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Article(id=1218551200014848696, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1218551189596197749, articleNumber=null, orderNo=null, doi=10.16438/j.0513-4870.2017-0614, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1498406400000, receivedDateStr=2017-06-26, revisedDate=1504540800000, revisedDateStr=2017-09-05, acceptedDate=null, acceptedDateStr=null, onlineDate=1768454845670, onlineDateStr=2026-01-15, pubDate=1515686400000, pubDateStr=2018-01-12, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1768454845670, onlineIssueDateStr=2026-01-15, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1768454845670, creator=13701087609, updateTime=1768454845670, updator=13701087609, issue=Issue{id=1218551189596197749, tenantId=1146029695717560320, journalId=1189982191388893191, year='2018', volume='53', issue='1', pageStart='1', pageEnd='162', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1768454843186, creator=13701087609, updateTime=1768456905168, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1218559838225879190, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1218551189596197749, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1218559838225879191, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1218551189596197749, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=141, endPage=146, ext={EN=ArticleExt(id=1218551200576885487, articleId=1218551200014848696, tenantId=1146029695717560320, journalId=1189982191388893191, language=EN, title=Cloning and expression analysis of MYB transcription factor genes in safflower, columnId=1190335348761793317, journalTitle=Acta Pharmaceutica Sinica, columnName=ORIGINAL ARTICLES, runingTitle=null, highlight=null, articleAbstract=

Safflower is a dried flower of the annual herbaceous plant safflower (Carthamus tinctorius L.). As a traditional Chinese medicine, it was widely used in the regulation of blood circulation. Flavonoids are the main active ingredients in safflower. MYB transcription factors are involved in the regulation of flavonoids. The cloning and expression analysis of MYB transcription factor genes in safflower is of great significance, not only for clarifying the regulation mechanism of flavonoids biosynthesis in safflower, but also for the artificial regulation of flavonoid biosynthesis in safflower. Based on the transcriptome data, we used iTAK to annotate the MYB transcription factors in safflower. The MYB transcription factors were cloned and their sequences were analyzed. Besides, their expressions were analyzed by a Real-time PCR. In the experiment, eight long fragment MYB transcription factors were screened and six MYB transcription factors was successfully cloned, named CtMYB-TF1, CtMYB-TF2, CtMYB-TF4, CtMYB-TF5, CtMYB-TF6 and CtMYB-TF7, respectively. The six MYB transcription factors had the core domain of MYB transcription factor family, and evolutionary analysis showed that the CtMYB-TF7 transcription factor was closely related to the factors AtMYBL2 and AtMYB12. Expression analysis showed that the expression of CtMYB-TF5, CtMYB-TF6 and CtMYB-TF7 was low in roots, stems and leaves, and was high in the flower. The results provide a foundation for study of mechanism of molecular regulation of safflower flavonoids.

, correspAuthors=Jin PEI, 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=Jiang CHEN, Xiao-hui TANG, Chao-xiang REN, Xiao CHEN, Wen HE, Si-yuan ZHANG, Qing-hua WU, Jin PEI), CN=ArticleExt(id=1218551202342687683, articleId=1218551200014848696, tenantId=1146029695717560320, journalId=1189982191388893191, language=CN, title=红花MYB转录因子基因克隆及表达分析, columnId=1190335348896011050, journalTitle=药学学报, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=

红花是菊科一年生草本植物红花（Carthamus tinctorius L.）的干燥花，为活血化瘀常用中药。黄酮类成分是红花主要有效成分，MYB转录因子广泛参与调控黄酮类成分的合成，对红花MYB转录因子克隆及表达分析为解析红花黄酮类成分的调控机制、调控黄酮类成分的合成具有重要意义。本研究基于在线二代转录组数据，首先利用iTAK软件对MYB转录因子进行注释，设计引物对长片段MYB转录因子基因进行克隆，其次对克隆到的MYB转录因子基因进行序列分析，再次利用实时荧光定量PCR对克隆的MYB转录因子基因表达进行分析。注释筛选得到8个长片段MYB转录因子基因，成功克隆到6个MYB转录因子基因，分别命名为CtMYB-TF1、CtMYB-TF2、CtMYB-TF4、CtMYB-TF5、CtMYB-TF6及CtMYB-TF7。序列分析表明，克隆到的6个MYB转录因子基因都具有MYB转录因子核心结构域，其中CtMYB-TF7转录因子同已报道黄酮类成分合成调控因子AtMYBL2及AtMYB12关系较近。表达分析表明，CtMYB-TF5、CtMYB-TF6及CtMYB-TF7在根、茎及叶中表达量低，在花中表达量高。研究结果为进一步研究红花黄酮类成分的分子调控奠定基础。

, correspAuthors=裴瑾, authorNote=null, correspAuthorsNote=
* 裴瑾, Tel/Fax:86-26-61800235, E-mail:
, copyrightStatement=版权所有©《药学学报》编辑部2018, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=VjyFj+hxgm3E1IZBRsWcRQ==, magXml=sMviK4OhM0heHPIJEGrVbA==, pdfUrl=null, pdf=miKRIQL4hHO3WN6ThEBk+A==, pdfFileSize=287379, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=uTadrTWjH66eew0EuB142g==, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=Y9PXuStjAT+zLktJxDCstg==, mapNumber=null, authorCompany=null, fund=null, authors=null, authorsList=陈江, 唐小慧, 任超翔, 陈骁, 何雯, 张思源, 吴清华, 裴瑾)}, authors=[Author(id=1218970735113061081, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551200014848696, 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=1218970735255667434, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1218551200014848696, authorId=1218970735113061081, language=EN, stringName=Jiang CHEN, firstName=Jiang, middleName=null, lastName=CHEN, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=1, 2, address=1. State Key Laboratory Breeding Base of Systematic Research, Development and Utilization of Chinese Medicine Resources, Chengdu 611137, China
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Construction of cDNA mixed template. A: The three periods (Ⅰ, Ⅱ and Ⅲ) for flower development; B: RNA electrophoresis for the three periods, M represents 2 000 bp DNA Marker

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Gene cloning of MYB transcription factors. 1-8 represent CtMYB-TF1, CtMYB-TF2, CtMYB-TF3, CtMYB-TF4, CtMYB-TF5, CtMYB-TF6, CtMYB-TF7 and CtMYB-TF8. M represents 2 000 bp DNA marker

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Protein domain analysis of MYB transcription factors. The gray frame indicates the full length of the corresponding MYB protein, the length is indicated by a number, and AA represent amino acids. The blue box represents the MYB DNA-binding domain

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Evolution analysis of MYB transcription factors. The gene name is shown in the figure, and the parentheses represent the NCBI accession number

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Tissue expression analysis of MYB transcription factor genes. The expression level of each gene is indicated by different grid lines, R represents root, S represents stem, L represents leaf, F1-F3 represents the three periods (Ⅰ, Ⅱ and Ⅲ)

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Gene name Primer sequence
CtMYB-TFTF1 F: TTTTCCGATGAACTCCAC
R: CCCAAACCCTAAATCTTTT
CtMYB-TFTF2 F: TTTTACAATCGCTGGTCC
R: AATCCCTCACTTCGTTCTT
CtMYB-TFTF3 F: TTTATCCCAACAACTCTACTTC
R: CCAATACCAGGCTCTTTCT
CtMYB-TFTF4 F: GAAGACAGGAGGCTTGAA
R: CACGATACGCATACCACA
CtMYB-TFTF5 F: GGCAAATTCGACGGATAA
R: GGCTGGATAACTTGGGTT
CtMYB-TFTF6 F: GCTGCGATCTAATACCAT
R: TTCACTAAAGTCTAACACCC
CtMYB-TFTF7 F: GACCTATCAGACGAGCAA
R: AACAGCCAACAGTTACGA
CtMYB-TFTF8 F: CATTTCCTTTCCTTTCTTTC
R: TTCATCTGCTTATTACTCCAA
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Primer sequences for MYB transcription factor gene cloning

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Gene name Primer sequence
CtMYB-TFTF1 F: TTTTCCGATGAACTCCAC
R: CCCAAACCCTAAATCTTTT
CtMYB-TFTF2 F: TTTTACAATCGCTGGTCC
R: AATCCCTCACTTCGTTCTT
CtMYB-TFTF3 F: TTTATCCCAACAACTCTACTTC
R: CCAATACCAGGCTCTTTCT
CtMYB-TFTF4 F: GAAGACAGGAGGCTTGAA
R: CACGATACGCATACCACA
CtMYB-TFTF5 F: GGCAAATTCGACGGATAA
R: GGCTGGATAACTTGGGTT
CtMYB-TFTF6 F: GCTGCGATCTAATACCAT
R: TTCACTAAAGTCTAACACCC
CtMYB-TFTF7 F: GACCTATCAGACGAGCAA
R: AACAGCCAACAGTTACGA
CtMYB-TFTF8 F: CATTTCCTTTCCTTTCTTTC
R: TTCATCTGCTTATTACTCCAA
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Gene name Primer sequence
CtMYB-TF1 QF:CTTCTTTGAGCCTATCCTTACC
QR:CTAATGGCACCTTCACCCT
CtMYB-TF2 QF:CGGAGGAGGACGAGAAGAT
QR:CGGCTGGTAGTTGTTGTTGA
CtMYB-TF4 QF:CTGTCGTCTGAGGTGGTGT
QR:AGGTTTAGTTTGAGCGTGTT
CtMYB-TF5 QF:ATTTGGCAACAGATGGAC
QR:GCAATAATCTTCGGGTCA
CtMYB-TF6 QF:GTCGTCTACCCAAACTGC
QR:AAGGAGGAGAACCAAGGA
CtMYB-TF7 QF:CGATACTCCAGAGCCGTTGA
QR:ACTTTCCGCACTCCCACA
Ct60S QF:CATCCATTATCCAACAATC
QR:AAGAGTAATCAGTCTCCA
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Primer sequence for Real-time PCR

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Gene name Primer sequence
CtMYB-TF1 QF:CTTCTTTGAGCCTATCCTTACC
QR:CTAATGGCACCTTCACCCT
CtMYB-TF2 QF:CGGAGGAGGACGAGAAGAT
QR:CGGCTGGTAGTTGTTGTTGA
CtMYB-TF4 QF:CTGTCGTCTGAGGTGGTGT
QR:AGGTTTAGTTTGAGCGTGTT
CtMYB-TF5 QF:ATTTGGCAACAGATGGAC
QR:GCAATAATCTTCGGGTCA
CtMYB-TF6 QF:GTCGTCTACCCAAACTGC
QR:AAGGAGGAGAACCAAGGA
CtMYB-TF7 QF:CGATACTCCAGAGCCGTTGA
QR:ACTTTCCGCACTCCCACA
Ct60S QF:CATCCATTATCCAACAATC
QR:AAGAGTAATCAGTCTCCA
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Gene name CDS length Amino acid Molecular weight Isoelectric point Average hydrophobicity GO annotation Pfam annotation
CtMYB-TF1 903 300 33 082.23 7.66 -0.684 DNA binding Myb-like protein
CtMYB-TF2 459 152 17 706.31 11.77 -0.804 DNA binding Myb-like protein
CtMYB-TF4 1 131 376 40 719.99 5.96 -0.653 DNA binding Myb-like protein
CtMYB-TF5 804 267 30 064.59 9.35 -0.997 DNA binding Myb-like protein
CtMYB-TF6 723 240 26 319.73 5.53 -0.414 DNA binding Myb-like protein
CtMYB-TF7 1 233 410 44 439.50 8.55 -0.736 DNA binding Myb-like protein
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Bioinformatics analysis of MYB transcription factor genes

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Gene name CDS length Amino acid Molecular weight Isoelectric point Average hydrophobicity GO annotation Pfam annotation
CtMYB-TF1 903 300 33 082.23 7.66 -0.684 DNA binding Myb-like protein
CtMYB-TF2 459 152 17 706.31 11.77 -0.804 DNA binding Myb-like protein
CtMYB-TF4 1 131 376 40 719.99 5.96 -0.653 DNA binding Myb-like protein
CtMYB-TF5 804 267 30 064.59 9.35 -0.997 DNA binding Myb-like protein
CtMYB-TF6 723 240 26 319.73 5.53 -0.414 DNA binding Myb-like protein
CtMYB-TF7 1 233 410 44 439.50 8.55 -0.736 DNA binding Myb-like protein
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红花MYB转录因子基因克隆及表达分析
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陈江 1, 2 , 唐小慧 1, 2 , 任超翔 1, 2 , 陈骁 2 , 何雯 2 , 张思源 2 , 吴清华 1, 2 , 裴瑾 1, 2, *
药学学报 | 研究论文 2018,53(1): 141-146
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药学学报 | 研究论文 2018, 53(1): 141-146
红花MYB转录因子基因克隆及表达分析
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陈江1, 2, 唐小慧1, 2, 任超翔1, 2, 陈骁2, 何雯2, 张思源2, 吴清华1, 2, 裴瑾1, 2, *
作者信息
  • 1.中药资源系统研究与开发利用国家重点实验室培育基地, 四川 成都 611137
  • 2.成都中医药大学药学院, 四川 成都 611137

通讯作者:

* 裴瑾, Tel/Fax:86-26-61800235, E-mail:
Cloning and expression analysis of MYB transcription factor genes in safflower
Jiang CHEN1, 2, Xiao-hui TANG1, 2, Chao-xiang REN1, 2, Xiao CHEN2, Wen HE2, Si-yuan ZHANG2, Qing-hua WU1, 2, Jin PEI1, 2, *
Affiliations
  • 1. State Key Laboratory Breeding Base of Systematic Research, Development and Utilization of Chinese Medicine Resources, Chengdu 611137, China
  • 2. Pharmacy College, Chengdu University of Traditional Chinese Medicine, Chengdu 611137, China
出版时间: 2018-01-12 doi: 10.16438/j.0513-4870.2017-0614
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摘要
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红花是菊科一年生草本植物红花（Carthamus tinctorius L.）的干燥花，为活血化瘀常用中药。黄酮类成分是红花主要有效成分，MYB转录因子广泛参与调控黄酮类成分的合成，对红花MYB转录因子克隆及表达分析为解析红花黄酮类成分的调控机制、调控黄酮类成分的合成具有重要意义。本研究基于在线二代转录组数据，首先利用iTAK软件对MYB转录因子进行注释，设计引物对长片段MYB转录因子基因进行克隆，其次对克隆到的MYB转录因子基因进行序列分析，再次利用实时荧光定量PCR对克隆的MYB转录因子基因表达进行分析。注释筛选得到8个长片段MYB转录因子基因，成功克隆到6个MYB转录因子基因，分别命名为CtMYB-TF1、CtMYB-TF2、CtMYB-TF4、CtMYB-TF5、CtMYB-TF6及CtMYB-TF7。序列分析表明，克隆到的6个MYB转录因子基因都具有MYB转录因子核心结构域，其中CtMYB-TF7转录因子同已报道黄酮类成分合成调控因子AtMYBL2及AtMYB12关系较近。表达分析表明，CtMYB-TF5、CtMYB-TF6及CtMYB-TF7在根、茎及叶中表达量低，在花中表达量高。研究结果为进一步研究红花黄酮类成分的分子调控奠定基础。

红花  /  MYB转录因子  /  基因克隆  /  表达分析  /  黄酮类成分

Safflower is a dried flower of the annual herbaceous plant safflower (Carthamus tinctorius L.). As a traditional Chinese medicine, it was widely used in the regulation of blood circulation. Flavonoids are the main active ingredients in safflower. MYB transcription factors are involved in the regulation of flavonoids. The cloning and expression analysis of MYB transcription factor genes in safflower is of great significance, not only for clarifying the regulation mechanism of flavonoids biosynthesis in safflower, but also for the artificial regulation of flavonoid biosynthesis in safflower. Based on the transcriptome data, we used iTAK to annotate the MYB transcription factors in safflower. The MYB transcription factors were cloned and their sequences were analyzed. Besides, their expressions were analyzed by a Real-time PCR. In the experiment, eight long fragment MYB transcription factors were screened and six MYB transcription factors was successfully cloned, named CtMYB-TF1, CtMYB-TF2, CtMYB-TF4, CtMYB-TF5, CtMYB-TF6 and CtMYB-TF7, respectively. The six MYB transcription factors had the core domain of MYB transcription factor family, and evolutionary analysis showed that the CtMYB-TF7 transcription factor was closely related to the factors AtMYBL2 and AtMYB12. Expression analysis showed that the expression of CtMYB-TF5, CtMYB-TF6 and CtMYB-TF7 was low in roots, stems and leaves, and was high in the flower. The results provide a foundation for study of mechanism of molecular regulation of safflower flavonoids.

safflower  /  MYB transcription factor  /  gene cloning  /  expression analysis  /  flavonoids
陈江, 唐小慧, 任超翔, 陈骁, 何雯, 张思源, 吴清华, 裴瑾. 红花MYB转录因子基因克隆及表达分析. 药学学报, 2018 , 53 (1) : 141 -146 . DOI: 10.16438/j.0513-4870.2017-0614
Jiang CHEN, Xiao-hui TANG, Chao-xiang REN, Xiao CHEN, Wen HE, Si-yuan ZHANG, Qing-hua WU, Jin PEI. Cloning and expression analysis of MYB transcription factor genes in safflower[J]. Acta Pharmaceutica Sinica, 2018 , 53 (1) : 141 -146 . DOI: 10.16438/j.0513-4870.2017-0614
正文
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红花是菊科一年生草本植物红花(Carthamus tinctorius L.)的干燥花, 自东汉时期张仲景《金匮要略》中“妇人杂病篇”就有使用记载, 其质柔软, 味辛, 性温, 归心、肝经, 具有活血通经、祛瘀止痛的功效, 为活血化瘀常用中药。历史上红花以四川“川红花”、河南“怀红花”最为出名[1]。当前红花在全国各地栽种范围较广, 新疆、云南、四川及河南等地均有种植, 特别是在新疆, 种植面积较大。
红花的化学成分按照结构包括黄酮、生物碱、聚炔、亚精胺、木脂素、倍半萜、有机酸、留醇和烷基二醇等类型。其中黄酮类成分, 包括羟基红花黄色素A (hydroxysafflor yellow A, HSYA)、槲皮素(quercetin)、柚皮素(naringenin)等是红花主要有效成分。研究表明HSYA具有抗凝、抗缺氧、降血压、改善心脑血管供血不足等作用[2, 3], 山奈酚[4]、槲皮素[5]等在活血化瘀上也有较好效果。
MYB转录因子广泛参与调控黄酮类成分的合成, 国内外已有多篇文章进行了综述[6-8]。在模式植物拟南芥中, AtMYB12能调控查尔酮合成酶基因(CHS)、查尔酮异构酶基因(CHI)及黄酮醇羟化酶基因(F3H)等基因表达, 促进调控黄酮类成分的合成[9]。其同源基因AtMYB11和AtMYB111也能调控黄酮类成分的合成[10]。AtMYBL2也被报道参与调控黄酮类成分的合成[11]。而在红花中, 克隆报道的MYB转录因子较少。至今仅见Guan等[12]克隆的CtMYB1一篇报道。
近年来, 为了深入研究红花黄酮类成分的合成调控, 对红花进行了大量的转录组测序工作[13-15]。本研究基于在线二代转录组数据, 利用iTAK软件对MYB转录因子基因进行注释, 设计引物对长片段MYB转录因子基因进行克隆, 其次对克隆到的MYB转录因子基因进行序列分析, 再次利用实时荧光定量PCR (real-time PCR)对克隆的MYB转录因子基因表达进行分析。通过对红花MYB转录因子基因克隆及表达分析, 为进一步解析红花黄酮类成分的调控机制、调控黄酮类成分的合成奠定基础。
材料与方法
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材料  红花为道地产区四川简阳地方品种, 经成都中医药大学药学院资源与鉴定系严铸云教授鉴定为红花(Carthamus tinctorius L.)。2016年3月底将红花种子种植于成都中医药大学温江校区药用植物园。红花3个时期的管状花混样用于MYB转录因子基因克隆(3个时期见图 1A)。红花根、茎、叶及3个时期的管状花用于MYB转录因子基因表达分析, 根、茎、叶的取样时间是在红花开花第一个时期。田间取样后立即液氮冷冻, 储存于-80 ℃冰箱备用。克隆用大肠杆菌DH5a菌株购自宝生物(TaKaRa, Dalian, China)。
仪器与试剂  主要仪器有PCR仪T100TM Thermal Cycler型(Bio-Rad, CA, USA), 高速低温离心机CT15RE型(HITACHI, Japan), 凝胶成像系统Gel DocTM XR+型(Bio-Rad, CA, USA), 实时荧光定量PCR仪Bio-Rad CFX96型(Bio-Rad, CA, USA); 主要试剂有2×TransTaq High Fidelity (HiFi) PCR SuperMix Ⅱ (Transgen, Beijing, China), 琼脂糖凝胶回收试剂盒、质粒提取试剂盒(Tiangen, Beijing, China), TRIzol (Invertrigen, MA, USA)及TRIzol伴侣试剂盒(Tiandz, Beijing, China), 感受态细胞制备试剂盒(TaKaRa, Dalian, China), 克隆载体pMD19-T (TaKaRa, Dalian, China), 引物合成及片段测序送擎科公司(Tsingke, Beijing, China)。
序列获取及引物设计  本研究采用的转录组数据主要参考Huang等[13]发表的在线数据, NCBI登录号SRA048496, 使用百迈克云平台(https://www.biocloud.net/), 选择无参模型进行分析, 利用iTAK软件[16]对调控因子进行注释, 筛选序列较长MYB转录因子基因, 以此序列利用Primer5对克隆引物进行设计。实验中筛选得到8条较长MYB转录因子基因序列, 设计的克隆引物见表 1。
cDNA混样模板的构建  将冷冻的红花材料液氮研磨, 用TRIzol进行裂解, 利用TRIzol伴侣试剂盒进行RNA提取, 具体实验方法见商品使用说明书。提取的RNA通过凝胶电泳进行完整度检测, 通过NanoDropTM分光光度计ND1000进行浓度测定。利用试剂盒RR047A在PCR仪上进行RNA的反转录。RNA提取和反转录过程中的耗材为去除RNA酶的材料。将3个时期的花RNA同时提取并进行反转录, 将得到的cDNA混样, 用于后续MYB转录因子基因的克隆。
基因克隆  以构建的混样cDNA为模板, 以2×TransTaq High Fidelity (HiFi) PCR SuperMix Ⅱ进行PCR反应。反应体系(20 μL): 2×TransTaq High Fidelity (HiFi) PCR SuperMix 10 μL, ddH2O 7 μL, 上、下游引物各1 μL, cDNA模板1 μL (50 ng·μL-1)。PCR反应条件为: 95 ℃预变性4 min, 95 ℃变性30 s, 56 ℃退火30 s, 72 ℃延伸2 min, 共34个循环, 72 ℃延伸5 min, 12 ℃保温。1%琼脂糖凝胶电泳后切取目的胶块, 利用胶回收试剂盒进行DNA片段回收, 回收片段利用T载体连接试剂盒连接到T载体中, 转化大肠杆菌进行克隆。
序列分析  使用ORFfinder (https://www.ncbi.nlm.nih.gov/orffinder/)对序列开放阅读框及氨基酸进行预测, 使用ProtParam (http://web.expasy.org/protparam/)对蛋白质进行分子量、等电点、平均疏水性进行分析, 使用GO (http://www.geneontology.org/)及Pfam (http://pfam.xfam.org/)对克隆基因进行功能注释。使用IBS1.0对序列结构域进行作图(http://ibs.biocuckoo.org/online.php)。使用DNAMAN软件(版本4.0, Lynnon Biosoft, Quebec, Canada)进行核苷酸序列和蛋白质比对的分析。使用MEGA4软件相邻连接法(neighbor-joining)构建系统进化树, bootstrap检验的重复次数为500次。
MYB转录因子Real-time PCR分析  依据克隆的MYB转录因子序列, 利用Primer 5设计Real-time PCR用特异性引物, 设计参数选择克隆产物大小为200~500 bp, 引物长度为20 ± 2 bp。引物序列见表 2, 其中红花60S rRNA为内参基因。设计的引物通过琼脂糖凝胶电泳测试引物的特异性。使用SYBR PrimeScript RT-PCR试剂盒进行Real-time PCR实验, 实验进行3次重复。反应体系(20 μL): 2×SYBR Ⅱ 10 μL, ddH2O 7 μL, 上、下游引物各1 μL, cDNA模板1 μL (约50 ng·μL-1)。PCR反应条件: 95 ℃预变性3 min, 95 ℃变性10 s, 57 ℃退火延伸30 s, 共43个循环, 65~95 ℃进行熔解曲线绘制。
结果与分析
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1 cDNA混样模板构建
高质量的cDNA模板是基因克隆的关键因素之一。为了尽可能对筛选的8个转录因子进行克隆, 实验选择红花不同时期的混样cDNA作为克隆模板。本研究选择了红花3个时期(图 1A), 对3个时期的管状花提取了RNA (图 1B)。结果可知, 3个样品提取的RNA条带清晰无降解。随后分别对3个时期的RNA进行反转录成cDNA, 将3个时期的cDNA样品等量混和, 保证后续基因克隆有高质量的模板。
2 MYB转录因子基因克隆
以构建的cDNA混样模板, 对筛选的8个MYB转录因子基因进行克隆(图 2)。由图可知, 6个转录因子基因(CtMYB-TF1、CtMYB-TF2、CtMYB-TF4、CtMYB-TF5、CtMYB-TF6及CtMYB-TF7)有清晰单一条带, 而2个转录因子(CtMYB-TF3及CtMYB-TF8)无明显条带。实验将CtMYB-TF1、CtMYB-TF2、CtMYB-TF4、CtMYB-TF5、CtMYB-TF6及CtMYB-TF7目的条带切胶回收后连接到pMD19-T转入DH5a进行克隆, 对阳性菌液送公司测序。返回结果表明, 实验成功对6个MYB转录因子基因进行了克隆。
3 序列分析
对克隆得到的6个MYB转录因子基因进行了序列分析(表 3)。结果表明, CtMYB-TF1的CDS长度为903 bp、CtMYB-TF2为459 bp、CtMYB-TF4为1 131 bp、CtMYB-TF5为804 bp、CtMYB-TF6为723 bp及CtMYB-TF7为1 233 bp, 都具有DNA结合功能, 属MYB类转录因子。实验利用Pfam对克隆的6个转录因子蛋白序列进行了结构域预测, 发现克隆到的6个MYB转录因子均有MYB类转录因子家族DNA结合区域(图 3)。研究已将克隆到6个MYB转录因子基因相关序列信息提交到NCBI GenBank中, 基因名(GenBank登录号)分别为: CtMYB-TF1 (MF156605)、CtMYB-TF2 (MF156606)、CtMYB-TF4 (MF156607)、CtMYB-TF5 (MF156608)、CtMYB-TF6 (MF156609)及CtMYB-TF7 (MF156610)。
4 MYB转录因子进化分析
本实验将克隆得到的转录因子对应氨基酸序列进行了进化分析, 实验选择了红花中已报道的CtMYB1[12]、拟南芥已报道的AtMYB12[9]及AtMYBL2[11], 以及拟南芥转录因子库(http://plntfdb.bio.uni-potsdam.de/v3.0/index.php?sp_id=ALY)中随机3个MYB转录因子共12个转录因子进行了进化分析。结果(图 4)发现, 实验克隆到的MYB转录因子间彼此关系较远, 特别是CtMYB-TF4、CtMYB-TF5、CtMYB-TF6、CtMYB-TF7之间关系, 暗示克隆到转录因子可能具有不同的生物学功能, 特别是CtMYB-TF6同其他克隆到的5个转录因子关系较远。其中仅发现CtMYB-TF7转录因子同AtMYB12和AtMYBL2进化关系相对较近, 暗示CtMYB-TF7可能参与黄酮类成分的调控功能。
5 MYB转录因子表达分析
实验利用Real-time PCR测定了克隆的6个MYB转录因子基因在根、茎、叶及花3个时期中相对表达量(图 5)。CtMYB-TF1在花中3个时期表达量相对较高, 在根、茎及叶中表达量较低; CtMYB-TF2在根和茎中表达量较高, 在叶和花3个时期中表达量较低; CtMYB-TF4在根、茎、叶及花中均有表达, 但表达量较低; CtMYB-TF5、CtMYB-TF6及CtMYB-TF7在根、茎及叶中表达量低, 在花3个时期表达量高。与其他部位和时期比较, CtMYB-TF6在花时期Ⅰ表达最高, CtMYB-TF5及CtMYB-TF7在花时期Ⅱ表达最高。
讨论
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本研究参考Huang等[13]发表的在线数据, 从筛选得到的8个较长片段的MYB转录因子基因序列, 成功克隆了6个MYB转录因子基因, 另外2个MYB转录因子基因没有克隆, 作者推测这2个MYB转录因子在花中表达量较低或不表达, 后面实验将可以构建红花其他部位的混样cDNA, 以此为模板对MYB转录因子基因进行克隆。
进化关系远近可以一定程度反应功能的相关性。实验将克隆到的6个MYB转录因子基因, 连同红花中已报道的CtMYB1[12]和拟南芥中AtMYB12[9]及AtMYBL2[11]等12个MYB转录因子进行了进化分析。结果发现, 克隆MYB转录因子之间彼此关系较远, 其中CtMYB-TF7与已报道的两个黄酮类成分调控转录因子AtMYB12[9]和AtMYBL2[11]关系较近, 这暗示了CtMYB-TF7可能参与了黄酮类成分合成的调控。同时, 实验利用RT-PCR对克隆的6个转录因子进行了组织表达分析, 分析表明CtMYB-TF5、CtMYB-TF6及CtMYB-TF7在根、茎及叶中表达量低, 在花中表达量较高。其中CtMYB-TF6在花时期Ⅰ表达量高, CtMYB-TF5及CtMYB-TF7在花时期Ⅱ表达量高。结合红花黄酮类成分主要在花中积累及进化关系, 表明CtMYB-TF7可能参与了红花黄酮类成分的调控功能, CtMYB-TF5和CtMYB-TF6可能参与红花花中其他生理功能。后续研究将开展相关实验, 验证CtMYB-TF7转录因子是否参与调控红花黄酮类成分的合成。
MYB转录因子家族成员众多, 广泛参与了植物的生长发育各个环节中。模式植物拟南芥已报道的转录因子就有近200个[17]。本实验利用在线二代数据仅分析到8个较长MYB片段, 当前三代测序技术已陆续推出且在中药资源研究方面有较好应用[18-20], 后续研究可以尝试利用三代测序仪器进行测序, 更全面的对红花MYB转录因子基因进行克隆。
基金
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  • 国家自然科学基金资助项目(81573544)
  • 四川省教育厅科研项目(17ZB0150)
  • 四川省科技支撑计划资助项目(2014SZ0156)
  • 成都中医药大学校基金资助项目(ZRYY1610)
  • 成都中医药大学校基金资助项目(ZRQN1647)
文献
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参考文献 引证文献
排序方式：
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2018年第53卷第1期
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文章信息
doi: 10.16438/j.0513-4870.2017-0614
  • 接收时间：2017-06-26
  • 首发时间：2026-01-15
  • 出版时间：2018-01-12
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  • 收稿日期：2017-06-26
  • 修回日期：2017-09-05
基金
国家自然科学基金资助项目(81573544)
四川省教育厅科研项目(17ZB0150)
四川省科技支撑计划资助项目(2014SZ0156)
成都中医药大学校基金资助项目(ZRYY1610)
成都中医药大学校基金资助项目(ZRQN1647)
作者信息
    1.中药资源系统研究与开发利用国家重点实验室培育基地, 四川 成都 611137
    2.成都中医药大学药学院, 四川 成都 611137

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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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