植物病原细菌通过复杂和精细的全局性调控网络来协调多个层面的毒性决定因子。在不同的植物病原细菌中,这些全局性的毒性调控网络控制着细菌的侵染策略、存活以及在面临寄主植物防卫系统的互作环境中实现成功侵染的病程。本文详细分析了植物病原细菌4个重要属(假单胞菌属、果胶杆菌属、黄单胞菌属和雷尔氏菌属)的模式病原菌主要的毒性调控系统,包括群体感应系统、双组分调控系统、转录激活调控子以及转录后、翻译后的调控机制。在此基础上,重点评价了一些模式菌株全局性毒性调控机制的异同点,总结了一些最新的研究进展,并绘制了精细的网络调控图。这些分析表明,虽然一些相同的调控系统控制着病原菌的毒性,但是在不同种以及种下的亚种或者致病变种中这些调控机制功能各异,对于病原菌全毒性的贡献也存在着明显的差异。
Plant pathogenic bacteria coordinate multifaceted virulence determinants through some complex and delicate global regulation networks. In the diverse species, these networks dictate bacterial infection strategies, survival, and the success in infection in the presence of host plant defense systems. In this paper, the major virulence regulation systems in the model pathogens in four genera (
植物病原细菌通过分泌多种毒性因子,干扰寄主植物正常的生理代谢过程或者抑制寄主植物的防卫反应,从而成功侵染其寄主植物。这些毒性因子通常包括胞外酶、胞外多糖、毒素、激素、脂多糖、黏附因子以及Ⅲ型效应蛋白等[
植物病原细菌的毒性调控具有一些共同的特点:(1) 细胞的群体感应具有保守性,在全局性调控网络中调控多个毒性相关表型;(2) 双组分信号转导系统,能够感知环境或者寄主来源的信号,通过磷酸化信号途径激活调控蛋白,调控下游靶标基因的表达;(3) 转录调控子或者sigma因子,能够在转录水平激活或者抑制基因的表达;(4) 转录后和翻译后调控机制,通过控制mRNA的稳定性或者通过调控靶标蛋白的稳定性来调控目标基因表达。
QS是指细菌自身能够产生可渗透性的信号分子,例如酰基化的高丝氨酸内酯类(acylated homoserine lactone,AHL)[
除木质部小菌属(
植物病原细菌AHLs介导的群体感应信号系统模式图
Model diagram of the AHLs-mediated QS system in plant pathogenic bacteria.
TCSTS是细菌感知胞外信号最主要的传导系统之一,一般由具有跨膜结构的组氨酸激酶(histidine kinase,HK)和位于胞内的感应调控子(response regulator,RR)组成。HK根据其磷酸基团转移的模式分为3种类型(
植物病原细菌双组分调控系统模式图
Model diagram of the two-component signal transduction systems in plant pathogenic bacteria.
随着更多植物病原细菌菌株全基因组序列的公布,许多双组分调控基因被发掘。目前,在假单胞菌属(
在已解析的植物病原细菌毒性调控网络中,特别是涉及T3SS调控途径中,需要一些转录调控子(例如AraC类调控子)和σ因子。虽然
Rsm转录后调控机制在一些重要的植物病原细菌中都存在,可能在毒性调控中起着共同或者相似的作用,这个调节体系由调节蛋白RsmA、RsmC和小RNA
翻译后机制通过蛋白降解来进行,这个降解过程一般发生在细胞内,需要依赖于ATP的蛋白酶。这类蛋白酶包含有4大类:ClpAP/ ClpXP[
在
假单胞菌属代表菌株的毒性调控网络
Virulence regulatory network of representative strains of phytopathogenic
在
黄单胞菌属代表菌株的毒性调控网络
Virulence regulatory network of representative strains of phytopathogenic
在青枯雷尔氏菌整个毒性调控网络中,PhcA作为全局性的调控因子,控制EPS、T3SS、胞外酶、QS系统以及噬铁素等多个毒性因子(
雷尔氏菌的代表菌株毒性调控网络
Virulence regulatory network of representative strains of phytopathogenic
在
果胶杆菌代表菌株的毒性调控网络
Virulence regulatory network of representative strains of phytopathogenic
植物病原细菌通过感知来自寄主植物、环境和QS信号,来协调其不同毒性相关基因的表达,从而表现出不同的致病表型。已有的研究证据表明,植物病原细菌的毒性调控网络以全局性调控因子或者依赖于QS调控为中心;双组分系统提供了来自于环境的输入信号,一些激活子或者sigma因子影响关键毒性因素,如T3SS或者PCWDEs基因的表达。但是,仍然有不少问题有待于研究:(1) 哪些寄主来源的信号能够激活T3SS基因的表达,不同属的细菌是否存在相似的机制, 这些信号在细菌的表面是如何被感知和传递的;(2) 为什么在不同属的病原细菌中,位于毒性调控网络中心的因子存在差异,在黄单胞菌中,位于中心的调控子是否为依赖于c-di-GMP的Clp,需要更多证据进行验证;(3) 在雷尔氏菌和黄单胞菌中都发现T4P组装蛋白能够调控T3SS基因的表达,但通过什么机制调控了T3SS,仍有待进一步研究,c-di-GMP是否参与T4P对T3SS的调控过程,也有待于进一步揭示;(4) Rsm转录后机制广泛存在于果胶杆菌、假单胞菌和黄单胞菌中,但
感谢上海交通大学农业与生物学院的李逸朗、黄梦桑、李颖、陈慧妍、梁靖聆、李子阳、高彦瑾和朗博在参考文献的搜集以及综述写作中给予的协助,感谢西南大学的张勇副教授在雷尔氏菌分型和毒性调控机理写作部分提供的宝贵意见。
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