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Article(id=1308842146724143799, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1308841897058198067, articleNumber=null, orderNo=null, doi=10.16438/j.0513-4870.2026-0297, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1773676800000, receivedDateStr=2026-03-17, revisedDate=1777392000000, revisedDateStr=2026-04-29, acceptedDate=null, acceptedDateStr=null, onlineDate=1789981884890, onlineDateStr=2026-09-21, pubDate=1783785600000, pubDateStr=2026-07-12, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1789981884890, onlineIssueDateStr=2026-09-21, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1789981884890, creator=13701087609, updateTime=1789981884890, updator=13701087609, issue=Issue{id=1308841897058198067, tenantId=1146029695717560320, journalId=1189982191388893191, year='2026', volume='61', issue='7', pageStart='2377', pageEnd='2820', issueExtLink='null', onlineDate='null', pubDate='1783785600000', pubDateStr='2026-07-12', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1789981825365, creator='13701087609', updateTime=1789984154811, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1308851667525792021, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1308841897058198067, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1308851667525792022, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1308841897058198067, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=2801, endPage=2809, ext={EN=ArticleExt(id=1308842146963219128, articleId=1308842146724143799, tenantId=1146029695717560320, journalId=1189982191388893191, language=EN, title=Cloning and functional characterization of sesquiterpene synthase genes from Centipeda minima, columnId=1190335348761793317, journalTitle=Acta Pharmaceutica Sinica, columnName=Original Articles, runingTitle=null, highlight=null, articleAbstract=

Centipeda minima (L.) A. Braun et Aschers. is a traditional medicinal plant. Sesquiterpenoids are its core bioactive components, and sesquiterpene synthases (STSs) are the key enzymes in sesquiterpene biosynthesis. In this study, using C. minima as material, nine full-length candidate CmTPSs genes were screened from published transcriptome data by local Blast alignment combined with phylogenetic analysis. Bioinformatics approaches were used to analyze their sequence features, phylogenetic relationships, and conserved motifs. TBtools was employed to construct gene expression heatmaps for investigating their tissue-specific expression patterns. The catalytic functions of CmTPS1, CmTPS3, CmTPS4, and CmTPS5 were further verified by gene cloning, heterologous expression in Saccharomyces cerevisiae, and GC-MS analysis. Results showed that the nine CmTPSs genes fell into five TPS subfamilies, namely TPS-a, TPS-b, TPS-c, TPS-e/f, and TPS-g, and displayed distinct tissue-specific expression in roots, stems, leaves, and flowers. CmTPS1 and CmTPS4 were specific caryophyllene synthases, while CmTPS3 and CmTPS5 were multifunctional synthases. This study provides key gene resources for elucidating the molecular mechanism of sesquiterpene biosynthesis in C. minima, and offers theoretical and technical support for the efficient production of its bioactive constituents and the development of its medicinal resources.

, authors=Hui-ru CHEN1, 2, Shun LIANG1, 2, Zhuo-yi ZHANG1, 2, Li-xin DUAN1, 2, Ai-jia JI1, 2, *, authorsList=Hui-ru CHEN, Shun LIANG, Zhuo-yi ZHANG, Li-xin DUAN, Ai-jia JI, authorCompany=null, correspAuthors=Ai-jia JI, authorNote=null, correspAuthorsNote=null, copyrightStatement=Copyright ©2026 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, fund=null), CN=ArticleExt(id=1308842148297007815, articleId=1308842146724143799, tenantId=1146029695717560320, journalId=1189982191388893191, language=CN, title=鹅不食草倍半萜合酶基因的克隆及功能表征, columnId=1190335348896011050, journalTitle=药学学报, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=

鹅不食草[Centipeda minima (L.) A. Braun et Aschers.] 为传统药用植物, 倍半萜类化合物是其核心活性成分, 而倍半萜合酶(STSs) 是倍半萜生物合成的关键酶。本研究以鹅不食草为材料, 基于已发表的转录组数据, 通过本地Blast比对结合系统发育树分析, 筛选获得9条完整的CmTPSs候选基因。采用生物信息学方法分析其序列特征、系统发育归属及保守基序, 利用TBtools软件绘制基因表达热图分析其组织特异性表达模式, 并通过基因克隆、酿酒酵母异源表达及GC-MS检测, 验证了CmTPS1、CmTPS3、CmTPS4、CmTPS5的催化功能。结果表明, 9条CmTPSs基因分属TPS-a、TPS-b、TPS-c、TPS-e/f、TPS-g 5个亚家族, 且在根、茎、叶、花中呈现明显的组织特异性表达; CmTPS1和CmTPS4为专一性Caryophyllene合酶, CmTPS3和CmTPS5为多功能合酶。本研究为解析鹅不食草倍半萜生物合成分子机制提供了关键基因资源, 也为其活性成分的高效制备及药用资源开发提供了理论与技术支撑。

, authors=陈慧茹1, 2, 梁顺1, 2, 章卓易1, 2, 段礼新1, 2, 季爱加1, 2, *, authorsList=陈慧茹, 梁顺, 章卓易, 段礼新, 季爱加, authorCompany=null, correspAuthors=季爱加, authorNote=null, correspAuthorsNote=
季爱加, E-mail:
, copyrightStatement=版权所有©《药学学报》编辑部2026, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=lMi/0/qGwmvzxhGi1RK5Hw==, magXml=g4DEDmdpaSN4iwRYFYx0UQ==, pdfUrl=null, pdf=8KNNMp8/bBOlD/ioZrE0mw==, pdfFileSize=6397213, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=dP2QCwKK8hs0OI5kM+fm+g==, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=5Cq+X0U2vLkd3E5CpyRXnQ==, mapNumber=null, fund=null)}, authors=[Author(id=1308842148687078095, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842146724143799, 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=1308842148775158482, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842146724143799, authorId=1308842148687078095, language=EN, stringName=Hui-ru CHEN, firstName=Hui-ru, middleName=null, lastName=CHEN, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=1, 2, address=1School of Pharmaceutical Sciences, Guangzhou University of Chinese Medicine, International Institute for Translational Chinese Medicine, Guangdong Engineering Research Center of Biosynthesis and Metabolism of Effective Components of Chinese Medicine, Guangzhou 510006, China
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A: Chromatograms of CmTPS3 products, β-elemene standard and neointermedeol standard; B: Mass spectrograms of CmTPS3 products, β-elemene standard and neointermedeol standard, figureFileSmall=Jp/m5tAQIk09mTGXxDV7CQ==, figureFileBig=Gf/Ouiqz4Rcd3/n6k5wedA==, tableContent=null), ArticleFig(id=1308842151170106112, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842146724143799, language=EN, label=null, caption=null, figureFileSmall=23FoZsiModWxPGDA0CXlMA==, figureFileBig=tEr6C8Y7ux5b+fOsUa8geQ==, tableContent=null), ArticleFig(id=1308842151237214977, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842146724143799, language=CN, label=Fig.7, caption= GC-MS detection of yeast-expressed products of CmTPS5 and positive gene AlTPS20. A: Chromatograms of CmTPS5 products and positive gene AlTPS20 products; B: Mass spectrograms of CmTPS5 products and positive gene AlTPS20 products, figureFileSmall=23FoZsiModWxPGDA0CXlMA==, figureFileBig=tEr6C8Y7ux5b+fOsUa8geQ==, tableContent=null), ArticleFig(id=1308842151295935234, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842146724143799, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
Strains and plasmidsDescriptionSource
pESC-URAYeast expression vector with URA selection markerStored in the lab
pESC-URA-CmTPS1pESC-URA-GAL1p-CmTPS1-CYC1tThis study
pESC-URA-CmTPS3pESC-URA-GAL1p-CmTPS3-CYC1tThis study
pESC-URA-CmTPS4pESC-URA-GAL1p-CmTPS4-CYC1tThis study
pESC-URA-CmTPS5pESC-URA-GAL1p-CmTPS5-CYC1tThis study
yYF601CEN.PK2-1C; erg9:: KanMX-CTR3p ERG9; leu2-3, 112:: His3MX6-GAL1p ERG19/GAL10p-
ERG8; ura3 52:: GAL1p-EfMvaSA110G/GAL10p EfMvaE; his3Δ1:: hphMX4-GAL1p ERG12/
GAL10p-IDI1; 308a: : GAL1p ERG20F96W-N127W		Presented by Xiaozhou
Luo team
ZY0yYF601; GAL80△: : Stored in the lab
ZY1ZY0; pESC-URA-CmTPS1This study
ZY2ZY0; pESC-URA-CmTPS3This study
ZY3ZY0; pESC-URA-CmTPS4This study
ZY4ZY0; pESC-URA-CmTPS5This study
), ArticleFig(id=1308842151354655491, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842146724143799, language=CN, label=Tab.1, caption=

Strains and plasmids used in this study

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Strains and plasmidsDescriptionSource
pESC-URAYeast expression vector with URA selection markerStored in the lab
pESC-URA-CmTPS1pESC-URA-GAL1p-CmTPS1-CYC1tThis study
pESC-URA-CmTPS3pESC-URA-GAL1p-CmTPS3-CYC1tThis study
pESC-URA-CmTPS4pESC-URA-GAL1p-CmTPS4-CYC1tThis study
pESC-URA-CmTPS5pESC-URA-GAL1p-CmTPS5-CYC1tThis study
yYF601CEN.PK2-1C; erg9:: KanMX-CTR3p ERG9; leu2-3, 112:: His3MX6-GAL1p ERG19/GAL10p-
ERG8; ura3 52:: GAL1p-EfMvaSA110G/GAL10p EfMvaE; his3Δ1:: hphMX4-GAL1p ERG12/
GAL10p-IDI1; 308a: : GAL1p ERG20F96W-N127W		Presented by Xiaozhou
Luo team
ZY0yYF601; GAL80△: : Stored in the lab
ZY1ZY0; pESC-URA-CmTPS1This study
ZY2ZY0; pESC-URA-CmTPS3This study
ZY3ZY0; pESC-URA-CmTPS4This study
ZY4ZY0; pESC-URA-CmTPS5This study
), ArticleFig(id=1308842151425958660, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842146724143799, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
PrimerSequence (5′→3′)
pESC-URA-CmTPS1-FaggagaaaaaaccccggatccATGGCGTCAACATGGACACTT
pESC-URA-CmTPS1-RgctagccgcggtaccaagcttTTATGACAGCCCTTGGATTGG
pESC-URA-CmTPS3-FaggagaaaaaaccccggatccATGGCAGCAGTTACACCCAATG
pESC-URA-CmTPS3-RgctagccgcggtaccaagcttTTACATGGGTAGAGAACCAACAAACA
pESC-URA-CmTPS4-FaggagaaaaaaccccggatccATGCACGCGATAGTAATTCTGTTAG
pESC-URA-CmTPS4-RgctagccgcggtaccaagcttTCATATGCTTATAGCATGCACTAACAA
pESC-URA-CmTPS5-FaggagaaaaaaccccggatccATGTCTGCTGAGCAAGATGAAGTT
pESC-URA-CmTPS5-RgctagccgcggtaccaagcttTCATATGCTGAGAGCATCAACAAGT
), ArticleFig(id=1308842151488873221, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842146724143799, language=CN, label=Tab.2, caption=

Primer sequences for constructing yeast expression vectors for C. minima genes

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PrimerSequence (5′→3′)
pESC-URA-CmTPS1-FaggagaaaaaaccccggatccATGGCGTCAACATGGACACTT
pESC-URA-CmTPS1-RgctagccgcggtaccaagcttTTATGACAGCCCTTGGATTGG
pESC-URA-CmTPS3-FaggagaaaaaaccccggatccATGGCAGCAGTTACACCCAATG
pESC-URA-CmTPS3-RgctagccgcggtaccaagcttTTACATGGGTAGAGAACCAACAAACA
pESC-URA-CmTPS4-FaggagaaaaaaccccggatccATGCACGCGATAGTAATTCTGTTAG
pESC-URA-CmTPS4-RgctagccgcggtaccaagcttTCATATGCTTATAGCATGCACTAACAA
pESC-URA-CmTPS5-FaggagaaaaaaccccggatccATGTCTGCTGAGCAAGATGAAGTT
pESC-URA-CmTPS5-RgctagccgcggtaccaagcttTCATATGCTGAGAGCATCAACAAGT
), ArticleFig(id=1308842151551787782, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842146724143799, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
Candidate gene IDGeneProtein length (aa)
DN841_c0_g4_i2CmTPS1589
DN31917_c0_g1_i3CmTPS2602
DN6136_c0_g1_i10CmTPS3559
DN86_c0_g1_i4CmTPS4564
DN6237_c0_g1_i3CmTPS5547
DN15510_c0_g1_i1CmTPS6586
DN3015_c0_g1_i24CmTPS7787
DN7857_c0_g1_i1CmTPS8867
DN7592_c0_g1_i1CmTPS9799
), ArticleFig(id=1308842151623090951, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842146724143799, language=CN, label=Tab.3, caption=

Nine candidate genes for CmTPSs

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Candidate gene IDGeneProtein length (aa)
DN841_c0_g4_i2CmTPS1589
DN31917_c0_g1_i3CmTPS2602
DN6136_c0_g1_i10CmTPS3559
DN86_c0_g1_i4CmTPS4564
DN6237_c0_g1_i3CmTPS5547
DN15510_c0_g1_i1CmTPS6586
DN3015_c0_g1_i24CmTPS7787
DN7857_c0_g1_i1CmTPS8867
DN7592_c0_g1_i1CmTPS9799
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鹅不食草倍半萜合酶基因的克隆及功能表征
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陈慧茹 1, 2 , 梁顺 1, 2 , 章卓易 1, 2 , 段礼新 1, 2 , 季爱加 1, 2, *
药学学报 | 研究论文 2026,61(7): 2801-2809
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药学学报 |研究论文 2026 , 61 (7) : 2801 -2809
鹅不食草倍半萜合酶基因的克隆及功能表征
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陈慧茹1, 2, 梁顺1, 2, 章卓易1, 2, 段礼新1, 2, 季爱加1, 2, *
作者信息
  • 1广州中医药大学中药学院, 国际中医药转化医学研究所, 中药功效成分生物合成与代谢广东工程研究中心, 广东 广州 510006
  • 2中医药广东省实验室 (横琴实验室), 广东 珠海 519000
通讯作者:
季爱加, E-mail:
Cloning and functional characterization of sesquiterpene synthase genes from Centipeda minima
Hui-ru CHEN1, 2, Shun LIANG1, 2, Zhuo-yi ZHANG1, 2, Li-xin DUAN1, 2, Ai-jia JI1, 2, *
Affiliations
  • 1School of Pharmaceutical Sciences, Guangzhou University of Chinese Medicine, International Institute for Translational Chinese Medicine, Guangdong Engineering Research Center of Biosynthesis and Metabolism of Effective Components of Chinese Medicine, Guangzhou 510006, China
  • 2Guangdong Provincial Laboratory of Traditional Chinese Medicine (Hengqin Laboratory), Zhuhai 519000, China
出版时间: 2026-07-12 doi: 10.16438/j.0513-4870.2026-0297
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摘要
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鹅不食草[Centipeda minima (L.) A. Braun et Aschers.] 为传统药用植物, 倍半萜类化合物是其核心活性成分, 而倍半萜合酶(STSs) 是倍半萜生物合成的关键酶。本研究以鹅不食草为材料, 基于已发表的转录组数据, 通过本地Blast比对结合系统发育树分析, 筛选获得9条完整的CmTPSs候选基因。采用生物信息学方法分析其序列特征、系统发育归属及保守基序, 利用TBtools软件绘制基因表达热图分析其组织特异性表达模式, 并通过基因克隆、酿酒酵母异源表达及GC-MS检测, 验证了CmTPS1、CmTPS3、CmTPS4、CmTPS5的催化功能。结果表明, 9条CmTPSs基因分属TPS-a、TPS-b、TPS-c、TPS-e/f、TPS-g 5个亚家族, 且在根、茎、叶、花中呈现明显的组织特异性表达; CmTPS1和CmTPS4为专一性Caryophyllene合酶, CmTPS3和CmTPS5为多功能合酶。本研究为解析鹅不食草倍半萜生物合成分子机制提供了关键基因资源, 也为其活性成分的高效制备及药用资源开发提供了理论与技术支撑。

鹅不食草  /  倍半萜  /  生物合成  /  功能表征

Centipeda minima (L.) A. Braun et Aschers. is a traditional medicinal plant. Sesquiterpenoids are its core bioactive components, and sesquiterpene synthases (STSs) are the key enzymes in sesquiterpene biosynthesis. In this study, using C. minima as material, nine full-length candidate CmTPSs genes were screened from published transcriptome data by local Blast alignment combined with phylogenetic analysis. Bioinformatics approaches were used to analyze their sequence features, phylogenetic relationships, and conserved motifs. TBtools was employed to construct gene expression heatmaps for investigating their tissue-specific expression patterns. The catalytic functions of CmTPS1, CmTPS3, CmTPS4, and CmTPS5 were further verified by gene cloning, heterologous expression in Saccharomyces cerevisiae, and GC-MS analysis. Results showed that the nine CmTPSs genes fell into five TPS subfamilies, namely TPS-a, TPS-b, TPS-c, TPS-e/f, and TPS-g, and displayed distinct tissue-specific expression in roots, stems, leaves, and flowers. CmTPS1 and CmTPS4 were specific caryophyllene synthases, while CmTPS3 and CmTPS5 were multifunctional synthases. This study provides key gene resources for elucidating the molecular mechanism of sesquiterpene biosynthesis in C. minima, and offers theoretical and technical support for the efficient production of its bioactive constituents and the development of its medicinal resources.

Centipeda minima  /  sesquiterpene  /  biosynthesis  /  functional verification
陈慧茹, 梁顺, 章卓易, 段礼新, 季爱加. 鹅不食草倍半萜合酶基因的克隆及功能表征. 药学学报, 2026 , 61 (7) : 2801 -2809 . DOI: 10.16438/j.0513-4870.2026-0297
Hui-ru CHEN, Shun LIANG, Zhuo-yi ZHANG, Li-xin DUAN, Ai-jia JI. Cloning and functional characterization of sesquiterpene synthase genes from Centipeda minima[J]. Acta Pharmaceutica Sinica, 2026 , 61 (7) : 2801 -2809 . DOI: 10.16438/j.0513-4870.2026-0297
正文
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鹅不食草[Centipeda minima (L.) A. Braun et Aschers.] 为菊科石胡荽属常用药用植物[1], 别名石胡荽、地芫荽、鸡肠草等, 在我国传统医药体系中应用悠久。历代本草记载其味辛性温, 具通鼻窍、散风寒、解毒消肿之效, 是治疗鼻渊、咳喘及炎症相关病症的经典药材[2]。现代药理学研究进一步证实, 鹅不食草富含挥发油、黄酮、萜类等多种活性成分, 进而展现出抗肿瘤、抗炎、抗氧化、抗菌、抗过敏、保肝、抗病毒等多元药理活性[3]。
萜类化合物是鹅不食草中种类最丰富的化合物, 目前已鉴定出117种, 涵盖单萜类、二萜类、倍半萜类和三萜类, 其中倍半萜类被认为是该植物的特征性成分及主要生物活性物质[4]。例如短叶老鹳草素(brevilin A, BA)、山金车内酯C (arnicolide C, ArnC)、山金车内酯D (arnicolide D, ArnD)、堆心菊灵等倍半萜类化合物, 因其显著的药理活性而被广泛研究并为人熟知; 值得注意的是, 鹅不食草中的倍半萜内酯以愈创木内酯类和伪愈创木内酯类为主, 是其核心活性组分[5]。
植物体内萜类化合物的生物合成主要通过两条空间上相互独立的代谢途径完成: 一条是定位于细胞质的甲羟戊酸途径(mevalonate pathway, MVA), 另一条是存在于质体中的2‑C‑甲基‑D‑赤藓糖醇‑4‑磷酸途径(2‑C‑methyl‑D‑erythritol 4‑phosphate pathway, MEP)。两条途径可通过IPP的跨质膜运输实现代谢交流, 共同参与植物萜类物质的合成调控[6]。倍半萜类化合物(C15) 结构多样且生物活性广泛, 其合成与积累主要依赖细胞质中的MVA途径。该途径以乙酰辅酶A为起始底物, 经多步酶促反应生成异戊烯基二磷酸(IPP) 及其异构体二甲基烯丙基二磷酸(DMAPP), 二者进一步缩合形成倍半萜的直接前体——法尼基焦磷酸(FPP)。倍半萜合酶(sesquiterpene synthases, STSs) 可催化FPP发生环化、重排等反应, 形成线性、单环、双环及三环等多种萜类核心骨架, 再经氧化还原、糖基化等后续修饰, 最终生成结构独特、功能多样的倍半萜类化合物[7]。
转录组测序技术可在全转录组水平高效识别萜类合成相关关键酶基因, 突破传统基因克隆的局限, 为非模式药用植物功能基因挖掘提供支撑[8]。合成生物学的发展为天然产物可持续供给提供新路径, 酿酒酵母(Saccharomyces cerevisiae) 因遗传背景清晰、操作简便、发酵成熟, 成为植物源倍半萜异源合成的理想底盘细胞[9, 10]。
因此, 本研究以鹅不食草为材料, 采用转录组测序筛选挖掘倍半萜合酶基因, 通过生物信息学分析明确其结构与功能, 构建酿酒酵母异源表达体系并验证基因功能。研究成果可阐明鹅不食草倍半萜生物合成途径, 丰富药用植物萜类合成分子基础, 为活性成分高效制备及药用资源绿色可持续利用提供理论与技术支撑, 具重要科学价值与应用前景。
材料与方法
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植物材料   本研究所用鹅不食草植物材料采自广东省梅州市, 经广州中医药大学中药学院药用植物学教研室彭光天博士鉴定确认。将采摘的新鲜植株用蒸馏水清洗干净后, 分别按根、茎、叶、花四个部位进行分离收集并做好对应标记, 随后置于液氮中速冻, 冷冻完成后转移至-80 ℃冰箱中保存, 以备后续实验使用。
菌株与质粒   本研究所用的菌株为在高产FPP的酿酒酵母工程菌yYF601 (中国科学院深圳先进技术研究院罗小舟研究员团队提供) 的基础上, 课题组对基因GAL80进行敲除所构建的菌株ZY0, 其中质粒载体和本研究所构建的菌株见表1。
生物信息学分析   采用MEGA X软件, 将CmTPSs的蛋白质序列与从文献及NCBI数据库中收集的已完成功能注释的TPS基因对应的蛋白质序列进行比对分析。基于比对结果, 采用邻接法(设置1 000次自举重复) 进行系统发育树构建与分析, 生成的系统发育树通过ChiPlot网站(ChiPlot) 进行美化处理, 以获得清晰规范的可视化结果; 基因表达量分析通过TBtools软件绘制热图进行可视化呈现; 使用GENEDOC软件对候选CmTPSs蛋白进行多重序列比对, 分析其保守基序。
鹅不食草RNA提取及cDNA获取    采用北京华越洋生物有限公司生产的植物总RNA提取试剂盒, 严格按照试剂盒说明书操作, 分别提取鹅不食草根、茎、叶、花四个组织部位的总RNA。提取完成后, 采用1%琼脂糖凝胶电泳检测总RNA的完整性, 通过NanoDrop One超微量分光光度计检测总RNA的浓度, 然后使用艾科瑞生物的Evo M-MLV Plus cDNA合成试剂盒构建cDNA文库, 构建完成后保存于-20 ℃冰箱, 作为后续基因克隆实验的模板。
鹅不食草倍半萜合酶基因克隆    基于鹅不食草转录组数据库中基因的注释信息, 筛选得到CDS全长序列, 使用Snapgene软件设计5′和3′端特异性引物, 见表2。以鹅不食草总RNA反转录得到的cDNA文库作为模板进行扩增反应。扩增反应体系: 2× Phanta Max Buffer 50 μL、dNTP Mix 2 μL、Phanta Max Super-Fidelity DNA Polymerase 2 μL、cDNA 1 μL、正反向引物各2 μL、无菌水补足至100 μL; 反应程序: 95 ℃ 3 min; 95 ℃ 15 s, 57 ℃ 15 s, 72 ℃ 1 min, 34个循环; 72 ℃ 5 min; 4 ℃保存。
PCR产物经1%琼脂糖凝胶电泳检测后, 选取条带位置及大小符合预期的目的条带, 采用胶回收试剂盒进行纯化回收。采用限制性核酸内切酶BamHⅠ/HindⅢ对pESC-URA载体进行双酶切处理, 将纯化后的PCR产物与线性化的pESC-URA载体通过同源重组方法连接, 连接产物转入大肠杆菌DH5α感受态细胞中。后续挑取单菌落进行菌液PCR鉴定, 筛选获得阳性菌液并送测; 提取测序结果正确的阳性菌液质粒, 获得pESC-URA-CmTPSs重组表达载体, 用于后续转化实验。
酵母感受态制备与转化    取保存的ZY0甘油菌, 在固体YPD培养基上划线进行活化处理; 挑取生长状态良好的单菌落, 接种至2 mL YPD液体培养基中, 置于摇床中过夜培养。次日清晨, 吸取适量复苏后的菌液转接至YPD培养基中, 继续置于摇床中培养, 直至菌液OD600值达到0.8~1.2, 即可开展后续转化操作。本实验采用醋酸锂化学转化法进行重组载体的转化[11]。
酵母转化子验证及诱导表达    当转化平板上长出单克隆菌落后, 为确认目的基因是否成功导入酵母底盘菌, 需采用基因特异性引物进行酵母菌落PCR验证。验证合格后, 在扩繁平板上挑取3个阳性转化子, 分别接种至含有3 mL相应筛选标记SD培养基的摇菌管中, 于30 ℃、200 r·min-1条件下过夜培养, 制备种子液。随后按1%的接种量, 将种子液转接至含有50 mL相应筛选标记SD培养基的摇瓶中, 发酵培养约24 h后, 向每瓶菌液中加入5 mL正十二烷[12], 使其覆盖于发酵液表面(以避免代谢产物在诱导表达过程中挥发), 之后置于酵母摇床上持续培养5 d。
代谢产物检测与鉴定    摇瓶培养结束后, 将发酵液连同萃取剂置于50 mL离心管中, 4 200 ×g离心5 min, 吸取上层正十二烷有机相, 用正己烷适当稀释并过滤处理, 稀释液用于后续定性或定量分析。
采用气相色谱-质谱联用仪[岛津GC-MS (QP2010SE)] 检测代谢产物。分析条件如下: 色谱柱型号为DB5MS (30 m × 0.25 mm, 0.25 μm), 载气为氦气, 流速为1.0 mL·min-1, 进样口温度为250 ℃, 不分流。升温程序: 初始温度为40 ℃保持3 min, 然后以10 ℃·min-1升温至130 ℃, 接着以3 ℃·min-1升温至160 ℃, 最后以40 ℃·min-1升温至250 ℃。此外, 电子能量为70 eV, 离子源温度为250 ℃, 接口温度为170 ℃, 扫描区间为m/z 50~600。根据NIST17质谱库搜索质谱成分, 质谱匹配因子(mass spectra match factor) > 85%的成分可初步鉴定为该化合物[13]。
结果
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1　 CmTPSs基因挖掘
本研究中鹅不食草的转录组数据来源于课题组前期一项已发表的研究[14], 为鉴定参与倍半萜生物合成途径的候选基因, 实验采用收集不同物种来源的已知功能TPSs基因的蛋白序列作为诱饵, 与鹅不食草的转录组数据进行本地Blast比对, 结合MEGA软件构建系统进化树, 初步筛选获得CmTPSs候选基因。通过本地Blast初步筛选得到9条序列, 在NCBI的ORF finder工具中查询每条序列最长的开放阅读框(open reading frame, ORF)。根据文献报道, 萜烯合酶基因的核苷酸序列长度通常为1 050~2 600 bp, 对应的氨基酸数量主要为350~860个, 据此判断, 初步筛选出的9条TPSs基因均可能为完整的TPSs基因, 其具体基因情况详见表3。
2　 CmTPSs基因表达量分析
在候选基因筛选过程中, 基因表达量水平[15]是重点关注指标之一。FPKM值是通过转录组测序获得的基因表达量量化指标, 可根据其差异判断对应基因的表达水平高低。图1展示了9条CmTPSs候选基因在鹅不食草根、茎、叶、花四个组织部位的表达水平差异, 结果显示: CmTPS1、CmTPS6在花组织中特异性表达; CmTPS2、CmTPS3在鹅不食草全株中均呈高表达; CmTPS4、CmTPS5、CmTPS9在根、茎组织中特异性高表达; CmTPS7在花、根、茎组织中均有表达; CmTPS8在叶、茎组织中表达。通过对候选基因差异表达量数据的分析, 可为后续基因扩增实验中选择对应组织部位的cDNA作为扩增模板提供重要参考依据。
3　系统进化树分析
利用MEGA软件构建系统发育树(图2), 结果显示: CmTPS1归属于TPS-b亚家族, 该亚家族虽为单萜合酶和异戊二烯合酶的特征性亚家族, 催化产物以环状单萜为主, 但已有研究表明部分TPS-b亚家族成员可兼具备倍半萜合酶活性[16-18]; 结合CmTPS1在鹅不食草花器官中呈特异性表达的特征, 推测其可能参与倍半萜合成。CmTPS2、CmTPS6归类于TPS-g亚家族, CmTPS3、CmTPS4、CmTPS5归属于TPS-a亚家族, CmTPS7、CmTPS8归类于TPS-e/f亚家族, CmTPS9归属于TPS-c亚家族。其中, TPS‑a是在双子叶植物与单子叶植物中均有分布且编码倍半萜合酶基因的最大亚家族; TPS‑b为单萜合酶和异戊二烯合酶特征性亚家族, 其催化产物以环状单萜为主。鉴于本研究重点关注鹅不食草的倍半萜合酶基因, 后续将选取CmTPS1、CmTPS3、CmTPS4、CmTPS5进行基因功能验证。
4　 CmTPSs多重序列比对
对候选CmTPSs蛋白进行多重序列比对(图3), 所有TPS全长序列均含有两个保守结构域, 分别位于序列的N端和C端。其中, N端为精氨酸保守结构域, 该结构域包含保守基序“R(R)X8W”, 此基序不仅与单萜的环化及异构化过程密切相关, 还对蛋白产物的结构稳定性具有重要作用; 在“R(R)X8W”基序上游, 还存在一段转运肽, 其功能是引导萜类合酶参与质体中的物质合成过程。C端为两个高度保守的天冬氨酸保守结构域, 包含“DDXXD”和“NSE/DTE”两个特征基序。其中, “DDXXD”基序的特定位点可促进二价离子与水分子的特异性结合并维持其稳定性; 这两个基序均能为Mg2+和Mn2+提供结合位点, 进而为萜类合酶的正常代谢活动提供保障[19]。研究发现, CmTPS1、CmTPS3、CmTPS4、CmTPS5这四个候选基因的保守基序均保持完整, 但部分氨基酸位点存在差异, 这暗示它们的功能可能存在分化, 其具体功能差异仍需通过后续实验进一步验证。
5　鹅不食草RNA提取、倍半萜合酶基因克隆和表达载体构建
分别提取鹅不食草根、茎、叶、花四个不同组织部位的总RNA (图4A), 采用1%琼脂糖凝胶电泳对其完整性进行检测。电泳结果显示, 在28S和18S rRNA对应位置出现两条清晰条带, 表明所提取的RNA样品完整性良好、质量合格, 可用于后续实验。针对4条CmTPSs候选基因, 设计特异性基因扩增引物; 以反转录获得的鹅不食草cDNA文库为模板, 进行PCR扩增反应。扩增产物经1%琼脂糖凝胶电泳检测, 结果显示在约1 000~2 000 bp内出现明亮条带, 且条带大小与预期目的基因片段一致, 表明成功扩增出4条CmTPSs基因(图4B)。将PCR扩增产物与载体连接后进行测序验证, 测序结果显示, 克隆获得的基因与目的基因的氨基酸序列完全一致; 随后采用质粒小提试剂盒提取阳性克隆质粒(图4C), 备用。
6　 CmTPSs功能验证
6.1　 CmTPS1、4的功能表征
对CmTPS1和CmTPS4的酵母表达产物进行GC-MS检测分析, 结果显示(图5), 与底盘菌株对照组相比, 上述两个倍半萜合酶的表达产物均出现一个新的特征产物峰, 该产物峰与caryophyllene标准品的保留时间及质谱图完全一致。据此可确定, 这两个倍半萜合酶均为caryophyllene合酶。
6.2　 CmTPS3的功能表征
对CmTPS3的酵母表达产物进行GC-MS检测分析, 结果显示(图6), 与底盘菌株对照组相比, 有5个新的产物峰出现, P1和P5分别与β-elemene和neointermedeol标准品具有相同的保留时间和质谱图, 剩余3个产物峰(P2、P3、P4) 因未能纯化分离, 且无对应标准品可购买, 暂无法确定其具体结构。综上所述, 确定CmTPS3是多功能倍半萜合酶。
6.3　 CmTPS5的功能表征
对CmTPS5的酵母表达产物进行GC-MS检测分析, 结果显示(图7), 与底盘菌株对照组相比, 其表达产物中出现5个新的特征产物峰。由于本实验室缺少相应标准品, 无法直接通过保留时间及质谱图比对确定产物结构。为此, 本研究构建了阳性对照基因AlTPS20的酵母表达载体进行比对验证(AlTPS20基因序列由中国中医科学院郭兰萍研究员馈赠, 该基因的酵母表达产物已完成结构鉴定并发表相关研究论文[20])。比对结果显示, 其中3个产物峰分别与(+/-)-β-isocomene、caryophyllene及α-caryophyllene的保留时间和质谱图一致; 剩余2个产物峰(P1、P2) 因未能纯化分离, 且无对应标准品可购买, 暂无法确定其具体结构。综合上述分析可确定, CmTPS5为多功能倍半萜合酶。
讨论
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本研究以传统药用植物鹅不食草为核心研究材料, 整合转录组数据挖掘、基因克隆、生物信息学分析及酿酒酵母异源表达等多项技术, 系统完成了9条CmTPSs候选基因的筛选和4条CmTPSs候选基因的克隆、鉴定及功能验证, 明确了各基因的序列特征、系统发育归属及催化功能。该研究不仅填补了鹅不食草倍半萜合成关键基因研究的空白, 更为鹅不食草倍半萜类成分的生物合成机制阐释、药用活性物质基础解析及绿色规模化制备提供了坚实的理论支撑与可行的技术路径。
通过本地Blast比对结合系统发育树分析, 从鹅不食草转录组数据库中筛选获得9条完整的CmTPSs候选基因, 其编码蛋白长度介于547~867 aa之间, 符合萜烯合酶基因核苷酸序列1 050~2 600 bp、氨基酸序列350~867 aa的典型特征, 证实其为完整的功能基因。系统发育树分析显示, 9条CmTPSs基因分别归属于TPS-a、TPS-b、TPS-c、TPS-e/f、TPS-g 5个亚家族, 其中CmTPS1 (TPS-b亚家族)、CmTPS3、CmTPS4、CmTPS5 (均为TPS-a亚家族) 与倍半萜合酶基因密切相关, 被确定为后续功能验证的目标基因。
基因表达量分析结果表明, 9条CmTPSs基因在鹅不食草根、茎、叶、花四个组织部位呈现明显的组织特异性表达特征: CmTPS1、CmTPS6在花组织中特异性表达, CmTPS2、CmTPS3在全株均呈高表达, CmTPS4、CmTPS5、CmTPS9在根、茎组织中特异性高表达, CmTPS7在花、根、茎组织中均有表达, CmTPS8在叶、茎组织中表达。CmTPSs基因的这种组织特异性表达特征, 为后续针对性选择基因扩增模板、优化异源表达效率提供了坚实的科学依据, 也为深入解析鹅不食草倍半萜类成分的组织特异性积累机制奠定了重要基础。
通过蛋白序列比对发现, 作为后续功能验证核心的CmTPS1、CmTPS3、CmTPS4、CmTPS5均含有完整的“R(R)X8W”“DDXXD”和“NSE/DTE”保守基序。值得注意的是, 这四个基因的保守基序虽均保持完整, 未出现关键位点缺失, 但部分氨基酸位点存在细微差异, 而氨基酸序列的特异性差异往往会影响蛋白的空间构象、催化活性及底物特异性, 这也暗示着CmTPS1、CmTPS3、CmTPS4、CmTPS5的催化功能可能存在分化, 为后续深入探究CmTPSs基因的结构与功能关联、解析其催化机制提供了重要研究线索。
酿酒酵母异源表达及GC-MS检测结果明确了4条CmTPSs基因的催化功能: CmTPS1和CmTPS4均为专一性caryophyllene合酶, 可高效催化底物法尼基焦磷酸(FPP) 生成caryophyllene; caryophyllene作为鹅不食草中的重要活性成分, 具有抗炎、抗肿瘤等药理作用[21], 该结果进一步阐明了鹅不食草药用活性的分子基础。CmTPS3为多功能倍半萜合酶, 可催化生成β-elemene和neointermedeol以及3种未明确结构的倍半萜产物, 其中β-elemene是公认的抗肿瘤活性成分[22, 23], 该发现丰富了鹅不食草活性倍半萜的合成路径。CmTPS5为多功能倍半萜合酶, 可催化合成(+/-)-β-isocomene、caryophyllene、α-caryophyllene及2种未明确结构的倍半萜产物, 其多功能催化特性暗示其在鹅不食草倍半萜类成分多样性形成中发挥核心作用。
本研究从鹅不食草中克隆并在酿酒酵母中功能验证了4条倍半萜合酶基因, 丰富了鹅不食草萜类生物合成相关基因的研究资料, 为进一步解析其倍半萜合成途径与分子机制提供了关键基因资源与实验依据。目前, 现有研究多集中于鹅不食草倍半萜类成分的分离、鉴定及活性评价, 对其生物合成关键基因及调控机制的研究较为薄弱。本研究通过转录组挖掘结合异源表达技术, 明确了CmTPSs基因的功能, 与现有成分分离研究形成互补, 进一步完善了药用植物萜类合成的分子基础。同时, 本研究选用课题组自主构建的高产FPP酿酒酵母工程菌ZY0作为底盘细胞, 成功实现了CmTPSs基因的异源高效表达, 克服了传统植物提取法产量低、周期长、环境影响大、成本高的弊端, 为鹅不食草活性倍半萜成分的规模化、绿色化制备提供了可行路径, 具有重要的工业应用前景。此外, CmTPS5的多功能催化特性及部分基因保守结构域的不完整性, 也为探究植物倍半萜合酶的功能分化及进化规律提供了新的研究素材。
同时, 本研究仍存在一定局限性, 需在后续研究中进一步优化完善: 一是CmTPS3催化生成的3种未知产物(P2、P3、P4) 和CmTPS5催化生成的2种未知产物(P1、P2), 因本实验室缺少对应标准品, 且受分离纯化技术限制, 暂未明确其具体结构, 无法进一步阐释其药用价值; 二是对CmTPSs基因的表达调控机制研究不足, 未明确其在鹅不食草生长发育不同时期、不同环境胁迫条件下的表达规律, 也未深入探究保守结构域缺失对催化功能、底物特异性的具体影响; 三是当前酿酒酵母异源表达体系的目标产物产量仍有较大提升空间, 尚未对基因密码子偏好性、代谢途径分支进行优化, 难以满足工业化生产需求; 四是未探究CmTPSs基因在鹅不食草体内的生理功能, 及其与其他萜类合成相关基因的协同作用机制。

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参考文献 引证文献
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2026年第61卷第7期
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doi: 10.16438/j.0513-4870.2026-0297
  • 接收时间：2026-03-17
  • 首发时间：2026-09-21
  • 出版时间：2026-07-12
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  • 收稿日期：2026-03-17
  • 修回日期：2026-04-29
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    1广州中医药大学中药学院, 国际中医药转化医学研究所, 中药功效成分生物合成与代谢广东工程研究中心, 广东 广州 510006
    2中医药广东省实验室 (横琴实验室), 广东 珠海 519000

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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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