CaWRKY39在辣椒响应疫霉菌侵染中的功能分析

doi:10.16420/j.issn.0513-353x.2023-1013

摘要/Abstract

摘要：

以辣椒自交系007EA为材料，克隆了1个WRKY转录因子基因CaWRKY39，其开放阅读框为984 bp，编码327个氨基酸，含有1个WRKY结构域和C2H2型的锌指结构。氨基酸序列比对和进化树分析发现CaWRKY39与茄子SmWRKY7、番茄SlWRKY4和马铃薯StWRKY7同源性较高。基因结构分析显示CaWRKY39含有3个外显子和2个内含子。CaWRKY39启动子中含有W-box，以及响应光信号、ABA、MeJA、SA和生长素的顺式作用元件。亚细胞定位结果显示，CaWRKY39定位于细胞核和细胞质。qRT-PCR结果显示CaWRKY39在辣椒叶片中表达量最高，其次是茎、根等组织；此外其表达量在疫霉菌侵染植株叶片后明显上升。进一步通过VIGS技术沉默CaWRKY39，发现TRV:CaWRKY39植株对疫病的抗性比TRV:00对照明显降低。这些结果表明CaWRKY39可能正调控辣椒对疫病的抗性。

关键词: 辣椒, CaWRKY39, 亚细胞定位, 疫病, 功能分析

Abstract:

A WRKY transcription factor gene CaWRKY39 was cloned from pepper（Capsicum annuum L.）inbred line 007EA. The open reading frame of CaWRKY39 was 984 bp，encoding a protein of 327 amino acids，which contained a WRKY domain and a C2H2 zinc finger structure. Amino acid sequence alignment and phylogenetic tree analysis showed that CaWRKY39 had the highest homology with SmWRKY7，SlWRKY4 and StWRKY7 and gene structure analysis showed that CaWRKY39 contained three exons and two introns. Promoter analysis showed that CaWRKY39 promoter included W-box，and cis-acting elements responsive to light，ABA，MeJA，SA and auxin. Subcellular localization results showed that CaWRKY39 was located in the nucleus and cytoplasm. qRT-PCR results showed that CaWRKY39 displayed the highest expression in leaf，followed by stem，root，and other tissues，and its expression level was significantly increased after plant leaves infected by Phytophthora capsici. The function of CaWRKY39 was further studied by VIGS technology. TRV:CaWRKY39-silenced pepper plants were much sensitive to P. capsici infection compared with the TRV:00 control plants. These results of this study showed that CaWRKY39 may be involved in the resistance of pepper to phytophthora blight.

Key words: pepper, CaWRKY39, subcellular localization, Phytophthora capsica, functional analysis

吴丹, 柳佳欣, 卓林熙, 李钰, 罗英, 周勇, 杨有新, 余婷. CaWRKY39在辣椒响应疫霉菌侵染中的功能分析[J]. 园艺学报, 2024, 51(10): 2297-2310.

WU Dan, LIU Jiaxin, ZHUO Linxi, LI Yu, LUO Ying, ZHOU Yong, YANG Youxin, YU Ting. Functional Analysis of CaWRKY39 Under Phytophthora capsici Infection in Pepper[J]. Acta Horticulturae Sinica, 2024, 51(10): 2297-2310.

图/表 11

参考文献 39

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用途 Application	引物名称 Primer name	正向引物（5′-3′） Forward	反向引物（5′-3′） Reverse
亚细胞定位 Subcellular localization	CaWRKY39-GFP	CTCGATACACCAAATCGACTCTAGTCTAGAATGGCTGTGGACTTAATGATG	CAGCTCCTCGCCCTTGCTCACCATGGTACCAGAAGATTCTAAAATAAGACTACTTGTTTC
亚细胞定位 Subcellular localization	GFP	GACGCCATTTCGCCTTTTCAG	GAACTTGTGGCCGTTTACGTCG
基因沉默 VIGS	CaWRKY39	TGAGTAAGGTTACCGAATTCTCTAGAAAAGCCTCCGTTGTCTTCTTCTTCAC	TTTAATGTCTTCGGGACATGCCCGGGATATCTGCCAGTTTCATGCTTATTGC
	CaPDS	TGAGTAAGGTTACCGAATTCTCTAGAAGGTCTTCTTTGGGAACTGA	TTTAATGTCTTCGGGACATGCCCGGGAGACAAACCACCCAAACCTG
	TRV1	ATTGAGGCGAAGTACGATGG	CCATCCACAATTATTTTCCGC
实时荧光定量 qRT-PCR	qCaWRKY39	ACTCCAATCCAACAAGTCCCT	TCATCTTCAGCAGCTTTCCAG
	qCaActin	GACGTGACCTAACTGATAACCTGAT	CTCTCAGCACCAATGGTAATAACTT
	qPcActin	TTTAGTTGGGGGTCTTGTACC	CCTCCACAACCAGCAACA

用途 Application	引物名称 Primer name	正向引物（5′-3′） Forward	反向引物（5′-3′） Reverse
亚细胞定位 Subcellular localization	CaWRKY39-GFP	CTCGATACACCAAATCGACTCTAGTCTAGAATGGCTGTGGACTTAATGATG	CAGCTCCTCGCCCTTGCTCACCATGGTACCAGAAGATTCTAAAATAAGACTACTTGTTTC
亚细胞定位 Subcellular localization	GFP	GACGCCATTTCGCCTTTTCAG	GAACTTGTGGCCGTTTACGTCG
基因沉默 VIGS	CaWRKY39	TGAGTAAGGTTACCGAATTCTCTAGAAAAGCCTCCGTTGTCTTCTTCTTCAC	TTTAATGTCTTCGGGACATGCCCGGGATATCTGCCAGTTTCATGCTTATTGC
	CaPDS	TGAGTAAGGTTACCGAATTCTCTAGAAGGTCTTCTTTGGGAACTGA	TTTAATGTCTTCGGGACATGCCCGGGAGACAAACCACCCAAACCTG
	TRV1	ATTGAGGCGAAGTACGATGG	CCATCCACAATTATTTTCCGC
实时荧光定量 qRT-PCR	qCaWRKY39	ACTCCAATCCAACAAGTCCCT	TCATCTTCAGCAGCTTTCCAG
	qCaActin	GACGTGACCTAACTGATAACCTGAT	CTCTCAGCACCAATGGTAATAACTT
	qPcActin	TTTAGTTGGGGGTCTTGTACC	CCTCCACAACCAGCAACA

元件 Element	序列 Sequence	位置/bp Location	数量 Number	位点功能 Function of site
TC-rich repeats	GTTTTCTTAC	-813	1	参与防御和胁迫反应的顺式作用元件 cis-Acting element involved in defense and stress responsiveness
LAMP-element	CTTTATCA	-168/-92	2	光响应元件的一部分 Part of a light responsive element
Box 4	ATTAAT	-1 583/-939/ -348	3	参与光响应的保守DNA模块的一部分 Part of a conserved DNA module involved in light responsiveness
ATCT-motif	AATCTAATCC	-1 456/-384	2	参与光响应的保守DNA模块的一部分 Part of a conserved DNA module involved in light responsiveness
circadian	CAAAGATATC	-1 520	1	参与昼夜节律调控的顺式作用元件 cis-Acting regulatory element involved in circadian control
3-AF1 binding site	TAAGAGAGGAA	-1 643	1	光响应元件 Light responsive element
AE-box	AGAAACTT	-1 474	1	光响应模块的一部分 Part of a module for light response
ABRE	ACGTG	-1 257	1	参与脱落酸反应的顺式作用元件 cis-Acting element involved in the abscisic acid responsiveness
ARE	AAACCA	-1 960/ -1 886	2	厌氧诱导所必需的顺式作用元件 cis-Acting regulatory element essential for the anaerobic induction
TGA-element	AACGAC	-653	1	生长素响应元件 Auxin-responsive element
AT-rich sequence	TAAAATACT	-1 861	1	最大诱导子介导的激活元件 Element for maximal elicitor-mediated activation
ATC-motif	AGTAATCT	-387	1	参与光响应的保守DNA模块的一部分 Part of a conserved DNA module involved in light responsiveness
AT1-motif	AATTATTTTTTATT	-359	1	光响应模块的一部分 Part of a light responsive module
AuxRR-core	GGTCCAT	-996	1	参与生长素响应的顺式作用元件 cis-Acting regulatory element involved in auxin responsiveness
TCA-element	CCATCTTTTT	-1 983/-140	2	参与水杨酸反应的顺式作用元件 cis-Acting element involved in salicylic acid responsiveness
G-Box	CACGTT	-1 257	1	参与光响应的顺式作用元件 cis-Acting regulatory element involved in light responsiveness
CGTCA-motif	CGTCA	-218	1	参与MeJA响应的顺式作用元件 cis-Acting regulatory element involved in the MeJA-responsiveness
I-box	GTATAAGGCC	-271	1	光响应元件的一部分 Part of a light responsive element

元件 Element	序列 Sequence	位置/bp Location	数量 Number	位点功能 Function of site
TC-rich repeats	GTTTTCTTAC	-813	1	参与防御和胁迫反应的顺式作用元件 cis-Acting element involved in defense and stress responsiveness
LAMP-element	CTTTATCA	-168/-92	2	光响应元件的一部分 Part of a light responsive element
Box 4	ATTAAT	-1 583/-939/ -348	3	参与光响应的保守DNA模块的一部分 Part of a conserved DNA module involved in light responsiveness
ATCT-motif	AATCTAATCC	-1 456/-384	2	参与光响应的保守DNA模块的一部分 Part of a conserved DNA module involved in light responsiveness
circadian	CAAAGATATC	-1 520	1	参与昼夜节律调控的顺式作用元件 cis-Acting regulatory element involved in circadian control
3-AF1 binding site	TAAGAGAGGAA	-1 643	1	光响应元件 Light responsive element
AE-box	AGAAACTT	-1 474	1	光响应模块的一部分 Part of a module for light response
ABRE	ACGTG	-1 257	1	参与脱落酸反应的顺式作用元件 cis-Acting element involved in the abscisic acid responsiveness
ARE	AAACCA	-1 960/ -1 886	2	厌氧诱导所必需的顺式作用元件 cis-Acting regulatory element essential for the anaerobic induction
TGA-element	AACGAC	-653	1	生长素响应元件 Auxin-responsive element
AT-rich sequence	TAAAATACT	-1 861	1	最大诱导子介导的激活元件 Element for maximal elicitor-mediated activation
ATC-motif	AGTAATCT	-387	1	参与光响应的保守DNA模块的一部分 Part of a conserved DNA module involved in light responsiveness
AT1-motif	AATTATTTTTTATT	-359	1	光响应模块的一部分 Part of a light responsive module
AuxRR-core	GGTCCAT	-996	1	参与生长素响应的顺式作用元件 cis-Acting regulatory element involved in auxin responsiveness
TCA-element	CCATCTTTTT	-1 983/-140	2	参与水杨酸反应的顺式作用元件 cis-Acting element involved in salicylic acid responsiveness
G-Box	CACGTT	-1 257	1	参与光响应的顺式作用元件 cis-Acting regulatory element involved in light responsiveness
CGTCA-motif	CGTCA	-218	1	参与MeJA响应的顺式作用元件 cis-Acting regulatory element involved in the MeJA-responsiveness
I-box	GTATAAGGCC	-271	1	光响应元件的一部分 Part of a light responsive element