植物执行者抗病基因研究进展

doi:10.11983/CBB24002

摘要/Abstract

摘要： 在与病原菌的长期斗争过程中, 植物进化出复杂而精细的免疫防御系统。抗病(resistant, R)基因的克隆和功能研究极大地促进了人们对植物免疫防御系统的理解。执行者(executor, E)基因作为一类新的植物抗病基因, 具有独特的抗病特点, 同时也是重要的抗病基因资源, 因此成为植物免疫领域的研究热点。近年来, E基因的克隆和功能机制研究取得了一系列重要进展, 但尚未见相关中文综述。该文全面总结了E基因的蛋白序列特征、与病原菌的互作机制、生物学功能及育种应用, 以期为深入理解植物-病原菌互作机制和作物抗病育种提供重要参考。

关键词: 植物-病原菌互作, 执行者基因, 基因克隆, 分子机制, 育种应用

Abstract: The survive of plants is mainly attributed to their complex and sophisticated immune defense system that has evolved through the process of battles against pathogens. The cloning and functional research of disease resistance (R) genes have greatly facilitated people’s understanding of plant immune defense systems. Executor (E) genes, as a new type of plant disease resistance genes, has unique disease resistance characteristics and is also an important resource for disease resistance genes, making it a research hotspot in the field of plant immunity. In recent years, significant progress has been made in the cloning and functional mechanism research of E genes, however, there is no Chinese review about them. This article comprehensively summarizes the protein sequence characteristics, interaction mechanisms with pathogens, biological functions and breeding applications of the E genes, aiming to provide important references for understanding the molecular mechanisms of plant-pathogen interactions and disease-resistant breeding of crops.

Key words: plant-pathogen interaction, executor genes, gene cloning, molecular mechanism, breeding application

何璐梅, 马伯军, 陈析丰. 植物执行者抗病基因研究进展. 植物学报, 2024, 59(4): 671-680.

Lumei He, Bojun Ma, Xifeng Chen. Advances on the Executor Resistance Genes in Plants. Chinese Bulletin of Botany, 2024, 59(4): 671-680.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB24002

https://www.chinbullbotany.com/CN/Y2024/V59/I4/671

图/表 5

表1 执行者(E)基因及其编码蛋白的基本信息

Table 1 Basic information of the executor (E) genes and their encoding proteins

基因	蛋白登录号	基因编码区长度(bp)	蛋白氨基酸数目(aa)	跨膜次数	物种	参考文献
Bs3	ABW82012	1366	342	0	Capsicum annuum	Römer et al., 2007; Schornack et al., 2008
Bs4C-R	AFW98885	495	164	4	C. pubescens	Strauß et al., 2012
Xa7	UMZ39519	342	113	2	Oryza sativa	Chen et al., 2021; Luo et al., 2021; Wang et al., 2021
Xa10	AGE45112	381	126	4	O. sativa	Tian et al., 2014
Xa23	AIX09985	342	113	3	O. rufipogon	Wang et al., 2015
Xa27	AFO69279	342	113	2	O. minuta	Wu et al., 2008

图1 执行者(E)基因编码蛋白的结构特征 (A) 用ClustalX软件进行5个E蛋白的氨基酸序列比对; (B) 用SOSUI软件预测的5个E蛋白的跨膜结构域(BS4C-R与XA10具有4个跨膜结构域, XA23具有3个, XA7与XA27均具有2个); (C) 用AlphaFold 2预测的5个E蛋白的三级结构(均含多个α-螺旋)。

Figure 1 Structural characteristics of proteins encoded by the executor (E) genes (A) Sequence alignment of the five E proteins using ClustalX; (B) Transmembrane domains of five E proteins predicted by SOSUI (BS4C-R and XA10 have four transmembrane domains, XA23 has three transmembrane domains, while XA7 and XA27 have two transmembrane domains); (C) 3D structure of the five E proteins predicted by AlphaFold 2 (each E protein contains multiple α-helix).

图2 转录激活类效应因子(TALE)的重复区高度变异的残基(RVD)与执行者(E)基因启动子效应蛋白结合元件(EBE)的配对关系绿色字母表示与RVD完美匹配的核苷酸, 红色字母表示与RVD不完美匹配的核苷酸, 携带*的RVD因缺失第13位氨基酸残基, 无碱基偏好性。

Figure 2 Recognition relationship between the repeat variable diresidue (RVD) of repeat domains in transcription activation- like effector (TALE) and the effector binding element (EBE) in promoter of executor (E) gene The green letters represent nucleotides that perfectly match RVD, while the red letters represent nucleotides do not perfectly match, RVD carrying * means no base preference due to the absence of the 13th amino acid residue.

表2 转录激活类效应因子(TALE)与靶基因启动子效应蛋白结合元件(EBE)的结合能力预测得分

Table 2 Prediction score of the binding ability of the transcription activation-like effector (TALE) to the effector binding element (EBE) in the target gene promoter

效应子	基因	EBE	匹配度得分值
AvrBs3	Bs3	Bs3-EBE_AvrBs3	11.05
AvrBs3	Upa20	Upa20-EBE_AvrBs3	11.42
AvrXa7	Xa7	Xa7-EBE_AvrXa7	19.65
AvrXa7	Sweet14	Sweet14-EBE_AvrXa7	26.72
PthXo3	Xa7	Xa7-EBE_PthXo3	29.95
PthXo3	Sweet14	Sweet14-EBE_PthXo3	31.87

图3 执行者(E)基因与其互作病原菌的模式图来源于水稻的E基因Xa7、Xa10、Xa23和Xa27能够增强水稻对白叶枯病菌Xoo的抗性, 来源于辣椒的E基因Bs3和Bs4C-R具有对辣椒斑点病菌Xcv的抗性。来源于大麦的Rph3基因赋予大麦对叶锈病菌(Puccinia hordei, Ph)的抗性, 而从拟南芥中克隆的WTS基因能增强对芸薹根肿菌(Plasmodiophora brassicae, Pb)的抗性。

Figure 3 Pattern diagram of executor (E) genes and their interacting pathogens The E genes Xa7, Xa10, Xa23 and Xa27 derived from rice enhance the resistance of rice to Xoo, while the E genes Bs3 and Bs4C-R derived from pepper enhance the resistance to pepper Xcv. The Rph3 gene originated from barley confers resistance to Puccinia hordei (Ph), while the WTS gene cloned from Arabidopsis enhances resistance to Plasmodiophora brassicae (Pb).

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