小麦<i>Tappc3A</i>基因克隆及功能预测

doi:10.7668/hbnxb.20193798

摘要/Abstract

摘要：

PEPC可催化磷酸烯醇式丙酮酸(PEP)生成草酰乙酸(OAA)进入三羧酸循环,对植物的生长发育和逆境适应发挥重要作用。但目前尚无关于PEPC参与植物器官发育的报道。发掘并研究小麦Tappc3A在花发育中的生物学功能,为探究小麦雄蕊同源转化为雌蕊性状的分子机制提供新的线索。通过PCR技术从CM28TP和HTS-1中克隆Tappc3A基因,使用生物信息学工具分析基因序列及系统进化关系,利用qRT-PCR技术分析Tappc3A在小麦幼穗不同发育阶段和不同生殖器官中的表达水平,通过原核表达分析Tappc3A基因编码的蛋白功能是否正常,利用小麦RNA-Seq数据库分析Tappc3A与其他调控花器官发育基因之间的共表达情况。Tappc3A基因的ORF长2 901 bp,编码966个氨基酸残基,具有典型的磷酸烯醇式丙酮酸羧化酶(PEPase)保守结构域,N端具有保守的丝氨酸(Ser,S)可逆磷酸化位点(SIDAQLR),C端具有植物型PEPC蛋白特征序列(QNTG),第774位氨基酸为C3植物PEPC典型的丙氨酸。聚类分析也表明,Tappc3A属于C3型PEPC家族。qRT-PCR分析表明,在小麦幼穗发育的3个阶段,二棱期至小花分化期和药隔时期,HTS-1中的Tappc3A基因的表达量高于CM28TP,且Tappc3A在雌蕊(P)和雌蕊化雄蕊(PS)中的表达量显著高于雄蕊(S)。原核表达结果显示,Tappc3A基因编码的蛋白能催化PEP生成OAA,并且在IPTG诱导后活性明显增强。基因共表达分析显示,Tappc3A可能参与了小麦花器官的形态发生。小麦Tappc3A可能参与小麦雌蕊发育,其在雄蕊中的过量表达可能与雄蕊同源转化为雌蕊性状形成有一定的关联。

关键词: 小麦, PEPC基因, 基因表达, 雌蕊发育

Abstract:

PEPC catalyzes phosphoenolpyruvate(PEP)to generate oxaloacetate(OAA)to participate the tricarboxylic acid(TCA),which plays an important role in development and stress adaptation of plant.However,there have been no reports of PEPC involvement in plant organ development.Exploration and study on biological function of Tappc3A gene in wheat flower development,and provides new clues to explore the molecular mechanism of homologous conversion of stamens into pistillody in wheat.The Tappc3A gene was cloned from CM28TP and HTS-1 by PCR,and the sequence and phylogenetic tree were analyzed by bioinformatics tools,and the expression level of Tappc3A gene in different developmental stages and different reproductive organs of wheat young spikes was analyzed by using qRT-PCR,and whether the protein function encoded by Tappc3A gene was analyzed by prokaryotic expression.The wheat RNA-Seq database was used to analyze the co-expression of Tappc3A gene and other genes that regulate flower organ development.The ORF of Tappc3A gene was 2 901 bp in length,encoding 966 amino acid residues,with a typical phosphoenolpyruvate carboxylase(PEPase)conserved domain,a conserved serine(Ser,S)reversible phosphorylation site(SIDAQLR)at the N-terminal,a plant-type PEPC protein signature sequence(QNTG)at the C-terminal,and the 774th amino acid was the typical alanine of C3 plant PEPC.Cluster analysis also showed that Tappc3A belonged to the C3 type PEPC family.qRT-PCR analysis,in three stages of wheat young spike development,showed that the expression of Tappc3A gene in HTS-1 was higher than that in CM28TP at the dichotomous stage to the florescence differentiation stage and the pharmacophore period,and the expression of Tappc3A gene was significantly higher in pistils(P)and pistillody stamens(PS)than stamens(S).The prokaryotic expression results showed that the protein encoded by Tappc3A gene,which catalyzes the production of OAA from PEP,and its activity was significantly enhanced after IPTG induction.The gene co-expression analysis showed that Tappc3A gene might be involved in the morphogenesis of wheat floral organs.Tappc3A gene might be involved in wheat pistil development,and its overexpression in stamens might be associated with the homologous transformation of stamens into pistillody trait formation.

Key words: Wheat, PEPC gene, Gene expression, Pistil development

向桂丽, 乌日娜, Yamamoto Naoki, 吴一超, 蒋进, 廖明莉, 魏淑红, 彭正松, 杨在君. 小麦Tappc3A基因克隆及功能预测[J]. 华北农学报, 2023, 38(4): 28-37. doi: 10.7668/hbnxb.20193798.

XIANG Guili, WU Rina, YAMAMOTO Naoki, WU Yichao, JIANG Jin, LIAO Mingli, WEI Shuhong, PENG Zhengsong, YANG Zaijun. Cloning and Functional Prediction of Tappc3A Gene in Wheat[J]. Acta Agriculturae Boreali-Sinica, 2023, 38(4): 28-37. doi: 10.7668/hbnxb.20193798.

http://www.hbnxb.net/CN/Y2023/V38/I4/28

图/表 8

表1 Tappc3A基因克隆、qRT-PCR及原核表达引物序列

Tab.1 Primers of Tappc3A gene used for gene cloning,qRT-PCR and prokaryotic expression

引物名称 Primer name	引物序列(5'—3') Primer sequence(5'—3')	退火温度/℃ Annealing temperature	引物用途 Primer purpose
Tappc3A-F	GGGGGAAGAGGGTTTTCTCG	56.0	克隆扩增
Tappc3A-R	CACACCATCTCACATAACATAACAGT
Tappc3A-qF	CAACATTTGGACTATCCCTTG	56.0	qRT-PCR
Tappc3A-qR	TTGGCGTTTCTCCTCTGAC
Tappc3A-edF	CCGGAATTCATGGCGCGCAA	61.5	原核表达
Tappc3A-edR	CCCAAGCTTTCAGCCGGTGT

图1 Tappc3A基因的PCR扩增及CDS序列 A.Tappc3A基因克隆:M.DL5000 Marker,1.HTS-1扩增产物,2.CM28TP扩增产物;B.Tappc3A基因的CDS序列。

Fig.1 PCR amplification and CDS sequence of Tappc3A gene A.Cloning of Tappc3A gene:M.DL5000 Marker,1.HTS-1 PCR product,2.CM28TP PCR product;B.CDS sequence of Tappc3A gene.

图2 Tappc3A与其他植物PEPC氨基酸序列比对方框分别为N端可逆磷酸化位点和C端植物型PEPC氨基酸序列; 下划线.保守的PEPC催化亚结构域;箭头.第 774 位氨基酸残基。

Fig.2 Amino acid sequence alignment of Tappc3A and PEPC in other plants Boxes are the reversible phosphorylationsite at the N terminus and plant-type PEPC motifs at the C terminus; Underlines.Conserved subdomains essential for PEPC catalysisis;Arrow.Amino acid residue in 774.

图3 Tappc3A蛋白序列分析 A.Tappc3A蛋白结构域;B.Tappc3A蛋白二级结构;C.蛋白质亲水性预测。红色区域.α-螺旋;蓝色区域.β-延伸链;黑色区域.无规则卷曲。

Fig.3 Analusis of the Tappc3A protein sequence A.Domain of the Tappc3A protein;B.Secondary structure prediction of the Tappc3A protein; C.Prediction of the hydrogen.The red part.Alpha helix;The blue part.β-extended strand;The black part.Random coil.

图4 Tappc3A与其他植物PEPC的系统进化树

Fig.4 Phylogenetic tree of Tappc3A and PEPC in other species

图5 Tappc3A基因在CM28TP和HTS-1幼穗发育的3个时期及P、PS、S的相对表达水平分析 1.二棱期至小花分化期0.2~0.5 cm;2.雌雄蕊原基形成期0.5~0.7 cm;3.药隔期0.7~1.0 cm;P.雌蕊;PS.雌蕊化雄蕊;S.雄蕊。不同字母的均值表示存在差异显著(P<0.05)。图6同。

Fig.5 Relative expression level of Tappc3A gene in CM28TP,HTS-1 and P,PS,S 1.Double ridge stage to floret differentiation stage 0.2—0.5 cm;2.Pistil and stamen primordiuxn formation stage 0.5—0.7 cm;3.Anther seperation stage 0.7—1.0 cm;P.Pistil;PS.Pistilloid-stamen;S.Stamen.Means with different letters are significant differences at the 0.05 probability level.The same as Fig.6.

图6 原核表达分析 A.重组质粒 pET-28a-Tappc3A的双酶切鉴定:M.Marker 15000;1.pET-28a-Tappc3A质粒经 EcoRⅠ、Hind Ⅲ双酶切;2.pET-28a-Tappc3A线性质粒。B.PEPC活性测定:VC.对照组;3A.试验组。

Fig.6 Prokaryotic expression analysis A.Identification of recombinant plasmid pET-28a-Tappc3A by double enzyme digestion:M.Marker 15000;1.pET-28a-Tappc3A digested by double enzyme EcoRⅠand Hind Ⅲ;2.pET-28a-Tappc3A linear plasmid.B.Enzymatic activity assay of PEPC:VC.Control group;3A.Experience group.

图7 Tappc3A共表达基因GO富集分析颜色表示富集程度;大小表示基因数量。

Fig.7 GO enrichment analysis of Tappc3A co-expression genes Color indicates enrichment degree;Size indicates number of gene.

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