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Article(id=1218551242020798470, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1218551209271677657, articleNumber=null, orderNo=null, doi=10.16438/j.0513-4870.2018-0709, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1533225600000, receivedDateStr=2018-08-03, revisedDate=1537891200000, revisedDateStr=2018-09-26, acceptedDate=null, acceptedDateStr=null, onlineDate=1768454855684, onlineDateStr=2026-01-15, pubDate=1544544000000, pubDateStr=2018-12-12, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1768454855684, onlineIssueDateStr=2026-01-15, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1768454855684, creator=13701087609, updateTime=1768454855684, updator=13701087609, issue=Issue{id=1218551209271677657, tenantId=1146029695717560320, journalId=1189982191388893191, year='2018', volume='53', issue='12', pageStart='1943', pageEnd='2134', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1768454847877, creator=13701087609, updateTime=1768457192749, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1218561044428017831, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1218551209271677657, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1218561044428017832, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1218551209271677657, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=2122, endPage=2128, ext={EN=ArticleExt(id=1218551243044208752, articleId=1218551242020798470, tenantId=1146029695717560320, journalId=1189982191388893191, language=EN, title=Differential pathway and gene expressions of endophytic Phialocephala fortinii inoculated on Rhodiola crenulata, columnId=1218551242851270749, journalTitle=Acta Pharmaceutica Sinica, columnName=ORIGINAL ARTICLES Pharmaceutics, runingTitle=null, highlight=null, articleAbstract=

The development and metabolism of medicinal plant is affected by many factors, among which the effect from endophytic fungi has been noticed recently and has become one of hot fields. In order to explore the effect of endophytic fungi on gene expression in R. crenulata, RNA-sequencing was used to find genes involved in metabolic pathways, and the differential genes were verified by real-time fluorescent quantitative PCR. The method of 2-△△Ct was used to analyze the relative expression levels of genes in related metabolic pathways, which was used to verify the result of transcriptomics sequencing. The results showed that the endophytic fungus, P. fortinii, could up-regulate the gene expression in lipid metabolic pathway of R. crenulata. In signal transduction pathway, the genes were influenced at different level but the gene expressions were significantly increased under control of Notch signaling pathway, which was 34 times of that in control. The gene expressions of environmental adaption pathway were up-regulated in R. crenulata after inoculation of P. fortinii. This study could provide help for further understanding on mechanism of plant-fungus interaction, root cause of geoherbalism of medicinal plant and exploring bio-function of endophytic fungi.

, correspAuthors=Jin-long CUI, authorNote=null, correspAuthorsNote=null, copyrightStatement=Copyright ©2018 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=Meng-liang WANG, Xiao-yin GAO, Jin-long CUI, Jun-hong WANG), CN=ArticleExt(id=1218551245132972371, articleId=1218551242020798470, tenantId=1146029695717560320, journalId=1189982191388893191, language=CN, title=内生菌Phialocephala fortinii影响大花红景天代谢的差异途径及基因表达, columnId=1218551243249729670, journalTitle=药学学报, columnName=研究论文 生药学, runingTitle=null, highlight=null, articleAbstract=

药用植物发育和代谢受到多种因素的影响，而内生真菌对植物的影响是近年来才得以重视并逐步成为热点。为探索内生真菌Phialocephala fortinii对大花红景天基因及代谢途径的影响，本实验通过RNA转录组测序筛选差异表达基因；并用Real-time PCR对基因表达进行验证，利用2-△△Ct的方法分析脂代谢、信号转导和环境适应性等重要代谢途径差异基因的相对表达量。结果表明，验证结果与测序结果一致，内生真菌P.fortinii能够促进大花红景天脂代谢途径中的差异基因表达量；它对信号转导途径中相关基因表达的影响不同，但控制Notch信号通路途径的基因表达量显著提高，为对照组的34倍；对适应性代谢途径差异基因表达上调，进而促进红景天环境适应能力。研究结果旨在为揭示内生真菌与红景天的互作机制、红景天道地性形成原因、内生真菌功能的开发等提供分子基础数据。

, correspAuthors=崔晋龙, authorNote=null, correspAuthorsNote=
* 崔晋龙, Tel:86-351-7016101, E-mail:
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PLoS One, 2016, 11:e0150591., articleTitle=null, refAbstract=null)], funds=[Fund(id=1218970841379951520, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, awardId=31270383, language=CN, fundingSource=国家自然科学基金资助项目(31270383), fundOrder=null, country=null), Fund(id=1218970841442866086, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, awardId=31670328, language=CN, fundingSource=国家自然科学基金资助项目(31670328), fundOrder=null, country=null), Fund(id=1218970841535140778, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, awardId=2014011029-1, language=CN, fundingSource=山西省自然科学基金资助项目(2014011029-1), fundOrder=null, country=null), Fund(id=1218970841610638255, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, awardId=晋教科[2017]14号, language=CN, fundingSource=山西省"1331工程"重点实验室-功能分子化学(晋教科[2017]14号), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1218970836648776295, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, xref=null, ext=[AuthorCompanyExt(id=1218970836652970600, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, companyId=1218970836648776295, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1. Institute of Applied Chemistry, Taiyuan 030006, China), AuthorCompanyExt(id=1218970836661359209, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, companyId=1218970836648776295, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1.山西大学应用化学研究所, 山西 太原 030006)]), AuthorCompany(id=1218970836762022515, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, xref=null, ext=[AuthorCompanyExt(id=1218970836782994036, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, companyId=1218970836762022515, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2. Institute of Biotechnology, Shanxi University, Taiyuan 030006, China), AuthorCompanyExt(id=1218970836795576951, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, companyId=1218970836762022515, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2.山西大学生物技术研究所, 山西 太原 030006)])], figs=[ArticleFig(id=1218970839450571599, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=EN, label=null, caption=null, figureFileSmall=WaV6L2I4jTT/u0y/ZwqQ+w==, figureFileBig=wdUgcU5Aiyl43pYkt/rC0Q==, tableContent=null), ArticleFig(id=1218970839584789335, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=CN, label=Figure 1, caption=

Agarose gel electrophoresis of total RNA extracted from R. crenulata. M, CK and CL represent maker, control group and treatment group, respectively

, figureFileSmall=WaV6L2I4jTT/u0y/ZwqQ+w==, figureFileBig=wdUgcU5Aiyl43pYkt/rC0Q==, tableContent=null), ArticleFig(id=1218970839714812765, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=EN, label=null, caption=null, figureFileSmall=AiNcL+vq7S5Z9Cr39OuidA==, figureFileBig=bsxJNTIYMbFyQ8bX/ButqA==, tableContent=null), ArticleFig(id=1218970839798698850, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=CN, label=Figure 2, caption=

The differentially expressed genes through transcriptome sequencing. A: The volcanic map of differential expression genes; B: The number of up-regulated and down-regulated genes of R. crenulata inoculated with P. fortinii compared to CK

, figureFileSmall=AiNcL+vq7S5Z9Cr39OuidA==, figureFileBig=bsxJNTIYMbFyQ8bX/ButqA==, tableContent=null), ArticleFig(id=1218970839878390629, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=EN, label=null, caption=null, figureFileSmall=jlxqSlhnYjrcuMyBUMK7FQ==, figureFileBig=ZfeNwNgiGDReNd7CNZneaQ==, tableContent=null), ArticleFig(id=1218970839979053927, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=CN, label=Figure 3, caption=

Differential gene expression and pathway in R. crenulata inoculated with P. fortinii based on RNA-seq and Q-PCR. Red and green mark was up-regulated and down-regulated genes and their relative quantity based on CK, respectively. In gene expression level illustration: The bars of PR1 and PR2 refer to left-X-axis; the bars of PISD, MGLL, DEGS, SNW1/SKIP, IAA, AUX1 and ERF1 refer to right-Y-axis

, figureFileSmall=jlxqSlhnYjrcuMyBUMK7FQ==, figureFileBig=ZfeNwNgiGDReNd7CNZneaQ==, tableContent=null), ArticleFig(id=1218970840067134314, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=EN, label=null, caption=null, figureFileSmall=ls/JE3Nmpeml5wO4KS4mTQ==, figureFileBig=C2FcSVNaX2vgIgU7ce9u+g==, tableContent=null), ArticleFig(id=1218970840155214702, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=CN, label=Figure 4, caption= Melting curves of genes in metabolic pathways and reference genes in <i>R. crenulata</i> inoculated with <i>P. fortinii</i>. 1-2: Reference genes (1: <i>PCS</i>; 2: <i>GAPDH</i>); 3-11: Target genes (3: <i>DEGS</i>; 4: <i>PR1</i>; 5: <i>IAA</i>; 6: <i>ERF1</i>; 7: <i>AUX1</i>; 8: <i>PR2</i>; 9:<i>SNW1</i>; 10: <i>PISD</i>; 11: <i>MGLL</i>). <i>Y</i>-axis and <i>X</i>-axis are - d(RFU)/dT and temperature, respectively , figureFileSmall=ls/JE3Nmpeml5wO4KS4mTQ==, figureFileBig=C2FcSVNaX2vgIgU7ce9u+g==, tableContent=null), ArticleFig(id=1218970840276849525, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=EN, label=null, caption=null, figureFileSmall=30AYvVQd+1w/lXMBkI/fwg==, figureFileBig=yPj4fnDfTdiih/3ZSr3jEg==, tableContent=null), ArticleFig(id=1218970840415261560, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=CN, label=Figure 5, caption=

Effect of endophytic fungi on gene expression of R. crenulata in the lipid metabolic pathway. n = 3, $\overline{x}±s$. *P < 0.05 vs control. PISD is involved in the metabolism of glycerophos-pholipids; MGLL is involved in glyceride metabolism; DEGS is involved in sphingolipid metabolism

, figureFileSmall=30AYvVQd+1w/lXMBkI/fwg==, figureFileBig=yPj4fnDfTdiih/3ZSr3jEg==, tableContent=null), ArticleFig(id=1218970840520119164, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=EN, label=null, caption=null, figureFileSmall=WlWGGlPxeJmS8crR//4rxw==, figureFileBig=eV6/Jwf58VYhFyHijZ41iA==, tableContent=null), ArticleFig(id=1218970840591422339, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=CN, label=Figure 6, caption=

Effect of endophytic fungi on gene expression in signal transduction pathway of R. crenulata. n = 3, $\overline{x}±s$. *P < 0.05, **P < 0.01 vs control. SNW1 is involved in the Notch signaling pathway which regulates the differentiation and development of cells, tissues and organs through the interaction between adjacent cells, and PR1, PR2, AUX1, IAA and ERF1 participant in plant hormone signaling pathway

, figureFileSmall=WlWGGlPxeJmS8crR//4rxw==, figureFileBig=eV6/Jwf58VYhFyHijZ41iA==, tableContent=null), ArticleFig(id=1218970840687891332, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=EN, label=null, caption=null, figureFileSmall=mXf1HfzW1Ba2efAw6mzeug==, figureFileBig=CglFLlkU3B91/YPwF/Irdw==, tableContent=null), ArticleFig(id=1218970840771777417, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=CN, label=Figure 7, caption=

Effect of endophytic fungi on the gene expression in environmental adaptation pathway of R. crenulata. n = 3, $\overline{x}±s$. *P < 0.05 vs control. PR1 and PR2 are involved in plant-fungus interaction

, figureFileSmall=mXf1HfzW1Ba2efAw6mzeug==, figureFileBig=CglFLlkU3B91/YPwF/Irdw==, tableContent=null), ArticleFig(id=1218970840868246414, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
Genetic codeGene nameGene symbolPrimer (5'–3')Amplicon size/bp
c3416_g1_i1Sphingolipid delta-4 desaturaseDEGSF: ACCAA TACTC CGATT CCA
R: GTCAT CCCAA CCTCC TTA		224
c67561_g1_i1Pathogenesis-related protein 1PR1F: GCGAC TTGGA TGTGC TAA
R: GACGA AACTG AAACT ACTCT GC		125
c74401_g1_i1Auxin-responsive protein IAAIAAF: CTGGA TGCTT GTAGG AGA
R: AAGTC GGTGA TTTGT TGA		174
c76552_g2_i1Ethylene-responsive transcription factor 1ERF1F: CTTTC GGTTC TTCCA ATT
R: CACCT CGGTA CGACT TCT		201
c78146_g1_i1Auxin influx carrierAUX1F: CACCA AGGTT TGTAG GGAGC
R: AAGGC ATTAG CCGTA GCG		238
c83837_g1_i2Pathogenesis-related protein 2PR2F: TCATT TGAAC TGACG ACGAA
R: TGTGC TAAGG TGAGG TGC		129
c89485_g1_i1SNW domain-containing protein 1SNW1/SKIPF: CTATT GCGAG ACAAG GACA
R: ACATT CACAT TCTTC GGTTT		211
c106081_g1_i1Phosphatidylserine decarboxylasePISDF: CTTCT ACCGC ACCCT AAA
R: TTCCC TTGAC TTGTT CCA		131
c139123_g1_i1Acylglycerol lipaseMGLLF: CTATA ATTCT GGCAT CGG
R: AGACT GTCAC GGAAC TGG		120
), ArticleFig(id=1218970840964715409, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551242020798470, language=CN, label=Table 1, caption=

Amplification primers with Q-PCR for differential genes of metabolic pathway in R. crenulata inoculated with P. fortinii

, figureFileSmall=null, figureFileBig=null, tableContent=
Genetic codeGene nameGene symbolPrimer (5'–3')Amplicon size/bp
c3416_g1_i1Sphingolipid delta-4 desaturaseDEGSF: ACCAA TACTC CGATT CCA
R: GTCAT CCCAA CCTCC TTA		224
c67561_g1_i1Pathogenesis-related protein 1PR1F: GCGAC TTGGA TGTGC TAA
R: GACGA AACTG AAACT ACTCT GC		125
c74401_g1_i1Auxin-responsive protein IAAIAAF: CTGGA TGCTT GTAGG AGA
R: AAGTC GGTGA TTTGT TGA		174
c76552_g2_i1Ethylene-responsive transcription factor 1ERF1F: CTTTC GGTTC TTCCA ATT
R: CACCT CGGTA CGACT TCT		201
c78146_g1_i1Auxin influx carrierAUX1F: CACCA AGGTT TGTAG GGAGC
R: AAGGC ATTAG CCGTA GCG		238
c83837_g1_i2Pathogenesis-related protein 2PR2F: TCATT TGAAC TGACG ACGAA
R: TGTGC TAAGG TGAGG TGC		129
c89485_g1_i1SNW domain-containing protein 1SNW1/SKIPF: CTATT GCGAG ACAAG GACA
R: ACATT CACAT TCTTC GGTTT		211
c106081_g1_i1Phosphatidylserine decarboxylasePISDF: CTTCT ACCGC ACCCT AAA
R: TTCCC TTGAC TTGTT CCA		131
c139123_g1_i1Acylglycerol lipaseMGLLF: CTATA ATTCT GGCAT CGG
R: AGACT GTCAC GGAAC TGG		120
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Treatment2.1571.97
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Quality detection of total RNA extracted from R. crenulata

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内生菌Phialocephala fortinii影响大花红景天代谢的差异途径及基因表达
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王梦亮 1 , 高小音 1, 2 , 崔晋龙 1, * , 王俊宏 1
药学学报 | 研究论文 生药学 2018,53(12): 2122-2128
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药学学报 | 研究论文 生药学 2018, 53(12): 2122-2128
内生菌Phialocephala fortinii影响大花红景天代谢的差异途径及基因表达
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王梦亮1, 高小音1, 2, 崔晋龙1, * , 王俊宏1
作者信息
  • 1.山西大学应用化学研究所, 山西 太原 030006
  • 2.山西大学生物技术研究所, 山西 太原 030006

通讯作者:

* 崔晋龙, Tel:86-351-7016101, E-mail:
Differential pathway and gene expressions of endophytic Phialocephala fortinii inoculated on Rhodiola crenulata
Meng-liang WANG1, Xiao-yin GAO1, 2, Jin-long CUI1, * , Jun-hong WANG1
Affiliations
  • 1. Institute of Applied Chemistry, Taiyuan 030006, China
  • 2. Institute of Biotechnology, Shanxi University, Taiyuan 030006, China
出版时间: 2018-12-12 doi: 10.16438/j.0513-4870.2018-0709
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摘要
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药用植物发育和代谢受到多种因素的影响，而内生真菌对植物的影响是近年来才得以重视并逐步成为热点。为探索内生真菌Phialocephala fortinii对大花红景天基因及代谢途径的影响，本实验通过RNA转录组测序筛选差异表达基因；并用Real-time PCR对基因表达进行验证，利用2-△△Ct的方法分析脂代谢、信号转导和环境适应性等重要代谢途径差异基因的相对表达量。结果表明，验证结果与测序结果一致，内生真菌P.fortinii能够促进大花红景天脂代谢途径中的差异基因表达量；它对信号转导途径中相关基因表达的影响不同，但控制Notch信号通路途径的基因表达量显著提高，为对照组的34倍；对适应性代谢途径差异基因表达上调，进而促进红景天环境适应能力。研究结果旨在为揭示内生真菌与红景天的互作机制、红景天道地性形成原因、内生真菌功能的开发等提供分子基础数据。

福廷瓶头霉  /  RNA-Seq  /  内参基因  /  实时荧光定量PCR  /  植物-真菌互作

The development and metabolism of medicinal plant is affected by many factors, among which the effect from endophytic fungi has been noticed recently and has become one of hot fields. In order to explore the effect of endophytic fungi on gene expression in R. crenulata, RNA-sequencing was used to find genes involved in metabolic pathways, and the differential genes were verified by real-time fluorescent quantitative PCR. The method of 2-△△Ct was used to analyze the relative expression levels of genes in related metabolic pathways, which was used to verify the result of transcriptomics sequencing. The results showed that the endophytic fungus, P. fortinii, could up-regulate the gene expression in lipid metabolic pathway of R. crenulata. In signal transduction pathway, the genes were influenced at different level but the gene expressions were significantly increased under control of Notch signaling pathway, which was 34 times of that in control. The gene expressions of environmental adaption pathway were up-regulated in R. crenulata after inoculation of P. fortinii. This study could provide help for further understanding on mechanism of plant-fungus interaction, root cause of geoherbalism of medicinal plant and exploring bio-function of endophytic fungi.

Phialocephala fortinii  /  RNA-Seq  /  internal reference genes  /  real-time fluorescent quantitative PCR  /  plant-fungus interaction
王梦亮, 高小音, 崔晋龙, 王俊宏. 内生菌Phialocephala fortinii影响大花红景天代谢的差异途径及基因表达. 药学学报, 2018 , 53 (12) : 2122 -2128 . DOI: 10.16438/j.0513-4870.2018-0709
Meng-liang WANG, Xiao-yin GAO, Jin-long CUI, Jun-hong WANG. Differential pathway and gene expressions of endophytic Phialocephala fortinii inoculated on Rhodiola crenulata[J]. Acta Pharmaceutica Sinica, 2018 , 53 (12) : 2122 -2128 . DOI: 10.16438/j.0513-4870.2018-0709
正文
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基因表达变化能直接或间接地影响药用植物代谢通路, 从而影响药用植物活性成分的积累[1]。然而, 药用植物基因表达受到多种因素的影响。其中, 内生真菌被认为是植物生长发育和代谢过程中不可或缺的组成部分, 可通过横向基因转移、诱导效应、生物转化等途径改变宿主基因表达和代谢途径, 对植物的生长、发育产生影响[2]。近年来, 陆续发现内生真菌在改变植物次生代谢产物的合成[3]、促进生长[4]、提高植物产量[5]和增强植物抵抗胁迫的能力[6]等方面具有重要作用。因此, 内生真菌与植物的互作关系, 已成为药用植物品质研究的前沿领域[7, 8]。
红景天(Rhodiola crenulata)属于景天科红景天属的多年生草本或亚灌木药用植物。其中, 主要品种为大红花景天, 生长于青藏高原, 被誉为“高原人参”, 并作为红景天唯一的基源植物被历届《中国药典》所收录, 主要具有抗氧化、抗辐射、抗缺氧等药理活性[6]。本研究是将前期发现的活性内生真菌P. fortinii接种于大花红景天, 从转录组水平探索该菌对大花红景天代谢途径中基因表达变化及互作机制, 为真菌-植物互作发育和代谢机制提供依据。
材料与方法
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材料  红景天组培苗:由本课题组建立的大花红景天组培苗体系进行转接[9], 稳定生长约一周后用于菌-苗互作实验。
供试菌株  分离自长白红景天根部的内生真菌, 经筛选, 具有促进大花红景天关键酶和次生代谢产物积累的作用[6, 10]。经鉴定, 隶属于福廷瓶头霉Phialocephala fortinii [11], GenBank库登记编号为KJ542299, 保藏于山西大学应用化学研究所[6]。
试剂与仪器  柱式RNA抽提纯化试剂盒(上海生工生物工程股份有限公司); 反转录试剂盒GoScriptTM Reverse Transcription Mix和实时荧光定量PCR试剂盒GoTaq® qPCRMaster Mix (北京赛奥生物技术有限公司); BIO-RAD CFx96TM Real-Time system (Bio-rad, USA)。
内生真菌与红景天组培苗共生培养  无菌条件下, 打取直径为0.5 cm的P. fortinii菌饼3块, 均匀接种于组培苗周围, 培养条件见文献[9]。以菌-苗紧密接触生长开始计时[12], 培养10天, 采集互作组培苗, 设置无菌组培苗作为对照(CK), 试验样品全部设置3个重复。将采集的样品在液氮中速冻30 min后, 保存于-80 ℃冰箱, 用于RNA的提取。
总RNA提取、cDNA合成及转录组测序  取约25 mg冻存样品(处理组和对照组)加入液氮迅速研磨成粉状, 采用柱式RNA抽提纯化试剂盒提取总RNA, 利用超微量分光光度计(Thermo Scientific NanoDrop Lite, 美国)和1%琼脂糖凝胶电泳对RNA质量进行检测。cDNA第一链的合成按照反转录试剂盒GoScript™ Reverse Transcription Mix试剂盒说明书要求进行。之后将构建好的cDNA文库送往上海派森诺生物科技股份有限公司进行无参转录组测序, 测序平台为Illumina Nextseq 500。
引物的合成及验证  利用Primer Premier 5软件(Premier公司, 加拿大)设计差异基因引物, 包括鞘脂类δ-4-脱氢酶基因(DEGS)、病程相关蛋白1基因(PR1)、生长素响应蛋白IAA基因(IAA)、乙烯响应转录因子1基因(ERF1)、生长素转运载体基因(AUX1)、病程相关蛋白2基因(PR2)、含SNW结构域蛋白1基因(SNW1/SKIP)、磷脂酰丝氨酸脱羧酶基因(PISD)和酰基甘油脂肪酶基因(MGLL)的引物(表 1), 然后通过琼脂糖凝胶电泳分析引物的特异性, 验证其引物能否进行特异性扩增。
qRT-PCR分析  单个内参基因表达不稳定, 而常常采用多个内参基因进行qRT-PCR校正, 以得到更为准确的结果[13], 本研究以植物螯合肽合成酶基因(PCS) (GenBank No. AJ548472.1)和甘油醛磷酸脱氢酶基因(GAPDH) (GenBank No. HQ156465)作为内参基因, 设计引物F1: 5'-TGAGGTGGCGACTG ATAACC-3', R1: 5'-GGGAGAACCCTCCGGTACAA-3'; F2: 5'-GCACGCCAATCCAGAACAAG-3', R2: 5'-TGGGCGTGATATCTCCTCGT-3'。
RT-PCR反应采用实时荧光定量PCR试剂盒GoTaq® qPCRMaster Mix, 在BIO-RAD CFx96TM荧光定量PCR仪上进行实验。反应总体系为20 μL: GoTaq® qPCRMaster Mix (2×) 10 μL、上下游引物(10 μmol·L-1)各0.6 μL、模板1 μL、Nuclease-Free Water 7.8 μL。扩增程序采用3步法(95 ℃预变性10 min; 95 ℃变性15 s, 55 ℃退火30 s, 72 ℃延伸30 s, 45个循环; 95 ℃ 15 s, 60 ℃ 15 s, 95 ℃ 15 s)。
数据处理  通过内参基因进行归一化比较, 首先利用Excel 2016软件计算对照组和处理组样品中目的基因和内参基因的相对表达量, 选用2-ΔΔCt的方法[14]计算处理组基因相对表达量, 生物学重复3次, 取平均值; 运用SPSS软件单因素分析(One Way ANOVA)数据差异性, 进行差异性分析显著性差异(P < 0.05)和极显著性差异(P < 0.01)。
结果与分析
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1 总RNA的质量检测
通过1.0%琼脂糖凝胶电泳对样品总RNA进行检测发现28S rRNA和18S rRNA条带清晰且无弥散现象, 表明RNA的完整性良好(图 1); 经过NanoDrop检测发现OD 260/280接近2.0左右(表 2), 说明RNA的纯度较高, 达到后续实验要求。
2 内生真菌P. fortinii与大花红景天互作的转录组测序
转录组测序结果表明, 相比于对照, 真菌P. fortinii对大花红景天的3 200个基因表达产生了影响, 其中上调基因3 048个, 下调基因152个(图 2)。其中, 经过KEGG数据库富集分析表明, 影响较大且最为重要的为脂代谢(lipid metabolism)、信号转导(signal transduction)和环境适应(environmental adaption) 3条代谢途径。因此本实验选取这3个途径中部分影响较大的基因为研究对象, 所研究基因如下:鞘脂类δ-4-脱氢酶基因(DEGS)、病程相关蛋白1基因(PR1)、生长素响应蛋白IAA基因(IAA)、乙烯响应转录因子1基因(ERF1)、生长素转运载体基因(AUX1)、病程相关蛋白2基因(PR2)、含SNW结构域蛋白1基因(SNW1/SKIP)、磷脂酰丝氨酸脱羧酶基因(PISD)和酰基甘油脂肪酶基因(MGLL) 9个基因(图 3)。
3 引物特异性分析
扩增产物经1.0%琼脂糖凝胶电泳检测, 条带大小与期望值相一致, 不存在其他非特异性扩增条带。进一步通过分析参与各代谢途径中基因的熔解曲线(图 4), 发现都只产生单一熔解峰, 且各重复样品之间曲线重复性良好, 说明引物满足实验要求。
4 红景天代谢途径中基因的表达分析
4.1 内生真菌对大花红景天脂代谢中基因表达的影响
内生真菌P. fortinii与大花红景天组培苗互作后, 红景天脂代谢基因表达的变化见图 5, 参与脂代谢差异基因的表达量均表现出升高, 其中参与甘油磷脂代谢的基因PISD与对照相比, 其相对表达量明显提高, 达到对照组的7.2倍(P < 0.05), 而涉及甘油酯代谢和鞘脂类代谢的基因MGLL和DEGS, 其表达量也有所提高。结果表明, 内生真菌能够促进宿主红景天脂代谢途径中基因表达。
4.2 内生真菌对大花红景天在信号转导途径中基因表达的影响
由图 6可知, 内生真菌P. fortinii显著提高了红景天中参与Notch信号通路途径基因的表达量, 即达到对照组的34倍(P < 0.05)。参与植物激素信号转导途径的5个基因中, PR1和PR2的相对表达量与对照组相比, 分别提高了3.8倍和0.96倍, 而其余3个基因AUX1、IAA和ERF1表达量均呈现下降趋势。结果表明, 内生真菌能够调节植物细胞的分化, 并且诱导激素调节的变化, 从而促进植物的生长和发育。
4.3 内生真菌对大花红景天在环境适应中基因表达的影响
由图 7可知, 活性内生真菌能够促进宿主植物红景天对环境的适应, 其中基因PR1的相对表达量明显提高, 为对照组的4.8倍, 且具有显著性差异(P < 0.05), 而PR2的表达量虽然呈现升高, 但与对照相比仅提高了0.96倍, 且两个基因均参与真菌与植物互作。研究表明, 内生真菌P. fortinii能够增强其宿主植物对生物胁迫的耐受性和对微生物的抗性。
讨论
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大花红景天是《神农本草经》、《四部药典》和《本草纲目》等医书中列为上品的高山药用植物, 具滋补、延缓衰老、抗疲劳等方面的药理功能[15]。近年来, 为了提高大花红景天的品质, 许多学者在基因调控[16]、活性成分的合成途径[17]及次级代谢产物调控[18]等方面进行了广泛的研究。但是, 影响大花红景天品质的因素尚不完全明确, 影响代谢途径中的基因表达及次级代谢物合成的机制尚不清楚。研究表明, P. fortinii能与植物根形成黑色有隔菌丝结构, 促进毛棉杜鹃幼苗磷、氮元素的吸收和生长[11]。研究发现, P. fortinii具有促进大花红景天生长、提高红景天苷等活性成分积累以及促进红景天苷合成途径中关键酶基因表达等的生物活性[6, 9, 19]。通过转录组测序表明, 内生真菌对宿主植物脂代谢、信号转导途径、环境适应性等途径中基因表达产生重要影响, 这对挖掘P. fortinii生物功能, 揭示该菌与红景天互作的分子机制, 对深入认识红景天道地性形成因素提供重要的补充。
脂代谢是生物体生命活动的主要代谢途径之一, 内生真菌促进红景天脂代谢途径中基因的表达, 可能与内生真菌的次级代谢有关, 内生真菌可通过产生多样化合物, 作为植物代谢物前体或直接成为其组成部分, 或被宿主吸收利用, 进而影响宿主生长的发育和代谢[20, 21]。①鞘脂不仅是生物膜结构的重要成分, 也是重要的生物活性分子, 鞘脂及其衍生物是许多细胞活动进程中重要的动力调节器, 参与了许多重要的信号转导过程, 进而使细胞产生各种不同的生物学功能。研究发现鞘脂δ-4-去饱和酶(DEGS)基因表达量提高, 这有助于促进不饱和鞘脂类不饱和脂肪酸分子的合成; 对生物膜稳定性、细胞信号、逆境反应、病原性防御和程序性细胞死亡进程中仍然发挥着重要作用[22]。②磷酯酰丝氨酸是细胞膜的活性物质, 具有延缓衰老的作用, 磷脂酰丝氨酸脱羧酶(PISD)能够将磷酯酰丝氨酸脱羧形成磷脂酰乙醇胺。研究发现PISD基因表达提高, 增强脱羧作用, 从而降低宿主抗衰老的功能。③酰基甘油脂肪酶(MGLL)是一类促进脂肪分解成甘油和脂肪酸的水解酶, 其基因表达量提高, 进一步能够促进甘油和脂肪酸的合成, 为宿主植物的生命活动提供能量等。
植物信号转导途径能够控制相关基因表达和引起特定的生理生化反应[20], 研究发现内生真菌与红景天互作后, 参与Notch信号通路的基因SNW1/SKIP相对表达量显著提高, 表明内生真菌P. fortinii可能通过调节宿主细胞、组织等的分化, 从而来促进植物的生长发育[23]; 激素调节是维持植物体稳态的调节方式, 本研究发现控制激素调节的基因表达量变化不一致, 生长素响应蛋白基因(IAA)、生长素转运载体基因(AUX1)和乙烯响应转录因子基因(ERF1)降低, 表明活性菌与植物互作一定时间后, 生长素的合成和转运能力降低, 从而可能会进一步影响植物的生长。
目前已有研究发现内生真菌能够提高宿主植物的抗性以及增强植物对环境的适应能力[24]。但是, 此研究发现病程相关蛋白基因(PR1和PR2)表达量提高, 说明活性内生真菌在一定程度上可能转换为病原菌, 从而能够引起宿主植物的防御反应[25], 进一步增强其植物的适应能力。本研究表明P. fortinii增强了宿主植物在环境适应方面的能力, 这可能是由于植物的防御和应激反应引起的。
尽管该研究从分子水平揭示了部分重要代谢途经的基因表达情况, 但是内生真菌影响红景天代谢的途径及基因还需要进一步研究。基因表达的调控不仅受到多时空因素的影响, 而且会呈现出多种复杂的影响结果。总之, 对于这类问题系统、深入的研究, 将为全面开发利用大花红景天内生真菌资源、寻找提高红景天品质的方法提供新途径。
基金
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  • 国家自然科学基金资助项目(31270383)
  • 国家自然科学基金资助项目(31670328)
  • 山西省自然科学基金资助项目(2014011029-1)
  • 山西省"1331工程"重点实验室-功能分子化学(晋教科[2017]14号)
文献
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2018年第53卷第12期
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文章信息
doi: 10.16438/j.0513-4870.2018-0709
  • 接收时间：2018-08-03
  • 首发时间：2026-01-15
  • 出版时间：2018-12-12
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出版历史
  • 收稿日期：2018-08-03
  • 修回日期：2018-09-26
基金
国家自然科学基金资助项目(31270383)
国家自然科学基金资助项目(31670328)
山西省自然科学基金资助项目(2014011029-1)
山西省"1331工程"重点实验室-功能分子化学(晋教科[2017]14号)
作者信息
    1.山西大学应用化学研究所, 山西 太原 030006
    2.山西大学生物技术研究所, 山西 太原 030006

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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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