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Article(id=1221483613724525516, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1221483606648734411, articleNumber=null, orderNo=null, doi=10.16438/j.0513-4870.2020-0705, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1588953600000, receivedDateStr=2020-05-09, revisedDate=1591200000000, revisedDateStr=2020-06-04, acceptedDate=null, acceptedDateStr=null, onlineDate=1769153987579, onlineDateStr=2026-01-23, pubDate=1607702400000, pubDateStr=2020-12-12, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1769153987579, onlineIssueDateStr=2026-01-23, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1769153987579, creator=13701087609, updateTime=1769153987579, updator=13701087609, issue=Issue{id=1221483606648734411, tenantId=1146029695717560320, journalId=1189982191388893191, year='2020', volume='55', issue='12', pageStart='2751', pageEnd='2998', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1769153985893, creator=13701087609, updateTime=1769154367876, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1221485208856085269, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1221483606648734411, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1221485208856085270, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1221483606648734411, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=2982, endPage=2988, ext={EN=ArticleExt(id=1221483614236230638, articleId=1221483613724525516, tenantId=1146029695717560320, journalId=1189982191388893191, language=EN, title=The mechanism of methyl jasmonate-induced accumulation of hydroxysafflor yellow A in safflower of different colors, columnId=1190335348761793317, journalTitle=Acta Pharmaceutica Sinica, columnName=Original Articles, runingTitle=null, highlight=null, articleAbstract=

To explore the mechanism hydroxysafflor yellow A (HSYA) biosynthesis and regulation, the effect of methyl jasmonate (MeJA) treatment on gene expression related to the biosynthesis of hydroxysafflor yellow A (HSYA) was analyzed, and expression differences in genes involved in HSYA biosynthesis in safflower of different colors was quantified. MeJA at concentrations of 0, 50, 100, and 200 μmol·L-1 was sprayed onto safflower florets to determine the optimal concentration of MeJA. Safflower was treated with 100 μmol·L-1 MeJA and florets were harvested 0, 3, 6, 12 and 24 h after treatment. The content of MeJA was determined by high performance liquid chromatography (HPLC). RNA was extracted from safflower florets treated with 100 μmol·L-1 MeJA for 6 h. The transcription of key genes involved in the biosynthesis of HSYA was quantified by qRT-PCR and differentially expressed genes were identified. The content of HSYA increased after treatment with MeJA, with 100 μmol·L-1 MeJA treatment for 6 h having the greatest effect on HSYA accumulation. qRT-PCR results showed that MeJA could significantly increase the transcription of HSYA biosynthesis genes including PAL2, PAL4, 4CL2, 4CL4, 4CL5, CHS3, CHS4 and CHI2. The content of HSYA differed between safflowers of different colors with a trend of red > orange-yellow > yellow > white. The results of qRT-PCR showed that the expression of CHS1 and CHI2 in red, orange and yellow safflower was significantly higher than that in white safflower. These results indicate that MeJA promotes the accumulation of HSYA by up-regulating the expression of genes involved in the biosynthesis of HSYA such as PAL2, PAL4, 4CL2, 4CL4, 4CL5, CHS3, CHS4 and CHI2, and the variation of HSYA content in safflower of different colors was related to a difference in the level of expression of CHS1 and CHI2.

, correspAuthors=Hui-zhen LIANG, authorNote=null, correspAuthorsNote=null, copyrightStatement=Copyright ©2020 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=Zheng-wei TAN, Lei LI, Hong-qi YANG, Yong-liang YU, Lan-jie XU, Wei DONG, Wei XIA, Xin-ming MA, Hui-zhen LIANG), CN=ArticleExt(id=1221483615918145643, articleId=1221483613724525516, tenantId=1146029695717560320, journalId=1189982191388893191, language=CN, title=茉莉酸甲酯诱导和不同花色红花中羟基红花黄色素A积累差异的机制分析, columnId=1190335348896011050, journalTitle=药学学报, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=

通过分析茉莉酸甲酯(methyl jasmonate，MeJA)处理对羟基红花黄色素A(hydroxysafflor yellow A，HSYA)生物合成相关基因表达影响及不同花色HSYA生物合成相关基因表达差异，为HSYA生物合成及调控提供参考。首先用0、50、100、200 μmol·L-1的MeJA对离体培养的红花花冠进行处理，找出最适MeJA处理浓度，然后用100 μmol·L-1 MeJA处理红花，在处理后的0、3、6、12、24 h不同时间点采样，用高效液相色谱法对HSYA的含量进行定量分析，找出最佳处理时间；以100 μmol·L-1 MeJA处理6 h的红花花冠提取RNA，通过qRT-PCR对参与HSYA生物合成的关键基因进行定量分析，找出表达差异基因；用HPLC和qRT-PCR对不同花色红花品系的HSYA含量及其相关合成基因进行定量分析，找出差异基因。结果表明，不同浓度的MeJA处理均能够显著增加HSYA的积累，其中100 μmol·L-1 MeJA处理6 h后HSYA的含量达到最高峰。qRT-PCR结果显示，MeJA处理红花花冠能够显著提高HSYA合成关键基因PAL2、PAL4、4CL2、4CL4、4CL5、CHS3、CHS4和CHI2的转录水平。不同花色红花中HSYA含量不同，呈现红色>橙黄色>黄色>白色的趋势，qRT-PCR结果表明CHS1和CHI2在红色、橙黄色、黄色红花中的表达量明显高于白色。以上研究表明MeJA对HSYA合成的调控主要通过影响PAL2、PAL4、4CL2、4CL4、4CL5、CHS3、CHS4和CHI2的转录水平来实现；CHS1和CHI2的基因表达差异是不同花色红花HSYA含量不同的主要因素。

, correspAuthors=梁慧珍, authorNote=null, correspAuthorsNote=
*梁慧珍, Tel:86-371-65738565, E-mail:
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PAL: Phenylalanine ammonia lyase; C4H: Innamate 4-hydroxylase; 4CL: 4-Coumaroyl CoA-ligase; CHS: Chalcone synthase; CGT: <i>C</i>-Glucosyltransferase; P450: Cytochrome P450; CHI: Chalcone isomerase; FNSⅠ: Flavone synthase Ⅰ; F3H: Flavanone 3-hydroxylase; DFR: Dihydroflavonol 4-reductase; ANS: Anthocyanidin synthase; UGT: UDP-glycosyltransferases , figureFileSmall=x3Px7ZDnLNK6x/x0tGRmDw==, figureFileBig=yCCLwkliv7ZGD2YQQm2RVA==, tableContent=null), ArticleFig(id=1221483620859036077, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1221483613724525516, language=EN, label=null, caption=null, figureFileSmall=G07ySF2LOydI6O0HIuiaYA==, figureFileBig=yGDWY6S3uJ+jFV+CbL2gGA==, tableContent=null), ArticleFig(id=1221483620951310770, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1221483613724525516, language=CN, label=Figure 2, caption= Effect of different concentration of MeJA (A) and different treatment time (B) on HSYA content. A: <sup>*</sup><i>P</i> < 0.05, <sup>**</sup><i>P</i> < 0.01 <i>vs</i> 0 μmol·L<sup>-1</sup>; B: <sup>**</sup><i>P</i> < 0.01 <i>vs</i> 0 h. MeJA: Methyl jasmonate , figureFileSmall=G07ySF2LOydI6O0HIuiaYA==, figureFileBig=yGDWY6S3uJ+jFV+CbL2gGA==, tableContent=null), ArticleFig(id=1221483621051974071, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1221483613724525516, language=EN, label=null, caption=null, figureFileSmall=CUvOLYB+KG4bZDWgoEqMLQ==, figureFileBig=nBcrjx7xzCDB4a92sC/qWQ==, tableContent=null), ArticleFig(id=1221483621131665850, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1221483613724525516, language=CN, label=Figure 3, caption= Effect of MeJA treatment on expression of related genes involved in HSYA biosynthesis. <sup>*</sup><i>P</i> < 0.05, <sup>**</sup><i>P</i> < 0.05 <i>vs</i> Mock , figureFileSmall=CUvOLYB+KG4bZDWgoEqMLQ==, figureFileBig=nBcrjx7xzCDB4a92sC/qWQ==, tableContent=null), ArticleFig(id=1221483621232329151, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1221483613724525516, language=EN, label=null, caption=null, figureFileSmall=7R5LnSg7eYnzDLbDimKmgw==, figureFileBig=9uTrWeIqqZHrc9GZ3wqiVQ==, tableContent=null), ArticleFig(id=1221483621370741190, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1221483613724525516, language=CN, label=Figure 4, caption= The phenotype of safflower with different colors (white, yellow, orange-yellow, red) , figureFileSmall=7R5LnSg7eYnzDLbDimKmgw==, figureFileBig=9uTrWeIqqZHrc9GZ3wqiVQ==, tableContent=null), ArticleFig(id=1221483621475598793, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1221483613724525516, language=EN, label=null, caption=null, figureFileSmall=K5b8tAB1BAPfItNZcqBXaQ==, figureFileBig=4bk/tymnSp5IxRGqg5LPug==, tableContent=null), ArticleFig(id=1221483621551096267, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1221483613724525516, language=CN, label=Figure 5, caption= The content of HSYA in safflower with different colors. W: White; Y: Yellow; OY: Orange-yellow; R: Red , figureFileSmall=K5b8tAB1BAPfItNZcqBXaQ==, figureFileBig=4bk/tymnSp5IxRGqg5LPug==, tableContent=null), ArticleFig(id=1221483621647565264, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1221483613724525516, language=EN, label=null, caption=null, figureFileSmall=P0devPE3WFLVs+9O9ZEgoA==, figureFileBig=aG33hyAXrPF0CCgcu7O4bw==, tableContent=null), ArticleFig(id=1221483621710479828, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1221483613724525516, language=CN, label=Figure 6, caption= Quantitative analysis of the expression of related genes involved in HSYA biosynthesis in safflower with different colors. <sup>**</sup><i>P</i> < 0.05 <i>vs</i> white safflower , figureFileSmall=P0devPE3WFLVs+9O9ZEgoA==, figureFileBig=aG33hyAXrPF0CCgcu7O4bw==, tableContent=null), ArticleFig(id=1221483621785977305, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1221483613724525516, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
Gene Forward primer (5′-3′) Reverse primer (5′-3′)
PAL1 TGGAAGCCGTAATCCAAGTC TCCGTGAAGAAATCAACCCT
PAL2 ACGAGGTGAAGAAGATGGTGG AGCCCTAGCTGATTCCGACA
PAL3 AAGTCCTATCGGCCCTCTTC CGTAATCGCTTCCATCCAAA
PAL4 GCTTGCAGATCATACCCATTA GGAACACCATTCCAACCCTC
PAL5 GAAATGGACCCACTGGAGAA CGTGTAGGGCTTTGTTTCTTG
C4H1 TTGGGAGCACGAAGAAGATG TGTCTCGATTGCAGCAACAT
4CL1 TGCTCTGCCATTCTCCTCTG TAGCCCTAATATCGTGACTTTG
4CL2 AGAACATTGAGAAACATAG GAAGGGAACTTCTTGGCACA
4CL3 TGTAAGAAGTCAATGTGTCA ACCGTGTTCATCAATAGTT
4CL4 TTCGCTAAGGAGCCATTTGA TGATTTGGTCACCACGGATA
4CL5 AGCACGCTGGAGGAAATGAT CAACTGGAACCGTCCGAGAA
CHS1 GGTACCGGGATCTCTAGAAGAT TTCTGATCTCGGCAATATCGGT
CHS2 GATCAATGGCAACGGTGTAGT AAATAGCCTTCGACCAAGCA
CHS3 AGACCAGATACACAGTAATG GCTTCCTTTGCCATCT
CHS4 CTCGTCTTCTGCACCACCTC TGTTCTCGGCTAGGTCCTT
CHS5 GACTCTTCGACAACAACT TGCCAACTTCTCTTTCAT
CHI1 TCCGACTCAACTACAATGGC TCCGGTAAACTTCACGAACC
CHI2 AACAACGCCCGATGGATTAG TTCGCTTCAGGAGAAACACC
Ct60s GGCAACACTGGTCTTGATCCAT CTGCGATTGTTGGATAATGGATG
), ArticleFig(id=1221483621920195035, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1221483613724525516, language=CN, label=Table 1, caption=

The primers sequences used for real-time PCR

, figureFileSmall=null, figureFileBig=null, tableContent=
Gene Forward primer (5′-3′) Reverse primer (5′-3′)
PAL1 TGGAAGCCGTAATCCAAGTC TCCGTGAAGAAATCAACCCT
PAL2 ACGAGGTGAAGAAGATGGTGG AGCCCTAGCTGATTCCGACA
PAL3 AAGTCCTATCGGCCCTCTTC CGTAATCGCTTCCATCCAAA
PAL4 GCTTGCAGATCATACCCATTA GGAACACCATTCCAACCCTC
PAL5 GAAATGGACCCACTGGAGAA CGTGTAGGGCTTTGTTTCTTG
C4H1 TTGGGAGCACGAAGAAGATG TGTCTCGATTGCAGCAACAT
4CL1 TGCTCTGCCATTCTCCTCTG TAGCCCTAATATCGTGACTTTG
4CL2 AGAACATTGAGAAACATAG GAAGGGAACTTCTTGGCACA
4CL3 TGTAAGAAGTCAATGTGTCA ACCGTGTTCATCAATAGTT
4CL4 TTCGCTAAGGAGCCATTTGA TGATTTGGTCACCACGGATA
4CL5 AGCACGCTGGAGGAAATGAT CAACTGGAACCGTCCGAGAA
CHS1 GGTACCGGGATCTCTAGAAGAT TTCTGATCTCGGCAATATCGGT
CHS2 GATCAATGGCAACGGTGTAGT AAATAGCCTTCGACCAAGCA
CHS3 AGACCAGATACACAGTAATG GCTTCCTTTGCCATCT
CHS4 CTCGTCTTCTGCACCACCTC TGTTCTCGGCTAGGTCCTT
CHS5 GACTCTTCGACAACAACT TGCCAACTTCTCTTTCAT
CHI1 TCCGACTCAACTACAATGGC TCCGGTAAACTTCACGAACC
CHI2 AACAACGCCCGATGGATTAG TTCGCTTCAGGAGAAACACC
Ct60s GGCAACACTGGTCTTGATCCAT CTGCGATTGTTGGATAATGGATG
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茉莉酸甲酯诱导和不同花色红花中羟基红花黄色素A积累差异的机制分析
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谭政委 1 , 李磊 1 , 杨红旗 1 , 余永亮 1 , 许兰杰 1 , 董薇 1 , 夏伟 1 , 马新明 2 , 梁慧珍 1, *
药学学报 | 研究论文 2020,55(12): 2982-2988
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药学学报 | 研究论文 2020, 55(12): 2982-2988
茉莉酸甲酯诱导和不同花色红花中羟基红花黄色素A积累差异的机制分析
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谭政委1, 李磊1, 杨红旗1, 余永亮1, 许兰杰1, 董薇1, 夏伟1, 马新明2, 梁慧珍1, *
作者信息
  • 1.河南省农业科学院芝麻研究中心, 河南 郑州 450002
  • 2.河南农业大学信息与管理科学学院, 河南 郑州 450002

通讯作者:

*梁慧珍, Tel:86-371-65738565, E-mail:
The mechanism of methyl jasmonate-induced accumulation of hydroxysafflor yellow A in safflower of different colors
Zheng-wei TAN1, Lei LI1, Hong-qi YANG1, Yong-liang YU1, Lan-jie XU1, Wei DONG1, Wei XIA1, Xin-ming MA2, Hui-zhen LIANG1, *
Affiliations
  • 1. Henan Sesame Research Center, Henan Academy of Agricultural Sciences, Zhengzhou 450002, China
  • 2. College of Information and Management Science, Henan Agricultural University, Zhengzhou 450002, China
出版时间: 2020-12-12 doi: 10.16438/j.0513-4870.2020-0705
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摘要
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通过分析茉莉酸甲酯(methyl jasmonate，MeJA)处理对羟基红花黄色素A(hydroxysafflor yellow A，HSYA)生物合成相关基因表达影响及不同花色HSYA生物合成相关基因表达差异，为HSYA生物合成及调控提供参考。首先用0、50、100、200 μmol·L-1的MeJA对离体培养的红花花冠进行处理，找出最适MeJA处理浓度，然后用100 μmol·L-1 MeJA处理红花，在处理后的0、3、6、12、24 h不同时间点采样，用高效液相色谱法对HSYA的含量进行定量分析，找出最佳处理时间；以100 μmol·L-1 MeJA处理6 h的红花花冠提取RNA，通过qRT-PCR对参与HSYA生物合成的关键基因进行定量分析，找出表达差异基因；用HPLC和qRT-PCR对不同花色红花品系的HSYA含量及其相关合成基因进行定量分析，找出差异基因。结果表明，不同浓度的MeJA处理均能够显著增加HSYA的积累，其中100 μmol·L-1 MeJA处理6 h后HSYA的含量达到最高峰。qRT-PCR结果显示，MeJA处理红花花冠能够显著提高HSYA合成关键基因PAL2、PAL4、4CL2、4CL4、4CL5、CHS3、CHS4和CHI2的转录水平。不同花色红花中HSYA含量不同，呈现红色>橙黄色>黄色>白色的趋势，qRT-PCR结果表明CHS1和CHI2在红色、橙黄色、黄色红花中的表达量明显高于白色。以上研究表明MeJA对HSYA合成的调控主要通过影响PAL2、PAL4、4CL2、4CL4、4CL5、CHS3、CHS4和CHI2的转录水平来实现；CHS1和CHI2的基因表达差异是不同花色红花HSYA含量不同的主要因素。

红花  /  茉莉酸甲酯  /  羟基红花黄色素A  /  表达差异

To explore the mechanism hydroxysafflor yellow A (HSYA) biosynthesis and regulation, the effect of methyl jasmonate (MeJA) treatment on gene expression related to the biosynthesis of hydroxysafflor yellow A (HSYA) was analyzed, and expression differences in genes involved in HSYA biosynthesis in safflower of different colors was quantified. MeJA at concentrations of 0, 50, 100, and 200 μmol·L-1 was sprayed onto safflower florets to determine the optimal concentration of MeJA. Safflower was treated with 100 μmol·L-1 MeJA and florets were harvested 0, 3, 6, 12 and 24 h after treatment. The content of MeJA was determined by high performance liquid chromatography (HPLC). RNA was extracted from safflower florets treated with 100 μmol·L-1 MeJA for 6 h. The transcription of key genes involved in the biosynthesis of HSYA was quantified by qRT-PCR and differentially expressed genes were identified. The content of HSYA increased after treatment with MeJA, with 100 μmol·L-1 MeJA treatment for 6 h having the greatest effect on HSYA accumulation. qRT-PCR results showed that MeJA could significantly increase the transcription of HSYA biosynthesis genes including PAL2, PAL4, 4CL2, 4CL4, 4CL5, CHS3, CHS4 and CHI2. The content of HSYA differed between safflowers of different colors with a trend of red > orange-yellow > yellow > white. The results of qRT-PCR showed that the expression of CHS1 and CHI2 in red, orange and yellow safflower was significantly higher than that in white safflower. These results indicate that MeJA promotes the accumulation of HSYA by up-regulating the expression of genes involved in the biosynthesis of HSYA such as PAL2, PAL4, 4CL2, 4CL4, 4CL5, CHS3, CHS4 and CHI2, and the variation of HSYA content in safflower of different colors was related to a difference in the level of expression of CHS1 and CHI2.

safflower  /  methyl jasmonate  /  hydroxysafflor yellow A  /  expression difference
谭政委, 李磊, 杨红旗, 余永亮, 许兰杰, 董薇, 夏伟, 马新明, 梁慧珍. 茉莉酸甲酯诱导和不同花色红花中羟基红花黄色素A积累差异的机制分析. 药学学报, 2020 , 55 (12) : 2982 -2988 . DOI: 10.16438/j.0513-4870.2020-0705
Zheng-wei TAN, Lei LI, Hong-qi YANG, Yong-liang YU, Lan-jie XU, Wei DONG, Wei XIA, Xin-ming MA, Hui-zhen LIANG. The mechanism of methyl jasmonate-induced accumulation of hydroxysafflor yellow A in safflower of different colors[J]. Acta Pharmaceutica Sinica, 2020 , 55 (12) : 2982 -2988 . DOI: 10.16438/j.0513-4870.2020-0705
正文
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红花(Carthamus tinctorius L.), 别名红蓝花、刺红花、草红花, 是菊科红花属一年生双子叶草本植物, 红花作为红花属中唯一的栽培种, 是集油用、药用和工业用于一体的特种经济作物[1, 2]。据《本草纲目》记载, 红花具有“活血、润燥、止痛、散肿通经”的作用。据统计, 《中国药典》2015年版收载含红花的中成药近百种, 中国药典2015版规定, 作为中药材使用的红花干燥花丝中HSYA含量应不低于1.0%[3], 因此, 其HSYA含量高低直接决定了红花花丝中药材的等级。药理学研究结果表明HSYA是红花中最主要的活性成分, 它在治疗心脑血管疾病、糖尿病、抗血栓、抗肿瘤、抗炎、护肝等方面都有很好的作用[4, 5]。
HSYA和其他类黄酮化合物一样也是通过苯丙氨酸代谢途径合成, 苯丙氨酸在苯丙氨酸解氨酶(phenylalanine ammonialyase, PAL)、肉桂酸-4-羟化酶(cinnamate 4-hydroxylase, C4H)、4-香豆酰辅酶A连接酶(4-coumaryol CoA ligase, 4CL)催化下生成香豆酰辅酶A, 再与3分子的丙二酸单酰辅酶A在查尔酮合酶(chalcone synthase, CHS)的作用下生成查尔酮, 一部分查尔酮在细胞色素P450氧化酶(cytochrome P450, CYP450)和C-糖基转移酶(C-glycosyltransferases, CGTs)的催化下生成HSYA和其他类醌式查尔酮类化合物, 一部分查尔酮在查尔酮异构酶(chalcone isomerase, CHI)的作用下生成柚皮素, 然后在一系列酶的催化下生成黄酮、黄酮醇、花青素等其他黄酮类化合物[6-8] (图 1)。CHS被认为是合成红花黄酮类化合物和HSYA的关键酶, 也是其限速酶, 但也有研究表明PAL、C4H、4CL的表达量高低也与类黄酮的积累有很大的相关性[9, 10], 但到目前为止, 对红花中黄酮类化合物的生物合成及调控的研究仍不够深入。
茉莉酸甲酯(methyl jasmonate, MeJA)作为一种植物内源激素和信号分子, 在植物的生长、发育、免疫调控、次生代谢、生物胁迫和非生物胁迫等方面都发挥着重要作用[11-14]。大量研究发现, MeJA通过激活或抑制相应转录因子的活性, 调控植物类黄酮[15]、二萜[16]、三萜[17]等多种植物次生代谢的关键酶的基因表达, 从而参与植物次生代谢的生物合成过程调控。有研究表明, MeJA能够诱导HSYA积累[18], 另外, 有研究发现不同花色红花花丝中HSYA含量不同[19], 推测MeJA对HSYA的诱导积累和不同花色红花中HSYA含量差异与HSYA生物合成相关基因的表达差异有关, 但具体是哪些(个)基因的相关研究还不够深入。本研究用豫红花1号作为MeJA处理材料, 对处理后的HSYA含量、相关合成基因进行定量分析, 同时对不同花色红花中的HSYA生物合成关键基因的表达量进行定量分析, 以期为MeJA调控HSYA生物合成及不同花色红花中HSYA含量差异的分子机制提供理论参考。
材料和方法
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材料  研究材料“豫红花1号”及管状花色颜色为白色(W)、黄色(Y)、橙黄色(OY)、红色(R)红花品系种植于河南农业科学院现代农业研究开发基地。茉莉酸甲酯(MeJA)购自上海生工有限公司, HSYA标准品购自中国食品药品检定研究院, 引物为河南尚亚生物技术有限公司合成。
样本处理   以“豫红花1号”作为MeJA处理材料, 为了减小实验误差, 选取在大田中开花第一天的顶果球红花花冠连同茎杆一起剪下, 放入1/2浓度的Hoagland标准营养液中, 移至光照培养箱中培养, 生长条件为(26 ± 1) ℃、16 h/8 h (光照/黑暗), 24 h后进行MeJA处理, 分别用0、50、100、200 μmol·L-1 MeJA喷施红花花序, 对照组用MeJA溶剂喷施, 每个处理设置3个生物学重复, 在处理后的0、3、6、12、24 h取样, 液氮速冻保存, 用于RNA提取和HSYA含量测定。
不同花色红花品系的取样   取不同花色红花品系开花第三天的花冠用于RNA提取和HSYA含量测定, 为了减小实验误差, 每个花色品系选取5~10株上的花序均匀混合, 取3个生物学重复。
HSYA标准曲线制定   称HSYA标准品10.0 mg, 加25%甲醇溶解并于10 mL容量瓶中定容, 经0.45 μm微孔滤膜过滤, 取滤液供HPLC测定使用。分别吸取HSYA对照品溶液0.25、0.50、0.75、1.0、2.0、4.0 mL置于10 mL容量瓶中, 加25%甲醇定容至刻度, 混匀, 取9 μL进样。色谱条件参考杨玉霞等[13]方法, 并稍作修改:色谱柱: Agilent C18柱(4.6 mm×250 mm, 5 μm); 检测波长: 403 nm; 流动相:甲醇-乙腈-0.7%的磷酸水溶液(26:2:72);流速: 1.0 mL·min-1; 柱温: 30 ℃; 进样量: 10 μL。
样本中HSYA含量测定  精密称取红花样品粉末100 mg, 精密量取25%甲醇10 mL, 称量, 25 ℃超声(300 W, 50 kHz)处理40 min, 取出后称重并补足失重。用0.45 μm滤膜过滤得供试品溶液。Agilent Zorbax Eclipse XDB-C18色谱柱(4.6 mm ×250 mm, 5 μm); 以甲醇-乙腈-0.7%磷酸溶液(26:2:72)为流动相等度洗脱; 检测波长403 nm; 流速1.0 mL·min-1; 柱温30 ℃; 进样量10 μL。
样品总RNA提取及逆转录反应   按照华越洋生物公司的Quick RNA Isolation Kit试剂盒说明书提取样品的总RNA; 按照天根生物科技有限公司的Fast Quant RT Super Mix说明书进行逆转录反应。
Real-time PCR分析  采用Eppendorf Mastercycler ep Realplex 2.2 Detection System进行实时荧光定量PCR, 根据红花转录组数据[20]和参考文献[21]设计PAL1、PAL2、PAL3、PAL4、PAL5、C4H1、4CL1、4CL2、4CL3、4CL4、4CL5、CHS1、CHS2、CHS3、CHS4、CHS5、CHI1、CHI2基因的特异引物, 以Ct60s作为内参基因, 引物序列如表 1所示。用天根生物科技公司的Super Real Pre Mix (SYBR Green)试剂盒进行荧光定量PCR反应, 扩增体系20 μL: 2×SuperReal Pre Mix 10 μL、上游引物(10 μmol·L-1) 0.5 μL、下游引物(10 μmol·L-1) 0.5 μL、cDNA模板5 μL、ddH2O 3 μL。荧光定量PCR条件设置反应程序: 95 ℃变性20 s、55 ℃退火20 s、72 ℃延伸20 s扩增45个循环。
数据分析   用2-ΔΔCT相对定量方法计算出各基因的相对表达量, 用标准曲线法对样品的HSYA含量进行定量分析。用单因素方差对样品的基因表达和HSYA含量进行统计分析, 结果以平均值±标准差的形式表示, P < 0.05表示差异具有显著水平。
结果与分析
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1 适度的MeJA处理能够促进红花管状花中HSYA的积累
为了确定MeJA对HSYA的积累是否有影响及最适处理浓度, 采用不同浓度(0、50、100、200 μmol·L-1)对红花进行喷施处理, 为了减少在自然环境中外界因素如光照、温度、风力等变化对激素处理的影响, 选取开花一致的红花花序, 连同茎秆一同剪下后置于营养液中稳定培养24 h后进行MeJA喷施处理, 在处理3 h后, 通过液相色谱对红花管状花中的HSYA含量进行定量分析, 结果显示MeJA处理能够显著增加HSYA的积累, 其中100 μmol·L-1的MeJA处理对HSYA促进效果最为明显(图 2A), 所以, 在后续的实验中选取该浓度MeJA对红花进行处理。
为了进一步验证适度的MeJA处理能够提高红花HSYA的含量, 用100 μmol·L-1的MeJA对红花进行了不同时间处理, 结果表明, 随着不同时长处理, 红花HSYA的积累呈现升高的趋势, 其中处理6 h后, HSYA的含量达到高峰(图 2B), 这一结果进一步证明了适度的MeJA处理能够促进红花中HSYA的积累。
2 MeJA处理对HSYA合成相关基因的影响
通过HSYA的结构推测其合成的最初底物是查尔酮, 为了进一步研究MeJA处理对红花HSYA积累的分子机制, 通过红花转录组数据及文献检索的方式, 找到了可能参与红花HSYA合成的相关基因, 以MeJA处理6 h的红花管状花为材料提取RNA, 并以喷施MeJA溶剂的红花为对照材料, 通过荧光定量PCR对参与查尔酮合成前后的18个基因进行定量分析。定量分析结果(图 3)表明, 与对照组(mock)相比, MeJA处理组中大部分基因的表达量都有所升高, 其中PAL2、PAL4、4CL2、4CL4、4CL5、CHS3、CHS4和CHI2的表达量显著升高。其中CHS是参与植物查尔酮合成的最为关键酶, 被认为是植物类黄酮合成途径的第一个酶, 而HSYA的合成是以查尔酮为底物, 因此, 推测MeJA主要通过提高PAL2、PAL4、4CL2、4CL4、4CL5、CHS3、CHS4、CHI2的转录水平参与调控HSYA的生物合成过程, 其中CHS3、CHS4的表达量升高可能是MeJA诱导HSYA积累的主要因素。
3 不同花色红花品种中HSYA含量的分析
研究室前期在红花种质资源收集与评价的研究过程中, 发现大量管状花花色变异的种质资源, 根据红花管状花颜色可以将红花种质资源大体分为4类, 分别为白色(W)、黄色(Y)、橙黄色(OY)和红色(R) (图 4), 对不同花色红花品系的HSYA含量进行测定, 结果表明, 白色红花中未检测到HSYA, 黄色(Y)、橙黄色(OY)和红色(R)中HSYA含量较高, 并呈现R > OY > Y > W的趋势, 但OY、Y和W中HSYA的含量差异并不明显(图 5)。
4 不同花色红花品种中HSYA合成关键基因表达分析
为了研究在不同红花品系中HSYA积累差异的分子机制, 对不同花色红花中HSYA合成的关键基因进行了定量分析, 荧光定量PCR分析结果显示, PAL3、PAL4、CL5、CHS1、CHS3和CHI2在红色、橙黄色和黄色红花品系中明显高于白色红花品系, 其中CHS1和CHI2的表达量达到显著水平(图 6), 但PAL3、PAL4、CL5、CHS1、CHS3和CHI2的基因表达量在红色、橙黄色和黄色红花品系中表达差异不明显。
讨论
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红花(Carthamus tinctorius L)是一种重要的药用植物, 其花丝和种子是传统中药材, 广泛应用于预防和治疗心脑血管疾病、骨质疏松症及妇科等疾病, 目前, 从红花中分离的化学成分超过100种, 主要包括类黄酮、生物碱、有机酸等[22], 其中醌式查尔酮糖苷类化合物是红花中特有的化学成分, 也构成红花红色素和黄色素的主要成分, 这些化合物具有很强的药理学活性, 其中HSYA是药理学活性最强的成分, 临床上广泛应用于治疗心、脑血管疾病的红花黄色素注射液的主要成分之一就是HSYA[23]。因此HSYA含量的高低直接决定了中药材红花花丝的等级标准, 如何提高红花HSYA含量也是红花遗传育种改良的主要目标之一, 但目前关于红花HSYA生物及合成调控的研究相对较少。
MeJA作为一种植物内源性激素, 在植物的次生代谢调控中起重要作用, MeJA广泛应用于悬浮培养细胞中诱导产生目标次生代谢产物, 如雷公藤素[24]、丹参酚酸类化合物[14]等。在本研究中, 适度外施MeJA处理红花花冠, 可以提高红花HSYA含量, 另有研究表明, MeJA处理红花可促使红花中芦丁、山柰酚-3-O-β-芸香糖苷、红花红色素等类黄酮化合物的积累[18], 这说明MeJA作为外源诱导子有利于红花类黄酮的积累。本研究为了更加深入地了解MeJA对HSYA的调控, 综合分析了18个与HSYA合成相关的基因, 结果显示, MeJA处理使PAL2、PAL4、4CL2、4CL4、4CL5、CHS3、CHS4和CHI2的转录水平明显调高, 植物中CHS家族一般包含多个基因, 这些基因通过不同的调控方式参与植物抗逆、生长发育等生理过程, Guo等[21]通过花粉管导入法在红花中过表达CHS1基因可以显著增加红花醌式查尔酮类化合物的含量, 本研究中发现MeJA处理后CHS3和CHS4的转录水平和HSYA含量都明显升高, MeJA对CHS1基因表达只有微弱的诱导作用, 这说明CHS3和CHS4也参与HSYA生物合成, 提示MeJA对HSYA的诱导作用主要通过CHS3和CHS4基因实现。
另外, 在红花种质资源的收集鉴定过程中, 发现大量花色变异的红花种质资源, 定量分析发现不同花色红花品系的HSYA含量有一定差异性, 红色、橙黄色和黄色红花品系中HSYA含量明显高于白色红花, 本研究通过对HSYA合成的关键基因的表达进行定量分析, 初步找到了影响HSYA在不同花色红花中含量差异的潜在候选基因, 与MeJA诱导PAL2、PAL4、4CL2、4CL4、4CL5、CHS3、CHS4和CHI2等基因的转录水平提高从而使HSYA含量升高的机制不同, CHS1和CHI2的转录水平在不同花色红花中存在明显差异, 红色、橙黄色和黄色红花品系中CHS1和CHI2基因的表达水平明显高于白色, 因为CHS催化产物是HSYA合成的直接底物, 所以推断CHS1在不同花色红花中的表达差异是不同花色红花HSYA含量差异的因素之一。但qRT-PCR结果显示CHS2、CHS3、CHS4和CHS5在白色红花品系中也有一定的表达量, 但白色红花中检测不到HSYA, 说明其他参与HSYA生物合成的下游基因如CGT、P450在白色红花中存在功能或表达缺失。
综上所述, 外源MeJA刺激可提高HSYA含量及该合成途径中关键基因的表达, 为今后进一步研究外源诱导子影响HSYA次生代谢途径机制提供了依据, 不同花色红花HSYA含量差异的机制探讨, 为红花品种的遗传改良提供了一定参考。
作者贡献:谭政委和梁慧珍设计了本实验, 李磊和杨红旗完成实验材料种植、采样和处理, 余永亮、许兰杰、董薇、夏伟完成了样品HSYA定量测定、基因的定量分析, 谭政委和李磊完成了数据分析、论文初稿的写作, 梁慧珍和马新明指导论文写作与修改。
利益冲突:本文的研究内容无任何利益冲突。
基金
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  • 现代农业产业技术体系建设专项资金资助(CARS-21)
  • 河南省农科院创新创意项目(2020CX03)
  • 河南省农科院优秀青年基金项目(2020YQ05)
  • 河南省农科院优秀青年基金项目(2020YQ28)
  • 河南省博士后基金项目(001803053)
  • 国家农业科研杰出人才及其创新团队“特种油料作物品质改良创新团队”(农财发(2016)45号)
文献
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2020年第55卷第12期
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doi: 10.16438/j.0513-4870.2020-0705
  • 接收时间：2020-05-09
  • 首发时间：2026-01-23
  • 出版时间：2020-12-12
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  • 收稿日期：2020-05-09
  • 修回日期：2020-06-04
基金
现代农业产业技术体系建设专项资金资助(CARS-21)
河南省农科院创新创意项目(2020CX03)
河南省农科院优秀青年基金项目(2020YQ05)
河南省农科院优秀青年基金项目(2020YQ28)
河南省博士后基金项目(001803053)
国家农业科研杰出人才及其创新团队“特种油料作物品质改良创新团队”(农财发(2016)45号)
作者信息
    1.河南省农业科学院芝麻研究中心, 河南 郑州 450002
    2.河南农业大学信息与管理科学学院, 河南 郑州 450002

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*梁慧珍, Tel:86-371-65738565, 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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