红花<i>CtANR2</i>和<i>CtANR3</i>基因的克隆、结构及表达模式分析

doi:10.7668/hbnxb.20193450

摘要/Abstract

摘要：

原花青素是植物中广泛存在的一类黄酮类化合物,是人类膳食的重要营养成分,在防治病虫害方面也发挥着重要作用,花青素还原酶(ANR)是合成原花青素的关键酶。以大果球红花品种为材料,克隆得到2个CtANR基因。生物信息学分析表明,CtANR2和CtANR3的编码区分别为1 020,1 023 bp,对应基因组序列中均含有5个外显子和4个内含子,第1个外显子长度不同,其余4个外显子长度一致,内含子长度差异较大。CtANR2和CtANR3基因编码蛋白质的氨基酸数目分别为339,340个,二级结构都主要由α-螺旋和无规则卷曲构成,都属于水溶性蛋白,但CtANR2蛋白不稳定,而CtANR3为稳定的亲水性蛋白。此外,2个蛋白质都不存在信号肽和跨膜结构,可能定位于细胞外。序列比对及系统进化分析表明,CtANR2和CtANR1同源性最高,亲缘关系最近,3个CtANR蛋白与菊科植物ANR蛋白进化关系最近,与茄科、锦葵科和桑科植物ANR蛋白也同属一个大分支。组织特异性表达分析发现,CtANR2和CtANR1组织表达模式相似,都是在花中的表达量最高,初期果球表达量最低,而CtANR3则在苞片中表达量最高,根和初期果球中表达量最低。三者在不同激素胁迫下呈现出不同的表达模式,CtANR2和CtANR1在不同激素处理后表达量均下降,而CtANR3在5种激素处理后表达量均有不同程度的升高。以上结果表明,红花3个CtANR基因可能在红花不同发育阶段及抵抗非生物胁迫中有不同的分工。

关键词: 红花, 花青素还原酶, 基因克隆, 基因结构, 表达模式

Abstract:

As part of flavone compounds,proanthocyanidins (PAs)play important roles in defense against pests and diseases,and they are also a kind of nutritionally valuable component of human diet.Anthocyanidin reductases (ANRs)are involved in the biosynthesis of PAs.We cloned two CtANR genes from Carthamus tinctorius L..Bioinformatic analysis showed that the full-length CDS sequences of CtANR2 and CtANR3 were 1 020 and 1 023 bp,respectively.Both of the genomic sequences contained five exons and four introns,the length of the first exon was different while the other four exons were the same,and the lengths of introns varied greatly.CtANR2 and CtANR3 genes encoded proteins with amino acid number of 339 and 340,respectively,and their secondary structures were mainly composed of α-helix and random curl.Both of them were hydrophilic proteins,CtANR2 protein was unstable,while CtANR3 was a stable hydrophilic protein.In addition,the two proteins both had no signal peptide sequence and transmembrane structure,and might be located outside the cells.Sequence alignment and phylogenetic analysis showed that CtANR2 and CtANR1 had the highest homology and the closest genetic relationship.The three CtANR proteins had the closest evolutionary relationship with ANR of Asteraceae,and also belonged to the same large branch with ANR of Solanaceae,Malvacede and Moraceae.Expression analysis showed that the tissue expression patterns of CtANR2 and CtANR1 were similar with the highest expression level in flowers and the lowest expression level in early fruit balls,while CtANR3 was highly expressed in bracts and almost not expressed in roots and early fruit balls.They displayed different expression patterns under different hormone treatments.The expressions of CtANR2 and CtANR1 were decreased after different hormone treatments,while the expression levels of CtANR3 were increased to different degrees after hormone treatments.These results suggest that CtANR genes might play different roles in different development stages and resistance to abiotic stresses of Carthamus tinctorius L..

Key words: Carthamus tinctorius L., Anthocyanidin reductase, Gene cloning, Gene structure, Expression profile

鲁丹丹, 谭政委, 余永亮, 李磊, 许兰杰, 杨红旗, 杨青, 董薇, 安素妨, 梁慧珍. 红花CtANR2和CtANR3基因的克隆、结构及表达模式分析[J]. 华北农学报, 2023, 38(1): 84-93. doi: 10.7668/hbnxb.20193450.

LU Dandan, TAN Zhengwei, YU Yongliang, LI Lei, XU Lanjie, YANG Hongqi, YANG Qing, DONG Wei, AN Sufang, LIANG Huizhen. Cloning,Structure and Expression Profile Analysis of CtANR2 and CtANR3 Genes from Carthamus tinctorius L.[J]. Acta Agriculturae Boreali-Sinica, 2023, 38(1): 84-93. doi: 10.7668/hbnxb.20193450.

http://www.hbnxb.net/CN/Y2023/V38/I1/84

图/表 11

表1 本研究所用引物信息

Tab.1 Information of primers used in this study

引物名称 Primer name	引物序列(5'—3') Primer sequence(5'—3')	用途 Usage
CtANR2-F	ATGGAAGAGAGTAAAAGCAAGTGCA	基因扩增
CtANR2-R	CTAGATATTGCCCGATCTCTTAGCAC
CtANR3-F	ATGGAAGAGAGTAAAAGCAGCAGGT	基因扩增
CtANR3-R	CTAGATATTACCAAATCTCTTTGCATATTC
qCtANR1-F	CGACAGGATTGAGGCAACAATCAA	实时荧光定量PCR
qCtANR1-R	AGCTGCAACAACAGAGGCGGTATA
qCtANR2-F	AACGACAGGATTGAGGCA	实时荧光定量PCR
qCtANR2-R	AACAACGGAGGCGGTATA
qCtANR3-F	AGTGAGGCGGCTCGTCTAT	实时荧光定量PCR
qCtANR3-R	CAAGGGTGTCCAGCAAGTTT
Ct60S-F	TGGAGCTCATCAAGAAGGG	内参基因
Ct60S-R	GGTAAGGACCACAAGACCGTA

表2 分子生物学网站及分析内容

Tab.2 Molecular biology websites and application

在线网站 Online website	网址 Website	分析内容 Application
GSDS 2.0	http://gsds.gao-lab.org/	基因结构
CD-Search	https://www.ncbi.nlm.nih.gov/Structure/cdd/wrpsb.cgi	保守结构域
ORF Finder	https://www.ncbi.nlm.nih.gov/orffinder/	最大开放阅读框
ProtParam	https://web.expasy.org/protparam/	理化性质
ProtScale	https://web.expasy.org/protscale/	亲疏水性
SOPMA	https://npsa-prabi.ibcp.fr/cgi-bin/secpred_sopma.pl	二级结构预测
TMHMM Server v.2.0	http://www.cbs.dtu.dk/services/TMHMM/	跨膜结构
SignalP 4.0 Server	http://www.cbs.dtu.dk/services/SignalP-4.0/	信号肽
ProtComp 9.0	http://linux1.softberry.com/berry.phtml	亚细胞定位

图1 2个CtANR基因的PCR扩增结果 M.DL 2000 Marker;1.CtANR2扩增产物;2.CtANR3扩增产物。

Fig.1 PCR amplification analysis of CtANR genes M.DL 2000 Marker;1.CtANR2 PCR product;2.CtANR3 PCR product.

图2 2个CtANR基因的结构分析

Fig.2 Genome structure analysis of CtANR genes

表3 CtANR2和CtANR3蛋白的理化性质及二级结构分析

Tab.3 Physical and chemical properties and secondary structures analysis of CtANR2 and CtANR3 proteins

蛋白质 Ptotein	理化性质 Physical and chemical properties					二级结构/% Secondary structure
蛋白质 Ptotein	氨基酸数目/个 Amino acid number	等电点 pI	分子质量/ku Molecular weight	亲水性指数 GRAVY	不稳定指数 Instability index	α-螺旋 Alpha helix	无规则卷曲 Random coil	延伸链 Extended strand	β-折叠 Beta turn
CtANR2	339	5.87	37.988 36	-0.231	42.92	40.71	37.76	14.45	7.08
CtANR3	340	6.32	38.188 59	-0.268	36.16	44.41	35.59	14.12	5.88

图3 CtANR2和CtANR3蛋白的氨基酸序列比对及保守结构域分析 A.CtANR2、CtANR3和CtANR1的氨基酸序列比对:#.活性位点;*.NADP辅酶结合位点;◆.CtANR2特异的NADP辅酶结合位点;&.CtANR3特异的NADP辅酶结合位点。B.保守结构域预测。

Fig.3 Amino acid sequence alignment and conserved domains prediction of CtANR2 and CtANR3 proteins A.Amino acid sequence alignment of CtANR2,CtANR3 and CtANR1:#.Active sites;*.Coenzyme NADP binding sites;◆.Specific coenzyme NADP binding sites of CtANR2;&.Specific coenzyme NADP binding sites of CtANR3.B.Conserved domains prediction.

图4 CtANR2和CtANR3 蛋白的生物信息学分析 A.亲疏水性分析;B.跨膜结构预测;C.信号肽预测(C-score.原始剪切位点的分值;S-score.信号肽的分值;Y-score.综合剪切位点的分值;Mean S.信号肽分值的平均值;D.识别分值)。

Fig.4 Bioinformatic analysis of CtANR2 and CtANR3 proteins A.Hydrophobicity analysis;B.Transmembrane structure prediction;C.Signal peptide prediction(C-score.Raw cleavage site score; S-score.Signal peptide score;Y-score.Combined cleavage site score;Mean S.The average S-score of possible signal peptides;D.Discrimination score).

表4 CtANR2和CtANR3蛋白亚细胞定位预测结果

Tab.4 Subcellular localization prediction results of CtANR2 and CtANR3 proteins

亚细胞结构 Destination of cell compartment	存在概率Probability
亚细胞结构 Destination of cell compartment	CtANR2	CtANR3
细胞核Nucleus	0	0
质膜 Plasma membrane	0	0
胞外基质 Extracellular matrix	2.53^*	2.31^*
细胞质 Cytoplasm	1.69	0.81
线粒体 Mitochondria	1.91	1.43
内质网 Endoplasmic reticulum	0.39	0.93
过氧化物酶体 Peroxisome	2.21	2.05
溶酶体 Lysosome	1.04	0.76
高尔基体 Golgi body	0.23	1.70
液泡 Vacuole	0	0

图5 不同植物CtANR蛋白的系统进化树

Fig.5 Phylogenetic analysis of CtANR proteins from different plant species

图6 CtANR2和CtANR3基因在不同品种红花不同组织部位及不同花期花瓣的表达分析 R.根;St.茎;L.叶;B.苞片;F.初期果球;S1~S5.不同花期。不同小写字母代表差异显著(P<0.05)。图7同。

Fig.6 Relative expression level of CtANR2 and CtANR3 genes in different tissues and different florescence petals of different varieties of Carthamus tinctorius L. R.Root;St.Stem;L.Leaf;B.Bract;F.Initial fruit ball;S1—S5.Different florescence.Different lowercase letters indicate significant differences (P<0.05). The same as Fig.7.

图 7 3个CtANR基因在不同激素处理下的表达分析

Fig.7 Relative expression analysis of CtANR genes under different hormone stresses

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红花CtANR2和CtANR3基因的克隆、结构及表达模式分析

Cloning,Structure and Expression Profile Analysis of CtANR2 and CtANR3 Genes from Carthamus tinctorius L.

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红花CtANR2和CtANR3基因的克隆、结构及表达模式分析

Cloning,Structure and Expression Profile Analysis of CtANR2 and CtANR3 Genes from Carthamus tinctorius L.

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