摘要
植物先天免疫在感受到病原微生物来源的分子后激活,这些分子包括病原物保守的病原相关分子模式(PAMPs)以及分泌到植物细胞内的效应蛋白(effector)。PAMPs引发的免疫反应(PTI)和效应蛋白诱导的免疫反应(ETI)是植物抵抗病原物攻击的强有力武器。毒性病原细菌通过Ⅲ型分泌系统将大量效应蛋白分泌进入宿主细胞,抑制宿主免疫反应从而有效生长和繁殖,但是大部分效应蛋白的生化功能和分子机制并不清楚。因此研究这些效应蛋白在寄主体内的靶蛋白和互作机制,将有助于我们深入理解细菌致病机理和植物免疫应答。
野油菜黄单胞菌野油菜致病变种(Xcc, Xanthomonas campestris pvcampestris)是一种在全球范围内引起十字花科植物黑腐病的重要病原细菌。AvrAC是一个广泛存在于Xcc菌株中的效应蛋白,利用Xcc和拟南芥互作模式系统,我们研究证实AvrAC在植物叶肉细胞中起到毒性效应蛋白的功能,能够抑制植物先天免疫反应增强病原细菌在寄主植物上的致病性。AvrAC可以与拟南芥免疫信号通路中两个重要的细胞质受体类激酶BIK1和RIPK直接互作,进一步对其生化功能研究表明AvrAC是一个尿苷5’-单磷酸转移酶,通过对底物BIK1和RIPK功能至关重要的激活环区保守的丝氨酸和苏氨酸进行尿苷单磷酸修饰来掩盖这两个位点,阻止它们的磷酸化,并抑制这两个受体激酶活性进而抑制由它们所介导的植物免疫信号转导。
在拟南芥Col-0的维管束组织中,AvrAC又能起到无毒效应蛋白的功能,可以诱导弱的ETI,并且部分依赖于植物ETI中的两个重要蛋白RAR1和EDS1。AvrAC也能够同细胞自噬蛋白ATG8家族基因互作,相关实验也表明AvrAC所激活的抗性可能和自噬发生相关。
AvrAC是目前报道的唯一一个具有尿苷单磷酸转移酶活性的细菌效应蛋白,我们的研究结果很好的阐明了Xcc是如何利用一种独特的生化和分子机制来调节植物免疫系统的。
Plant innate immunity is activated upon the perception of conservedPathogen-Associated Molecular Patterns (PAMPs) at the cell surface or ofpathogen effector proteins inside the plant cell. Together, PAMP-triggeredimmunity (PTI) and effector-triggered immunity (ETI) constitute powerfuldefenses against various phytopathogens. A hallmark of many virulentphytopathogenic bacteria is their ability to deliver a plethora of effectorproteins into host cell by conversed type Ⅲ secretion system to promoteparasitism. While a number of effector proteins have been shown to inhibithost immunity, the biochemical function and molecular basis remainsunknown for the vast majority of these effectors. Therefore, identification ofplant targets and research of their interactions will make us know more aboutthe mechanisms of bacterial virulence and plant resistance.
Xanthomonas campestris pv campestris (Xcc) is a causal agent of blackrot diseases on numerous crucifer plants such as Brassica and Arabidopsis inthe world. The type Ⅲ effector protein AvrAC exists in all three sequencedstrains of Xcc. Using the Arabidopsis-Xcc as a model system, our researchdemonstrates that AvrAC plays a role in virulence in mesophyll tissues ofArabidopsis. It can strongly inhibit plant PTI and ETI and contribute to Xccvirulence in Arabidopsis by specifically targeting Arabidopsis BIK1andRIPK, two receptor-like cytoplasmic kinases (RLCKs) known to mediateimmune signaling. Further analysis shows that AvrAC is an uridylyltransferase that adds UMP to and masks conserved phosphorylation sites inthe activation loop of BIK1and RIPK. This leads to reduced kinase activity ofBIK1and RIPK and consequently inhibits downstream signaling.
In vascular tissue of Arabidopsis ecotype Col-0, AvrAC also can berecognized by some kind of protein to trigger a weak ETI. Our results indicatethat AvrAC induced resistence is partially dependent on RAR1and EDS1, twoimportant components respectively involved in R protein stability and ETIsignaling pathway. AvrAC has ability to interact with ATG8family genesinvolved in autophagosome fromation, which suggests that avirulence function of AvrAC maybe relates to plant cell autophagy process.
AvrAC is the only effector protein possesses uridylyl transferase activityamong all sequenced phytopathogenic bacteria at present. The work describedhere illustrates a unique biochemical mechanism by which the Xcc bacteriummodulates the plant innate immune system.
引文
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