.263070
4.083466
4.763664
5.714258
6.535149
95149
9.51486
121486
), ArticleFig(id=1210518257082175615, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1210518244239217288, language=CN, label=Table 3, caption=

Program of UPLC gradient elution

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Time/minAcetonitrile (A)/%0.05% Phosphoric acid (B)/%
01486
1.362377
3.263070
4.083466
4.763664
5.714258
6.535149
95149
9.51486
121486
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过表达LIS基因对甘草毛状根中甘草酸生物合成的调控研究
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柯铃钰 , 陈姿伊 , 丁文文 , 张智新 , 何平 * , 刘颖 *
药学学报 | 研究论文 2022,57(12): 3686-3694
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药学学报 | 研究论文 2022, 57(12): 3686-3694
过表达LIS基因对甘草毛状根中甘草酸生物合成的调控研究
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柯铃钰, 陈姿伊, 丁文文, 张智新, 何平* , 刘颖*
作者信息
  • 北京中医药大学生命科学学院, 北京 102488

通讯作者:

*何平, Tel: 86-10-53912163, E-mail: ;
刘颖, E-mail:
Overexpression of (3S)-linalool synthase gene (LIS) regulates the glycyrrhizic acid biosynthesis in Glycyrrhiza uralensis hairy roots
Ling-yu KE, Zi-yi CHEN, Wen-wen DING, Zhi-xin ZHANG, Ping HE* , Ying LIU*
Affiliations
  • School of Life Sciences, Beijing University of Chinese Medicine, Beijing 102488, China
出版时间: 2022-12-12 doi: 10.16438/j.0513-4870.2022-0666
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芳樟醇合酶[(3S)-linalool synthase, LIS] 是单萜合成途径上的关键酶。本课题组前期在甘草转录组研究中发现LIS基因与甘草酸(glycyrrhizic acid, GA) 的生物合成密切相关, 因此本研究拟利用甘草毛状根培养体系, 进一步解析LIS对甘草酸生物合成的调控作用。本研究克隆了甘草LIS基因(GenBank注册号: MZ169552); 采用基因融合法构建过表达LIS基因的植物双元表达载体pCA-LIS; 利用发根农杆菌(Agrobacterium rhizogenes ATCC15834) 介导法诱导甘草毛状根系; 通过实时荧光定量PCR (RT-qPCR) 进行LIS基因的表达水平分析; 最终采用UPLC法测定各甘草毛状根系中的甘草酸含量。结果显示, 过表达LIS基因的甘草毛状根系中, LIS基因的表达水平显著高于野生型, 而甘草酸含量则显著低于野生型和阴性对照组, 表明LIS基因的表达水平与甘草酸含量呈负相关。本研究首次克隆得到甘草LIS基因, 通过反向遗传学策略证实了LIS基因对甘草酸生物合成的负调控作用, 为进一步完善甘草酸生物合成的分子调控网络提供了依据。

甘草  /  甘草酸  /  芳樟醇合酶  /  基因过表达  /  毛状根

(3S)-Linalool synthase (LIS) is a key enzyme involved in the monoterpene biosynthetic pathway. Based on our previous transcriptome study, the expression level of LIS gene was exceedingly related to glycyrrhizic acid (GA) biosynthesis. Therefore, we used hairy root culturing to further investigate the effect of LIS on the GA biosynthesis. A LIS gene (GenBank accession number: MZ169552) was cloned from Glycyrrhiza uralensis. The plant binary overexpression vector pCA-LIS was constructed by gene fusion. G. uralensis hairy roots overexpressing LIS were induced by the Agrobacterium rhizogenes ATCC15834. The expression levels of LIS were analyzed by real-time quantitative PCR (RT-qPCR) and the contents of GA in hairy root lines were determined by UPLC. It was found that in the hairy root lines overexpressing LIS, the expression levels of LIS were significantly higher than that in the wild type, while the contents of GA were remarkably lower than those in the wild type and negative control. These findings indicate that the expression level of LIS is negatively correlated with the accumulation of GA. In this study, LIS was cloned from G. uralensis for the first time and the negative regulatory effect of LIS on GA biosynthesis was confirmed by reverse genetics. This work provides support for further improvement of the molecular regulatory network of GA biosynthesis in G. uralensis.

Glycyrrhiza uralensis  /  glycyrrhizic acid  /  (3S)-linalool synthase  /  overexpression of gene  /  hairy root
柯铃钰, 陈姿伊, 丁文文, 张智新, 何平, 刘颖. 过表达LIS基因对甘草毛状根中甘草酸生物合成的调控研究. 药学学报, 2022 , 57 (12) : 3686 -3694 . DOI: 10.16438/j.0513-4870.2022-0666
Ling-yu KE, Zi-yi CHEN, Wen-wen DING, Zhi-xin ZHANG, Ping HE, Ying LIU. Overexpression of (3S)-linalool synthase gene (LIS) regulates the glycyrrhizic acid biosynthesis in Glycyrrhiza uralensis hairy roots[J]. Acta Pharmaceutica Sinica, 2022 , 57 (12) : 3686 -3694 . DOI: 10.16438/j.0513-4870.2022-0666
正文
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甘草(Glycyrrhiza uralensis) 始载于《神农本草经》, 是我国最常用的大宗药材之一, 具有清热解毒、缓急止痛、调和诸药等功效[1]。现代药理学研究表明, 甘草中的三萜类化合物甘草酸(glycyrrhizic acid, GA) 是其发挥功效的主要活性成分, 已证实其具有抗癌[2]、抗炎[3]、抗病毒[4]等多种药理活性, 因此历版《中华人民共和国药典》均将其规定为中药材甘草的检测指标[1]。然而, 本课题组前期调查显示, 约50%市售甘草药材中甘草酸含量未达到国家药典规定的最低标准(2.0%)[5]。因此, 提高栽培甘草中甘草酸含量对于保障甘草药材质量具有重要意义。
高等植物中萜类化合物的生物合成始于一对同分异构体, 即: 二甲基烯丙基焦磷酸(dimethylallyl pyrophosphate, DMAPP) 和异戊烯基焦磷酸(isopentenyl pyrophosphate, IPP), 它们经连续缩合形成不同萜类化合物的前体, 再通过各种骨架的排列及修饰, 从而产生类异戊二烯结构和功能的多样性[6]。如图 1所示, IPP和DMAPP是由细胞质中的甲羟戊酸(mevalonate, MVA) 途径和质体中的2-C-甲基-D-赤藻糖-4-磷酸(2-C-methyl-D-erythritol 4-phosphate, MEP) 途径共同合成的[7]。通常认为, 来源于MVA途径的DMAPP和IPP主要用于合成倍半萜、三萜、甾醇等化合物, 而MEP途径则提供DMAPP和IPP用于合成单萜、二萜、四萜等化合物[8]。甘草酸属于三萜类化合物, 由MVA途径合成, 但同时受MEP途径的影响[9]。DMAPP和IPP通过牻牛儿牻牛儿基焦磷酸合酶(geranylgeranyl pyrophosphate synthase, GGPS) 缩合生成单萜途径前体牻牛儿基焦磷酸(geranyl pyrophosphate, GPP), 而芳樟醇合酶[(3S)-linalool synthase, LIS] 是催化GPP进一步形成芳樟醇[(+)-linalool] 的关键酶[10]。研究显示, 在金宣茶(C. sinensis cv. Jinxuan) 和樟树(Cinnamomum camphora) 中芳樟醇的释放量与LIS的表达量均呈正相关[11, 12]。LIS属于萜烯合酶(terpene synthase, TPS) 家族, 目前在GenBank中已注册4 200多条植物TPS基因序列, 包括: 北美云杉(Picea sitchensis)[13]、月季(Rosa chinensis)[14]、小麦(Triticum aestivum)[15]、甜豌豆(Lathyrus odoratus)[16]等。本课题组前期开展了甘草的转录组研究, 挖掘到影响甘草酸生物合成的15个核心差异表达基因[17], LIS即为其中之一。然而, 在甘草中尚未克隆得到该基因。因此, 本研究拟克隆甘草LIS基因, 并解析其对甘草酸生物合成的调控作用。
植物毛状根的形成是由发根农杆菌(Agrobaeterium rhizogenes) 侵染而产生的一种病理表现。由于毛状根具有生长快速稳定[18]、无需额外激素[19]、可合成植物次生代谢产物[20]等优点, 已成为研究植物基因功能的良好遗传转化体系, 如: 在苦荞(Fagopyrum tataricum) 毛状根中过表达FtF3GT1基因可显著提高毛状根中总黄酮的含量[21]; 在西伯利亚益母草(Leonurus sibiricus L.) 毛状根中过表达AtPAP1基因则可诱导高水平酚酸的积累[22]。就甘草而言, 本课题组前期已利用毛状根体系对甘草转录组数据中挖掘到的生长素应答蛋白基因(auxin-responsive protein IAA gene, ARPI)、β-香树酯醇合成酶基因(β-amyrin synthase gene, β-AS)、UDP-葡萄糖4-差向异构酶基因(UDP-galactose/glucose-4-epimerase gene, UGE)、查尔酮异构酶基因(chalcone isomerase, CHI)、阿魏酸5-羟化酶基因(ferulate 5-hydroxylase, F5H) 等陆续开展了研究, 解析了这些功能基因对于甘草酸生物合成的调控作用[9, 23-25]。因此, 本研究也将利用毛状根体系对LIS的功能开展研究, 解析该基因在甘草萜类化合物代谢中的作用, 为进一步构建甘草酸生物合成的分子调控网络提供依据。
材料与方法
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试剂与耗材  DNA/RNA一站式提取试剂盒购于上海生工生物工程股份有限公司; 大肠杆菌DH5α感受态细胞、GoldenView、DNA Marker (BM2000、BM15000)、高纯质粒小量快速提取试剂盒(离心柱型)、BM无缝克隆试剂盒、琼脂糖凝胶纯化回收试剂盒均购于北京博迈德科技发展有限公司; 卡那霉素(kanamycin, Kan) 购于Genview公司; pMD-19T载体、限制性内切酶Bgl II、Spe I等购于TaKaRa公司; 头孢噻肟钠(cefotaxime sodium, Cef) 和氨苄青霉素(ampicillin, Amp) 购于北京拜尔迪生物技术有限公司; 甲醇(分析纯) 购于天津西典化学科技有限公司; 乙腈(色谱纯) 购于赛默飞世尔科技(中国) 有限公司; 甘草酸单铵盐(MUST-16011310) 购于成都曼斯特生物科技有限公司, 纯度为99.45%; NovoStart® SYBR qPCR SuperMix Plus、NovoScript® Plus All-in-one 1st Strand cDNA Synthesis SuperMix (gDNA Purge) 购于上海近岸科技有限公司。
仪器  A300型PCR扩增仪(杭州朗基科学仪器有限公司); 恒温振荡培养箱DLHR-Q200 (北京东联哈尔滨仪器制造有限公司); 1-13000型离心机(美国Sigma公司); Gene PulserMXcell穿孔仪(美国BIO-RAD公司); DYY-8型稳压稳流电泳仪(上海琪特分析仪器有限公司); 凝胶成像系统(北京百晶生物技术有限公司)。
PCR引物及程序  引物合成及测序均由上海生工生物工程股份有限公司完成。所采用的全部引物信息如表 1所示, 全部PCR程序如表 2所示。
甘草LIS基因克隆  采用RNA提取试剂盒从新鲜甘草根样中提取总RNA, 采用cDNA合成试剂盒逆转录获得cDNA。利用NCBI在线工具PrimerBLAST (https://blast.ncbi.nlm.nih.gov/Blast.cgi) 设计引物I (表 1), 采用PCR程序1 (表 2) 对LIS基因进行扩增, 其25 μL体系为: cDNA 1.0 μL, LA Taq酶0.25 μL, LIS-F1 (10 μmol·L-1) 0.5 μL, LIS-R1 (10 μmol·L-1) 0.5 μL, dNTPs (2.5 μmol·L-1) 2.0 μL, 10×LA PCR buffer (Mg2+) 2.5 μL, ddH2O 18.25 μL。对PCR产物进行胶回收纯化, 连入pMD-19T载体, 16 ℃反应过夜。通过热激法将连接产物转化入大肠杆菌DH5α感受态细胞, 在LB (含Amp 50 mg·L-1) 平板上筛选阳性克隆, 并进行PCR验证, 将验证正确的菌液送测序。
甘草LIS生物信息学分析  采用EsPASy-ProtParam对甘草LIS序列进行理化性质分析; 采用SOPMA对甘草LIS进行二级结构分析; 采用Swiss-Model构建甘草LIS三级结构模型; 采用SignalP 6.0 Server对甘草LIS进行信号肽预测; 采用NCBI的CD-search对甘草LIS中的保守结构域进行预测; 从GenBank数据库中挑选19条来自不同种属的萜烯合酶蛋白序列, 使用MEGA 11软件对以上序列和本文克隆获得的甘草LIS进行聚类分析, 采用近邻聚类算法(neighbor-joining, NJ) 构建系统进化树。
过表达LIS植物双元表达载体的构建  在pCAMBIA1305.1载体上, 于Bgl II及Spe I酶切位点插入LIS基因, 具体步骤如下: 依据BM无缝克隆试剂盒设计引物II (表 1), 采用PCR程序2 (表 2) 扩增带有载体同源臂的目标基因; 采用限制性内切酶Bgl II和Spe I在37 ℃酶切pCAMBIA1305.1质粒1 h; 对扩增获得的目标基因与线性化载体进行胶回收纯化; 采用BM无缝克隆试剂盒对以上二者(载体与基因的摩尔比为1∶3) 进行连接, 50 ℃、30 min; 通过热激法将连接产物转入大肠杆菌DH5α感受态细胞中, 在含有Kan (50 mg·L-1) 的LB平板上筛选阳性克隆; 对阳性克隆进行PCR及测序验证; 将验证正确的质粒命名为pCA-LIS。
重组发根农杆菌ATCC15834的构建  采用电转法(C: 25 μF, PC: 200 Ω, U: 2 400 V) 将重组质粒pCA-LIS导入发根农杆菌ATCC15834感受态细胞, 在TY平板(含Kan 50 mg·L-1) 上筛选阳性克隆。采用引物I (表 1) 和PCR程序1 (表 2) 对阳性菌落进行PCR验证, 然后送测序验证。将验证正确的工程菌接种于TY液体培养基中, 在25 ℃、180 r·min-1条件下振荡培养至对数生长期, 离心后重悬于等体积的6, 7-V液体培养基[26]中。
过表达LIS基因甘草毛状根的诱导及验证  在超净工作台中切取甘草无菌苗胚轴作为外植体材料, 浸泡于上述6, 7-V重悬菌液中, 侵染30 min后转接于6, 7-V平板, 黑暗条件下共培养2天。用Cef水溶液(500 mg·L-1) 浸泡外植体5 min, 无菌水冲洗后重新接种于含Cef (500 mg·L-1) 的6, 7-V平板上, 诱导过表达LIS基因甘草毛状根。同时, 采用发根农杆菌ATCC15834诱导野生型甘草毛状根, 采用携带pCAMBIA1305.1空质粒的发根农杆菌诱导阴性对照组甘草毛状根。每7天转接继代除菌, 逐代降低Cef的浓度, 直至发根农杆菌完全除净。使用DNA提取试剂盒提取各甘草毛状根系DNA, 采用引物III (表 1) 和PCR程序3 (表 2) 对毛状根样品的rolC基因进行PCR验证; 采用引物IV (表 1) 和PCR程序4 (表 2) 对过表达LIS基因甘草毛状根样品进行PCR验证。
甘草毛状根的液体培养  对以上验证正确的甘草毛状根进行液体培养, 步骤如下: 在无菌条件下称取生长状态良好的各甘草毛状根系样品2.0 g, 转接入6, 7-V液体培养基中, 每根系3重复, 25 ℃、110 r·min-1条件下振荡培养, 每隔1周换液1次, 3周后收集毛状根样品, 洗净, 用于后续LIS基因表达水平及甘草酸含量测定。
甘草毛状根中LIS的表达水平分析  使用RNA提取试剂盒提取上述各毛状根系的总RNA, 采用一步法逆转录合成cDNA, 其反应体系为: 甘草RNA 6.0 μL, gDNA Purge 1.0 μL, NovoScript® Plus 1st Strand cDNA Synthesis SuperMix 10.0 μL, RNase-Free H2O 3.0 μL, 反应条件为: 50 ℃下孵育15 min, 75 ℃再孵育5 min, 以甘草β-actin基因作为内参, 利用RT-qPCR对甘草毛状根样品中LIS基因的表达水平进行分析。采用引物V (表 1) 和PCR程序5 (表 2) 对β-actin进行扩增, 采用引物VI (表 1) 和PCR程序6 (表 2) 对LIS基因进行扩增, 反应体系为: 基因1.0 μL, 上游引物(10 μmol·L-1) 1.0 μL, 下游引物(10 μmol·L-1) 1.0 μL, 2×NovoStart® SYBR qPCR SuperMix Plus 5.0 μL, ddH2O 2.0 μL。采用2-△△CT法计算LIS基因的相对表达水平。
UPLC法测定甘草毛状根中甘草酸含量  采用本课题组前期建立的UPLC法对各甘草毛状根样品中的甘草酸进行含量测定[27]。供试品制备: 将毛状根样品在60 ℃下烘干至恒重, 粉碎后过60目筛; 精密称取各甘草毛状根样品粉末0.1 g, 置于50 mL量瓶中, 加入50%甲醇水溶液, 超声提取30 min (频率40 kHz, 功率500 W); 冷却后用50%甲醇水溶液补至50 mL, 经0.45 μm微孔滤膜过滤, 取续滤液作为UPLC供试品。线性: 精密称取甘草酸单铵盐4.52 mg, 用50%甲醇水溶液配制成质量浓度为0.085 8 mg‧mL-1的贮备液, 并稀释成浓度为0.085 8、0.068 6、0.051 5、0.042 9、0.017 2、0.008 58、0.004 3 mg‧mL-1的梯度溶液进行线性分析。甘草酸含量测定: 采用Waters UPLC ACQUITY色谱系统(配备UPLC ACQUITY PDA eλ检测器) 和ACQUITY UPLC BEH C18色谱柱(2.1 mm × 100 mm, 1.7 μm), 以乙腈(A)-0.05%磷酸液(B) 为流动相进行梯度洗脱, 程序见表 3, 柱温为40 ℃、流速为0.3 mL·min-1、进样量为1 μL, 于250 nm波长检测甘草酸。
结果与分析
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1 基因克隆与生物信息学分析
由图 2a可知, PCR扩增得到了长度约为1 800 bp的特异性条带, 测序结果显示, 该条带实际长度为1 837 bp, 其中开放阅读框(open reading frame, ORF) 全长为1 743 bp, 使用在线BLAST比对显示该片段与豆科大豆Glycine max的LIS cDNA序列相似度为83.76%, 其氨基酸序列的相似度为74.74%, 表明所克隆序列为甘草LIS基因序列。在GenBank上对所获序列进行注册, 注册号为: MZ169552。由甘草LIS蛋白序列理化性质分析可知: 该蛋白分子式为C2936H4620N806O870S32; 理论分子质量为66 155.78 Da; 等电点为6.36; 不稳定指数为45.19 (> 40), 可初步判断其为不稳定蛋白; 在哺乳动物网织红细胞内的最长半衰期为30 h; 亲水性为-0.295 (< 0), 可认为其是亲水性蛋白。二级结构分析如图 2b所示, 甘草LIS含有69.41%的α-螺旋(蓝色)、3.79%的延长链(红色) 和27.07%的无规卷曲(黄色), 三级结构模型如图 2c所示。跨膜区域预测结果表明该蛋白高概率位于膜外; 信号肽预测结果显示, 对信号肽和切割位点的预测值均为0, 说明其结构中不含信号肽; 保守结构域预测结果显示, 甘草LIS属于萜烯合酶/环化酶家族蛋白, 具有Terpene_cyclase_plant_C1保守结构域。聚类分析结果如图 2d所示: 裸子植物(紫色)、单子叶植物(绿色)、双子叶植物(橙色) 的TPS序列各自聚为一支; 甘草LIS序列(红色框) 与豆科植物红豆(Abrus precatorius)、野生大豆(Glycine soja)、密花豆(Spatholobus suberectus) 的TPS序列相距最近, 而与3种裸子植物相距最远, 此聚类结果与各物种的亲缘关系相一致。
2 过表达LIS基因甘草毛状根的诱导及验证
按照图 3所示方法构建过表达甘草LIS基因的植物双元表达载体pCA-LIS, 对其进行PCR验证, 结果如图 4a所示: 扩增获得了长度约为1 800 bp的片段, 与目标序列LIS长度相符, 进一步测序结果显示该片段与上述注册序列甘草LIS (MZ169552) 具有100%的一致性, 表明携带甘草LIS基因的双元表达载体构建成功。图 4b是侵染后第7和28天的甘草毛状根样品, 包括: 野生型甘草毛状根(wild type, WT)、含pCAMBIA1305.1空质粒的阴性对照毛状根(negative control, NC) 和过表达LIS基因的甘草毛状根(LIS+), 均长势良好。图 4c为各甘草毛状根系中rolC基因的PCR验证结果, 扩增获得了长度约为600 bp的条带, 测序结果显示其与发根农杆菌rolC基因(DQ160187.1) 具有100%的一致性。图 4d为LIS+毛状根中外源基因的PCR扩增结果, 扩增获得了长度约为500 bp的条带, 与预期长度相符, 测序结果显示该条带5′端266 bp与pCAMBIA1305.1载体骨架上GUS基因序列100%一致, 3′端276 bp与甘草LIS序列(MZ169552) 100%一致, 表明毛状根中的LIS基因序列为外源导入, 而非毛状根自带。以上结果证明, 已诱导获得过表达LIS基因的甘草毛状根系。
3 甘草毛状根中LIS基因的表达水平分析
通过RT-qPCR测定甘草毛状根中LIS基因的表达水平, 使用GraphPad Prism 8软件对其进行样品间单因素方差分析, 结果如图 5所示: 5个LIS+根系中LIS基因的相对表达量均显著高于WT。
4 甘草毛状根中甘草酸的含量检测
各甘草毛状根系经液体培养21天后的生长情况如图 6a所示, 各毛状根系均长势良好。毛状根系中甘草酸含量测定的UPLC色谱图如图 6b所示, 其中1为甘草酸对照品色谱图, 显示甘草酸保留时间为6.532 min, 2~4依次为WT、NC和LIS+甘草毛状根系的色谱图。甘草酸标准曲线回归方程为: Y = 2 683 455.37X - 1 660.17 (R2 = 0.999 9), 线性范围为: 0.004 3~0.085 8 μg·mL-1。图 6c为各甘草毛状根系中甘草酸含量的柱状图, 运用GraphPad Prism 8软件对甘草酸含量进行样品间单因素方差分析, 结果显示所有LIS+样品中甘草酸含量均显著低于WT和NC样品。
讨论
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本研究克隆了甘草LIS基因, 构建了过表达载体pCA-LIS, 通过发根农杆菌介导法诱导甘草胚轴外植体形成了野生型、阴性对照及过表达LIS基因的甘草毛状根系, 通过RT-qPCR检测LIS基因的表达水平, 并采用UPLC法测定各甘草毛状根系中的甘草酸含量。结果表明, 过表达LIS基因的甘草毛状根系中, LIS基因的表达量显著高于野生型, 而甘草酸含量则显著低于野生型和阴性对照组, 表明LIS基因的表达水平与甘草酸含量呈负相关。
如前所述, 高等植物的萜类合成涉及MVA和MEP途径, 前者以乙酰辅酶A作为起始物, 主要合成倍半萜、三萜及其他多萜; 后者则以丙酮酸和3-磷酸甘油醛为起始物, 主要合成单萜、二萜及类胡萝卜素等萜类化合物[28]。乙酰辅酶A的生物合成除了泛酸这一通路之外, 作为有氧呼吸的一环, 也可从葡萄糖开始, 经由丙酮酸氧化而生成。LIS作为单萜途径上的关键酶, 其基因过表达上调了MEP途径, 促进了丙酮酸和3-磷酸甘油醛的消耗, 降低了乙酰辅酶A的合成, 使得进入MVA途径的前体化合物减少, 从而导致甘草酸的含量下降。此外, 植物中各种萜类化合物是由IPP和DMAPP按照不同比例缩合而成的[29], 如: 位于胞质中的MVA途径合成的IPP和DMAPP通常按照2∶1的比例合成倍半萜及三萜的前体化合物——法尼基焦磷酸(farnesyl pyrophosphate, FPP); 而位于质体中的MEP途径则通常按照IPP∶DMAPP为6∶1的比例来合成单萜途径前体GPP和四萜途径前体牻牛儿牻牛儿基焦磷酸(geranylgeranyl pyrophosphate, GGPP)[30]。虽然MVA和MEP途径在亚细胞区域中是彼此分开的, 但有研究表明植物中这两条途径之间存在相互关联, 胞质中的IPP能够进行跨质体膜的运输[31]。因此, 当LIS过表达时, 其前体GPP的过量消耗将导致MEP途径上IPP的需求增多, 促使MVA途径合成的IPP流向MEP途径, 造成三萜途径前体FPP的合成量减少, 这也是导致甘草酸含量下降的可能原因。
近些年来, 大量研究均表明功能基因能够影响药用植物有效成分的生物合成, 例如: 在丹参(Salvia miltiorrhiza) 毛状根中过表达SmHMGR1、SmHMGR2和SmHMGR3可显著提高丹参酮的含量[32]; 在青蒿(Artemisia annua) 中过表达AaTAR2可正向调节青蒿素和类黄酮的生物合成[33]。同样地, 在本课题组前期研究中也发现, 过表达CHS和CHI可有效提高甘草毛状根中总黄酮的水平[24, 34]; 而过表达β-AS和UGE则可使其甘草酸含量显著升高[23]。今后, 将继续以甘草毛状根为实验材料, 对影响甘草酸生物合成的其他关键基因开展研究, 完善甘草酸生物合成的分子调控网络, 为甘草的大规模优培优选奠定基础。
作者贡献: 柯铃钰撰写了论文; 刘颖和何平构思并设计了实验方案; 柯铃钰、陈姿伊和丁文文合作开展了相关实验, 对本论文同等贡献; 张智新协助分析了实验数据; 所有作者均阅读并参与修改了这篇文章。
利益冲突: 本文作者均没有利益冲突。
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2022年第57卷第12期
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doi: 10.16438/j.0513-4870.2022-0666
  • 接收时间：2022-06-03
  • 首发时间：2025-12-24
  • 出版时间：2022-12-12
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  • 收稿日期：2022-06-03
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    北京中医药大学生命科学学院, 北京 102488

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*何平, Tel: 86-10-53912163, E-mail: ;
刘颖, E-mail:
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https://castjournals.cast.org.cn/joweb/yxxb/CN/10.16438/j.0513-4870.2022-0666
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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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