大豆<i>GmFLK</i>基因的克隆及特性分析

doi:10.7668/hbnxb.20193922

摘要/Abstract

摘要：

FLK(Flowering Locus KH Domain)基因在植物开花调控过程中具有重要的作用,其功能在模式植物拟南芥中已经鉴定。为揭示大豆GmFLK基因功能,从大豆Williams 82中克隆获得GmFLK基因,并对其编码蛋白进行结构分析和生物信息学分析。在烟草叶片中检测该蛋白的亚细胞定位情况,在大豆中分析其表达模式。结果表明,GmFLK基因全长6 806 bp,含有6个外显子,5个内含子,CDS序列长1 341 bp,编码446个氨基酸。蛋白分子量大小为47.031 ku,等电点为5.46。GmFLK蛋白中α-螺旋、β-折叠和无规则卷曲所占比例分别为24.44%,14.80%和60.76%。进化分析结果显示,GmFLK与Glyma.03g248200同源关系最近,其次为苜蓿Medtr7g115340、花生ArahyQL2VNI、 ArahyUPVY9X和拟南芥AtFLK。亚细胞定位结果表明,GmFLK蛋白在细胞核、细胞质和细胞膜中均有表达。GmFLK基因启动子区域含有10类顺式作用元件,其中6类与光反应有关。GmFLK基因在花、叶、根、种子和茎中均有表达,其中在叶和种子中表达相对较高,在花中表达相对较低。GmFLK基因在长日照和短日照条件下表达模式相似,但在一天中不同时间点表达不同,在光照后4 h表达相对较低,在光照后0,12 h表达相对较高,由此推测,大豆GmFLK基因可能参与大豆开花调控途径。

关键词: 大豆, GmFLK, 基因克隆, 亚细胞定位, 基因表达

Abstract:

FLK(Flowering Locus KH Domain)plays an important role in flowering time regulation,and its function has been characterized in Arabidopsis.In order to identify the function,GmFLK was isolated from soybean Williams 82 and the encoded protein was predicted by structural and bioinformatics analysis.The subcellular localization of this protein was examined in tobacco leaves and the expression pattern of this gene was analyzed in soybean.The results showed that the GmFLK was 6 806 bp in length,containing 6 exons and 5 introns.The CDS was 1 341 bp,encoding 446 amino acids.The molecular weight of the protein was 47.031 ku,and the isoelectric point was 5.46.GmFLK protein contained three structures,α-Helix,β-Strand and Coil,which had 24.44%,14.80% and 60.76%,respectively.Phylogenetic analysis revealed that GmFLK had a close relationship with Glyma.03g248200,followed by Medicago Medtr7g115340,Arachis hypogaea ArahyQL2VNI, ArahyUPVY9X,and Arabidopsis AtFLK.Subcellular localization analysis showed that GmFLK protein was expressed in the nucleus,cytoplasm and cell membrane.GmFLK promoter contained 10 types of cis-acting elements,and 6 types were considered to be involved in photoperiod response.In addition,GmFLK was expressed in flower,leaf,root,seed and stem,and GmFLK showed relative high expression levels in leaf and seed,and relative low levels in flower.The expression patterns of GmFLK under long day and short day photoperiod conditions were similar.However,GmFLK showed different expression levels throughout the day and night,and exhibited low expression level after 4 h,lights-on,and high expression level after 0 and 12 h,lights-on.In summary,our study reveals that GmFLK might be a key factor involved in flowering time regulation.

Key words: Soybean, GmFLK, Gene clone, Subcellular localization, Gene expression

赵胜男, 高美欣, 于朝杭, 聂凯悦, 王浩然, 孟杰, 朱虹, 李帅. 大豆GmFLK基因的克隆及特性分析[J]. 华北农学报, 2023, 38(3): 26-34. doi: 10.7668/hbnxb.20193922.

ZHAO Shengnan, GAO Meixin, YU Zhaohang, NIE Kaiyue, WANG Haoran, MENG Jie, ZHU Hong, LI Shuai. Cloning and Characteristic Analysis of Soybean GmFLK Gene[J]. Acta Agriculturae Boreali-Sinica, 2023, 38(3): 26-34. doi: 10.7668/hbnxb.20193922.

http://www.hbnxb.net/CN/Y2023/V38/I3/26

图/表 10

表1 本试验中所用引物序列

Tab.1 Primers used in this study

引物名称 Primer name	序列(5'-3') Sequences(5'-3')	长度/bp Length	功能 Function
GmFLK-F	gagaacacgggggacgagctcATGGCTGAAGTGGATCAGTACACG	45	基因克隆
GmFLK-R	catgtcgactctagaggatccCTAGTACCCATAGTTTGCACCATAAAC	48
GmFLK-qRT-F	GGTTGGGCGGATTCAGGTAT	20	qRT-PCR
GmFLK-qRT-R	GCCCGTGTACTGATCCACTT	20
GmCons4-qRT-F	CGGTGGTTCTATCTTGGCATC	21
GmCons4-qRT-R	GTCTTTCGCTTCAATAACCCTA	22

图1 GmFLK基因保守结构域和基因结构分析

Fig.1 Conserved domain and gene structure analysis of GmFLK gene

图2 GmFLK蛋白二级结构和三维结构预测

Fig.2 Secondary and three dimensional structure analysis of GmFLK protein

图3 GmFLK系统进化树分析

Fig.3 Phylogenetic tree analysis of GmFLK and its homologous proteins

图4 GmFLK蛋白及其同源蛋白序列比对

Fig.4 Sequence alignment of GmFLK protein and its homologous proteins

表2 GmFLK启动子区域顺式作用元件分析

Tab.2 Cis-acting element analysis of the GmFLK promoter

编号 Number	名称 Name	位置 Location	数量 Quantity	序列 Sequence	功能 Function
1	ARE	-422,-785	2	AAACCA	对厌氧诱导顺式作用元件
2	TGACG-motif	-1 023	1	TGACG	参与MeJA响应的顺式作用调控元件
3	CAAT-box	-1 776,-1 188,-846,-491,-374,-296	6	CAAAT	启动子和增强子区域常见的顺式作用元件
	CAAT-box	-1 567,-561	2	CCAAT	启动子和增强子区域常见的顺式作用元件
3	CAAT-box	-1 746,-1 566,-1 792,-1 162,-603,-560,-461,-438,-352,-156,-50	11	CAAT	启动子和增强子区域常见的顺式作用元件
4	TATA-box	-1 073,-1 058,-1 921,-1 902,-1 768,-1 404,-1 312,-1 133,-1 120	9	TATA	核心启动子元素在转录开始的-30左右
4	TATA-box	-1 059,-1 136,-1 134	3	ATATAT	核心启动子元素在转录开始的-30左右
4	TATA-box	-1 075,-835,-1 904,-552,-510,-448,-347	7	ATTATA	核心启动子元素在转录开始的-30左右
4	TATA-box	-614,-1 405	2	ATATAA	核心启动子元素在转录开始的-30左右
4	TATA-box	-1 406,-1 135	2	TATATA	核心启动子元素在转录开始的-30左右
5	Box 4	-1 773,-1 340,-1 667,-1 283	4	ATTAAT	参与光响应的保守DNA模块的一部分
6	GATA-motif	-1 949,-1 092	2	AAGGATAAGG	光响应元件的一部分
7	GT1-motif	-673	1	GGTTAA	光敏元件
8	I-box	-1 947	1	gGATAAGGTG	光响应元件的一部分
9	Sp1	-265	1	GGGCGG	光敏元件
10	circadian	-570	1	CAAAGATATC	参与昼夜节律控制的顺式作用调节元件

图5 GmFLK基因在不同组织及不同光周期条件下表达分析 A.GmFLK基因在不同组织表达分析;B.GmFLK基因在不同光周期条件下表达分析:0~20为光照开启后取样时间,LD.长日照,SD.短日照。

Fig.5 The expression of GmFLK in different tissues and different photoperiod conditions A.The expression of GmFLK gene in different tissues;B.The expression of GmFLK gene throughout the day under different photoperiod conditions:0-20 indicate the time after lights on,LD.Long day condition,SD.Short day condition.

图6 GmFLK基因亚细胞定位载体构建 A.GmFLK基因PCR扩增电泳图:M.DL2000 Marker,1-4.GmFLK基因CDS片段扩增,约1 300 bp;B.pCAMBIA1300空载体和GmFLK表达载体经Sac Ⅰ和BamH Ⅰ双酶切:M.DL10000 Marker,1.pCAMBIA1300未酶切质粒,2.pCAMBIA1300经Sac Ⅰ和BamH Ⅰ双酶切后的线性质粒,3.GmFLK表达载体未酶切质粒,4.GmFLK表达载体经Sac Ⅰ和BamH Ⅰ双酶切后的线性质粒;C.GmFLK表达载体转化大肠杆菌:M.DL2000 Marker,1.阳性对照,2.阴性对照,3-6.单克隆菌液PCR;D.GmFLK表达载体转化农杆菌:M.DL2000 Marker,1.阳性对照,2.阴性对照,3-6.单克隆菌液PCR。

Fig.6 Construction of GmFLK gene subcellular localization plasmid A.Amplification of GmFLK gene:M.DL2000 Marker,1-4.Amplification of GmFLK CDS fragment,approximately 1 300 bp;B.pCAMBIA1300 and GmFLK expression vector plasmid digested using SacⅠand BamHⅠ:M.DL10000 Marker,1.pCAMBIA1300 plasmid,2.pCAMBIA1300 plasmid digested using SacⅠand BamHⅠ,3.GmFLK expression vector,4.GmFLK expression vector digested using SacⅠ and BamHⅠ;C.Transformation of GmFLK gene into Escherichia coli:M.DL2000 Marker,1.Positive control,2.Negative control,3-6.PCR of clones;D.Transformation of GmFLK gene into Agrobacterium tumefaciens:M.DL2000 Marker,1.Positive control,2.Negative control,3-6.PCR of clones.

图7 GmFLK基因亚细胞定位载体模式图

Fig.7 Model of GmFLK gene subcellular localization vector

图8 GmFLK蛋白亚细胞定位分析

Fig.8 Subcellular localization analysis of GmFLK protein

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