大豆GmGST基因簇基因序列分析及诱导表达分析

doi:10.13560/j.cnki.biotech.bull.1985.2024-0980

摘要/Abstract

摘要：

目的探究GmGST基因结构及其表达特征，为开展GmGST在大豆抗大豆花叶病毒病中的功能奠定基础。方法运用RT-PCR从抗病材料X149中克隆第7号染色体上同一位点GmGSTU12（Glyma.07G139700）、GmGSTU13（Glyma.07G139800）、GmGSTU16（Glyma.07G140100）、GmGSTU47（Glyma.07G140200），并对其进行生物信息学分析；利用RT-qPCR分析GmGST在抗病材料X149中受大豆花叶病毒株系SC15、激素（MeJA、ABA、ETH和SA）、H₂O₂诱导表达情况和组织特异性表达情况；同时，采用紫外分光光度计检测抗病材料X149受SC15侵染后谷胱甘肽S-转移酶（GSTs）活性。结果基因扩增结果表明，GmGSTU12、GmGSTU13、GmGSTU16、GmGSTU47的开放阅读框长度分别为678、678、952和669 bp；生信分析表明，4个基因都属于GST的Tau家族，分别为无信号肽的细胞质、叶绿体和细胞核蛋白，其中，GmGSTU12、GmGSTU13、GmGSTU47为稳定的亲水蛋白，GmGSTU16为疏水性蛋白，均是一种无跨膜结构蛋白；RT-qPCR分析表明，GmGSTU13在叶中的表达量最高，GmGSTU12、GmGSTU16、GmGSTU47在根中的表达量最高；4个基因均受SC15和H₂O₂诱导上调表达；此外，GmGSTU12和GmGSTU13主要响应MeJA诱导表达；GmGSTU16主要响应MeJA和SA诱导表达；GmGSTU47主要响应MeJA和ABA诱导表达；GSTs活性检测表明受SC15诱导12 h后GSTs活性显著上升，且在48 h活性最大。结论 4个GmGST基因均属于GST Tau类家族，均能响应SC15、外源激素（MeJA、ABA、ETH、SA）和H₂O₂诱导表达。

关键词: 大豆, 大豆花叶病毒, GmGST, 基因扩增, 生信分析, 表达分析

Abstract:

Objective To investigate the gene structure and expression characteristics of GmGST, laying the foundation for studying its function in soybean resistance to soybean mosaic virus. Method The gene GmGSTU12 (Glyma.07G139700), GmGSTU13 (Glyma.07G139800), GmGSTU16 (Glyma.07G140100), and GmGSTU47 (Glyma.07G140200) were cloned from the resistant material X149 using RT-PCR. The method of bioinformatics was used to analyze GmGST bioinformatics. RT-qPCR was used to analyze the expressions of GmGST genes induced by SC15, hormones (MeJA, ABA, ETH and SA), H₂O₂and tissue-specific expression in the resistant material X149, and a UV spectrophotometer was used to detect the activities of glutathione S- transferase (GSTs). Result The results of gene amplification showed that the lengths of the open reading frames of GmGSTU12, GmGSTU13, GmGSTU16 and GmGSTU47 were 678, 678, 952, and 669 bp, respectively. Bioinformatics analysis showed that the four genes belonged to the Tau family of GSTs, which were cytoplasmic, cytoplasmic, chloroplast and nuclear proteins without signal peptides, among which GmGSTU12, GmGSTU13 and GmGSTU47 were stable hydrophilic proteins, and GmGSTU16 were hydrophobic proteins, all of which were non-transmembrane structural proteins. RT-qPCR analysis showed that the expression of GmGSTU13 was the highest in the leaves, and the expressions of GmGSTU12,GmGSTU16 and GmGSTU47 in the roots were the highest. All four genes were up-regulated by SC15 and H₂O₂. In addition, GmGSTU12 and GmGSTU13 were mainly expressed in response to MeJA induction. GmGSTU16 was mainly expressed in response to the induction of both MeJA and SA. GmGSTU47 was mainly expressed in response to the induction of both MeJA and ABA. The dectection of GSTs activity showed that GSTs activity increased significantly at 12 h after SC15 induction and reached its maximum at 48 h. Conclusion All four GmGST genes belong to the Tau class of the GST family and can be induced by SC15, exogenous hormones (MeJA, ABA, ETH, SA), and H₂O₂.

Key words: soybean, soybean mosaic virus, GmGST, gene amplification, bioinformatics analysis, expression analysis

赵婧, 郭茜, 李睿琦, 雷滢炀, 岳爱琴, 赵晋忠, 殷丛丛, 杜维俊, 牛景萍. 大豆GmGST基因簇基因序列分析及诱导表达分析[J]. 生物技术通报, 2025, 41(5): 129-140.

ZHAO Jing, GUO Qian, LI Rui-qi, LEI Ying-yang, YUE Ai-qin, ZHAO Jin-zhong, YIN Cong-cong, DU Wei-jun, NIU Jing-ping. Sequence Analysis and Induced Expression Analysis of GmGST Gene Cluster Genes in Soybean[J]. Biotechnology Bulletin, 2025, 41(5): 129-140.

图/表 17

表1 扩增引物信息

Table 1 Primer information for amplification

基因 Gene	正向引物 Forward primer （5′‒3′）	反向引物 Reverse primer （5′‒3′）	退火温度 Annealing temperature （℃）
GmGSTU12	TTACCAGCTATACTTGTTCC	ATTCAAACAGCAATAACTCAA	55
GmGSTU13	GGTTAGAAGTGCTACAATACAAA	AATTCAAACAGCAGTAACTCAAC	56
GmGSTU16	ACCTTAGGACCCATCACAGAC	GATTGTGTGAACTACTTGTCG	58
GmGSTU47	AGTCAAGAGAAGAAGTGAAGTG	AGGTTGTTGACATTTTCGCT	55

表2 在线软件网址

Table 2 Online software URLs

软件名称 Software name	软件网址 Software URL
SignalP4.1	http：//www.cbs.dtu.dk/services /SignalP-4.1/
PlantCARE	http：//bioinformatics.psb.ugent.be/webtools/plantcare/html/
ProtParam	http：//web.expasy.org/protparam/
SOPMA	https：//npsaprabi.ibcp. fr/cgibin/npsa_automat.pl？page=/NPSA/npsa_gor4.html
Phyre2	http：//www.sbg.bio.ic.ac.uk/phyre2/html
Evolview	http：//www.omicsclass.com/article/671
MEME	http：//meme suite.org/
GSDS2.0	http：//gsds.cbi.pku.edu.cn/
TMHMM-2.0	http：//www.cbs.dtu.dk/services/TMHMM/
WoLF PSORT	https：//wolfpsort.hgc.jp/
Pfam	https：//www.ebi.ac.uk/interpro/entry/pfam

表3 实时荧光定量引物序列

Table 3 Primers' sequences for real-time quantitative PCR

基因 Gene	正向引物 Forward primer （5′‒3′）	反向引物 Reverse primer （5′‒3′）
GmTublin	GGAGTTCACAGAGGCAGAG	CACTTACGCATCACATAGC
GmGSTU12	TGTGTGCAGGGTCCAGATTG	GCTATGGGCTGCTCATTGTG
GmGSTU13	GCGAAGAATCCCTTAACCACC	AGCGGGCTTTGAAGAAGGTA
GmGSTU16	GGGTCCAGATTGCCCTCAAA	ACGGGGTTGGATTTGAGAAGT
GmGSTU47	CAGGGTGCAGATTGCTCTCA	CAAGGGACTCGGATATGGGC

表4 RT-qPCR反应体系

Table 4 RT-qPCR reaction system

组分 Component	体积 Content/μL
MonAmp™ RapidStart Universai SYBR^® Green qPCR Mix	10
Forward primer	0.4
Reverse primer	0.4
cDNA	2
ddH₂O	7.2

图1 GmGST基因的克隆M: Trans4.5K marker; 1: GmGSTU12;2: GmGSTU13; 3: GmGSTU16; 4: GmGSTU47

Fig. 1 Cloning of GmGST genes

图2 GmGST基因的CDS序列比对

Fig. 2 CDS sequence alignment of GmGST genes

表5 GmGST 蛋白的跨膜结构、信号肽和亚细胞定位

Table 5 Transmembrane helices、signal peptide and subcellular localizationof GmGST proteins

蛋白名称 Protein name	信号肽 Signal peptide	细胞定位 Subcellular localization
GmGSTU12	无 No	细胞质Cytoplasm
GmGSTU13	无 No	细胞质Cytoplasm
GmGSTU16	无 No	叶绿体Chloroplast
GmGSTU47	无 No	细胞核Nucleus

图3 GmGST的启动子顺式作用元件分析ABRE：脱落酸响应元件；GARE-motif：赤霉素响应元件；G-box：光响应元件；TCA-element：水杨酸响应元件；MBS：干旱胁迫响应元件；ARE：厌氧诱导元件；ERE：乙烯响应元件；CGTCA-motif：茉莉酸响应元件；WUN-motif：伤害响应元件

Fig. 3 Analysis of promoter cis-acting elements of the GmGST geneABRE: Abscisic acid response element; GARE-motif: gibberellin response element; G-box: light-responsive element; TCA-element: salicylic acid response element; MBS: drought stress response element; ARE: anaerobic recovery element; ERE: ethylene responsive element; CGTCA-motif: jasmonic acid response element; WUN-motif: damage response element

表6 GmGST基因序列信息和理化性质分析

Table 6 Analysis of GmGST gene sequence information and physicochemical properties

基因名称 Gene name	基因全长 Gene length （bp）	CDS序列 CDS sequence （bp）	氨基酸数 Amino acid amounts	分子式 Molecular formula	分子量 Mw （kD）	等电点 pI	不稳定系数 Instability index	亲水性平均值 GRAVY
GmGSTU12	1 473	678	225	C₁₂₁₈H₁₈₅₅N₂₉₃O₃₃₄S₂	26.01	5.39	30.27	-0.130
GmGSTU13	1 700	678	225	C₁₂₁₈H₁₈₆₄N₂₉₂O₃₃₅S₂	26.02	5.30	30.22	-0.138
GmGSTU16	1 269	952	316	C₁₆₆₄H₂₅₅₉N₃₉₉O₄₆₄S₈	35.66	5.74	37.76	0.007
GmGSTU47	1 989	669	222	C₁₁₈₄H₁₈₃₁N₂₉₁O₃₂₇S₄	25.50	6.13	36.38	-0.175

表7 GmGST的二级结构预测

Table 7 Prediction of secondary structure and subcellular localization of GmGST

蛋白质名称 Protein name	α螺旋 Alpha helix （%）	延伸链 Extended strand （%）	无规则卷曲 Random coil （%）
GmGSTU12	54.22	11.56	34.22
GmGSTU13	54.67	11.56	33.78
GmGSTU16	59.49	10.93	29.58
GmGSTU47	54.50	10.36	35.14

图4 GmGST蛋白三级结构（A）、基因结构（B）分析与蛋白的保守元件（C）和保守基序（D）分析

Fig. 4 Tertiary structure (A), gene structure (B) analysis, and conserved element (C) and conserved sequence (D) analysis of GmGST protein

图5 GmGST 蛋白系统进化分析AtGST：拟南芥；HbGST：橡胶树；TcGST：可可树；PtGST：胡杨果；ZmGST：玉米；GsGST：野生型大豆；MtrGST：苜蓿；CcGST：辣椒；PvGST：菜豆；PDGST：毛果杨

Fig. 5 Analysis of the phylogenetic evolution of GmGST proteinsAtGST: Arabidopsis thaliana; HbGST: Hevea brasiliensis; TcGST: Theobroma cacao; PtGST: Populus euphratica; ZmGST: Zea mays; GsGST: Glycine soja Siebold; MtrGST: Medicago sativa; CcGST: Capsicum annuum L.; PvGST: Phaseolus vulgaris; PDGST: Populus pilosa

图6 大豆GmGST基因受SC15诱导表达模式分析*： P<0.05， **： P<0.01， ***： P<0.001. The same below

Fig. 6 Analysis of the expression pattern of GmGST gene induced by SC15 in soybean

图7 大豆谷胱甘肽S-转移酶活性

Fig. 7 Activity of soybean glutathione S-transferase

图8 GmGST基因家族在不同组织中的表达分析

Fig. 8 Expression analysis of the GmGST gene family in different tissues

图9 激素处理大豆GmGST基因的表达模式分析A：MeJA处理；B：ABA处理C：ETH处理D：SA处理

Fig. 9 Analysis of the expression pattern of GmGST gene in hormone-treated soybeanA: MeJA treatment; B: ABA treatment; C: ETH treatment; D: SA treatment

图10 H2O2处理大豆GmGST基因的表达模式分析

Fig. 10 Analysis of the expression pattern of GmGST gene in H2O2-treated soybean

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