棉花<i>GhFUL1</i>基因及其启动子的克隆和功能分析

doi:10.7668/hbnxb.20193992

摘要/Abstract

摘要：

为探究GhFUL1基因在棉花分枝发育进程中的功能,从棉花中克隆MADS-box家族GhFUL1基因及其启动子,对该基因进行功能研究,并对其启动子进行活性分析。结果表明,棉花中GhFUL1基因开放阅读框726 bp,编码241个氨基酸。进化树分析表明,GhFUL1与其他物种中可可的TcAGL8-1亲缘关系最近。烟草亚细胞定位分析结果表明,GhFUL1定位在细胞核和细胞膜。不同组织材料qRT-PCR分析表明,GhFUL1基因在茎尖中表达量最高。构建过表达载体转化拟南芥,结果表明,转基因拟南芥阳性株系中GhFUL1基因表达量显著增加,转基因株系的开花时间、花器官形态与野生型相比无明显变化,但是开花时侧枝数增加。qRT-PCR结果表明,细胞分裂素氧化/脱氢酶基因AtCTK1和AtCTK6表达量减少,推测GhFUL1基因可能通过调控细胞分裂素合成途径影响分枝。利用启动子分析网站PlantCARE预测可知,GhFUL1启动子区域不仅有TATA-box、CAAT-box核心元件,还有参与光响应、生长素响应以及胁迫应答等顺式作用元件。将GhFUL1启动子构建到表达载体pBI121检测其启动子活性,通过对拟南芥转基因阳性株系进行GUS染色,发现GhFUL1启动子在拟南芥幼苗期的茎尖分生组织和成熟期的花器官中特异性表达。初步揭示GhFUL1基因及其启动子转化拟南芥后在其分枝发育过程中具有一定的功能。

关键词: 棉花, GhFUL1, 亚细胞定位, 功能分析, 启动子活性, GUS染色

Abstract:

To investigate the function of GhFUL1 gene in branching development of cotton,the GhFUL1 gene in the MADS-box family and its promoter were cloned from cotton,and the function and promoter activity of the gene were analyzed.The results indicated that the open reading frame (ORF) of GhFUL1 gene was 726 bp and encoded 241 amino acids.Phylogenetic tree analysis showed that the amino acid sequence of FUL subclade was conserved,and GhFUL1 owned the highest similarity to Cacao TcAGL8-1 except for the homologous proteins in cotton.Subcellular localization result showed that GhFUL1 was localized in nucleus and membrane.The expression level of different cotton tissues was examined,and the result showed that GhFUL1 was highly expressed in shoot apical meristem.Overexpressing GhFUL1 in Arabidopsis thaliana increased the number of lateral branches by decreasing the expression of cytokinin oxidation/dehydrogenase genes AtCTK1 and AtCTK6.Compared with the wild type,transgenic plants did not result in significant changes of the flowering time and floral organ development.The present results suggested that GhFUL1 might affect the number of branches by regulating the cytokinin synthesis pathway.The promoter of GhFUL1 was predicted to include the TATA-box and CAAT-box elements,and cis-acting elements related to light response,auxin response and stress response by PlantCARE website.Then the promoter of GhFUL1 was cloned and constructed into expression vector pBI121 to detect promoter activity.The GUS staining of transgenic Arabidopsis lines showed that GhFUL1 promoter was specifically expressed in shoot apical meristem at seedling stage and flower organ at maturity.This study revealed that the gene and promoter of GhFUL1 played an important role in the Arabidopsis branching development process after transformation.

Key words: Cotton, GhFUL1, Subcellular localization, Function analysis, Promoter activity, GUS staining

张晓红, 许佩阳, 闫绪, 张贝贝, 于婵婵. 棉花GhFUL1基因及其启动子的克隆和功能分析[J]. 华北农学报, 2023, 38(5): 60-68. doi: 10.7668/hbnxb.20193992.

ZHANG Xiaohong, XU Peiyang, YAN Xu, ZHANG Beibei, YU Chanchan. Cloning and Functional Analysis of GhFUL1 Gene and Its Promoter in Cotton[J]. Acta Agriculturae Boreali-Sinica, 2023, 38(5): 60-68. doi: 10.7668/hbnxb.20193992.

http://www.hbnxb.net/CN/Y2023/V38/I5/60

图/表 10

表1 GhFUL1基因克隆及表达分析引物

Tab.1 Primers used in GhFUL1 gene cloning and expression analysis

引物名称 Primer name	引物序列(5'-3') Primer sequence(5'-3')	用途 Purpose
GhFUL1-F	ATGGGGAGGGGTAGGGTTCAG	基因克隆
GhFUL1-R	CTCAATACGAGGAATCATCCA	基因克隆
pGhFUL1-F	CAAAAGGCTAATTCAAGCTCA	启动子克隆
pGhFUL1-R	TTTTCTTCCAAATTTCGACC	启动子克隆
qGhFUL1-F	CAATTGCCAAGATTCATCCTC	荧光定量PCR
qGhFUL1-R	CCCTCTTCTCTTCCATTGCT	荧光定量PCR
GhACTIN4-F	ATCCTCCGTCTTGACCTTG	荧光定量PCR
GhACTIN4-R	TGTCCGTCAGGCAACTCAT	荧光定量PCR
AtUBQ5-F	CTCGCCGACTACAACATCCAG	荧光定量PCR
AtUBQ5-R	CCTTCCTCAAACGCTGAACC	荧光定量PCR
AtCKX7-F	GGTTTTGCTACAGCCAGCTCC	荧光定量PCR
AtCKX7-R	GGTCAGCACCGTTGACAAAC	荧光定量PCR
AtCKX1-F	TACCTTCATTTGACCGTTGGAG	荧光定量PCR
AtCKX1-R	CCTGTAACAATCTCTAGCTGG	荧光定量PCR
AtCKX6-F	TATCAGAGCTAAGCTACGTGAC	荧光定量PCR
AtCKX6-R	GATCGGAGTTTTACCTCAGACAC	荧光定量PCR

图1 GhFUL1基因克隆及基因结构分析 A.GhFUL1基因克隆:1.GhFUL1基因扩增产物,M.DNA Marker;B.GhFUL1基因结构。

Fig.1 GhFUL1 gene cloning and the gene structure analysis A.GhFUL1 gene clone:1.Amplified product of the GhFUL1 gene,M.DNA Marker;B.GhFUL1 gene structure.

表2 GhFUL1同源基因信息

Tab.2 GhFUL1 homologous gene information

基因名 Gene name	基因ID Gene ID	染色体ID Chromosome ID	起始位点 Start position	终止位点 End position	链 Strand	氨基酸相似度/% Amino acid identity
GhFUL1-D	Gh_D07G0671	D07	7915954	7933806	-	95.47
GhFUL1-A	Gh_A07G0605	A07	8448050	8466601	-	100.00
GhFUL2-D	Gh_D03G0922	D03	31494711	31500162	+	82.26
GhFUL2-A	Gh_A03G0634	A03	17989506	17995387	+	81.88

图2 GhFUL1氨基酸序列比对(A)及进化树分析(B)

Fig.2 The amino acid multiple sequence alignment (A) and phylogenetic analysis (B) of GhFUL1 protein

图3 GhFUL1组织特异性表达分析柱上不同小写字母表示在0.05水平差异显著。

Fig.3 Tissue specific expression analysis of GhFUL1 gene Different lowercase letters above the bars indicate significant difference at the 0.05 probability level.

图4 GhFUL1蛋白的亚细胞定位分析

Fig.4 Subcellular localization analysis of GhFUL1 protein

图5 转基因拟南芥表型鉴定及相关基因表达量分析 A.转基因拟南芥表型鉴定(WT为野生型);B.野生型拟南芥的莲座叶;C.转基因拟南芥的莲座叶;D.转基因拟南芥基因组DNA中GhFUL1检测(N和P分别表示阴性和阳性对照,数字代表不同拟南芥转基因株系,M表示DNA Marker);E.转基因拟南芥中GhFUL1基因表达水平;F.转基因拟南芥中细胞分裂素氧化/脱氢酶基因的表达量变化。柱上不同小写字母和大写字母分别表示在0.05和0.01水平差异显著。

Fig.5 Phenotype identification and expression analysis of related genes in transgenic Arabidopsis A.Phenotypic identification of transgenic Arabidopsis(WT indicates wild type);B.The rosette leaves of wild-type Arabidopsis;C.The rosette leaves of transgenic Arabidopsis;D.Detection of GhFUL1 in the genomic DNA of transgenic Arabidopsis(N and P indicate negative and positive control,number indicates the different transgenic lines,M indicates DNA Marker);E.Relative expression level of GhFUL1 in different transgenic lines;F.The expression profiles of genes involved in regulating cytokinin.Different lowercase and capital letters above the bars indicate significant difference at the 0.05 and 0.01 levels,respectively.

表3 GhFUL1顺式作用元件预测

Tab.3 The predicted cis-acting elements of GhFUL1

元件 Element	链 Strand	数量 Number	序列 Sequence	功能注释 Function annotation
TATA-box	+/-	51	TATATA	核心启动子元件
CAAT-box	+/-	22	CCAAT	启动子和增强子区域中常见的顺式作用元件
TGACG-motif/CGTCA-motif	+/-	2	TGACG/CGTCA	参与MeJA反应的顺式作用元件
ATCT-motif/Box4	+	2	AATCTAATCC/ATTAAT	参与光响应的保守DNA模块的部分元件
AT-rich element	-	1	ATAGAAATCAA	AT-rich DNA结合蛋白结合位点
TGA-element	+	1	AACGAC	生长素响应元件
TCA-element	+	1	CCATCTTTTT	参与SA响应的顺式作用元件
ARE	+/-	2	AAACCA	厌氧诱导必须元件
GT1-motif	+	4	GGTTAA	光响应元件
CCAAT-box	+	1	CAACGG	MYBHv1结合位点

图6 GhFUL1克隆及转基因拟南芥鉴定 A.GhFUL1克隆;B.转基因株系GUS基因PCR检测;C.转基因株系启动子序列PCR检测。M.DNA Marker,数字代表不同拟南芥转基因株系。

Fig.6 Cloning of GhFUL1 and identification of transgenic Arabidopsis A.Cloning of GhFUL1;B.PCR detection of transgenic plants with GUS gene;C.PCR detection of transgenic plants with GhFUL1.M.DNA Marker,and the numbers indicate different transgenic lines.

图7 转GhFUL1-GUS拟南芥的GUS染色 A.播种后7 d的幼苗;B.成熟莲座叶; C.花序;D.荚。标尺为1 000 μm。

Fig.7 GUS staining in different tissues of transgenic Arabidopsis with GhFUL1-GUS A.The seedling after 7 days of seeding;B.Mature rosette leaf; C.Inflorescence;D.Pod.Scale bar=1 000 μm.

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棉花GhFUL1基因及其启动子的克隆和功能分析

Cloning and Functional Analysis of GhFUL1 Gene and Its Promoter in Cotton

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