Laboratory of Phytopathology, Wageningen University and Research Centre, 6708 PB, Wageningen, The Netherlands.
Proc Natl Acad Sci U S A. 2010 Apr 20;107(16):7610-5. doi: 10.1073/pnas.1002910107. Epub 2010 Apr 5.
Most fungal effectors characterized so far are species-specific and facilitate virulence on a particular host plant. During infection of its host tomato, Cladosporium fulvum secretes effectors that function as virulence factors in the absence of cognate Cf resistance proteins and induce effector-triggered immunity in their presence. Here we show that homologs of the C. fulvum Avr4 and Ecp2 effectors are present in other pathogenic fungi of the Dothideomycete class, including Mycosphaerella fijiensis, the causal agent of black Sigatoka disease of banana. We demonstrate that the Avr4 homolog of M. fijiensis is a functional ortholog of C. fulvum Avr4 that protects fungal cell walls against hydrolysis by plant chitinases through binding to chitin and, despite the low overall sequence homology, triggers a Cf-4-mediated hypersensitive response (HR) in tomato. Furthermore, three homologs of C. fulvum Ecp2 are found in M. fijiensis, one of which induces different levels of necrosis or HR in tomato lines that lack or contain a putative cognate Cf-Ecp2 protein, respectively. In contrast to Avr4, which acts as a defensive virulence factor, M. fijiensis Ecp2 likely promotes virulence by interacting with a putative host target causing host cell necrosis, whereas Cf-Ecp2 could possibly guard the virulence target of Ecp2 and trigger a Cf-Ecp2-mediated HR. Overall our data suggest that Avr4 and Ecp2 represent core effectors that are collectively recognized by single cognate Cf-proteins. Transfer of these Cf genes to plant species that are attacked by fungi containing these cognate core effectors provides unique ways for breeding disease-resistant crops.
迄今为止,大多数已鉴定的真菌效应子都是种特异性的,有利于在特定的宿主植物上致病。在感染其宿主番茄时,Cladosporium fulvum 分泌的效应子在缺乏同源 Cf 抗性蛋白的情况下作为毒力因子发挥作用,并在其存在的情况下诱导效应子触发的免疫。在这里,我们表明,C. fulvum Avr4 和 Ecp2 效应子的同源物存在于其他 Dothideomycete 类的致病性真菌中,包括 Mycosphaerella fijiensis,它是香蕉黑星病的病原体。我们证明,M. fijiensis 的 Avr4 同源物是 C. fulvum Avr4 的功能同源物,它通过与几丁质结合来保护真菌细胞壁免受植物几丁质酶的水解,尽管整体序列同源性较低,但在番茄中触发 Cf-4 介导的过敏反应 (HR)。此外,在 M. fijiensis 中发现了 C. fulvum Ecp2 的三个同源物,其中一个在缺乏或含有推定同源 Cf-Ecp2 蛋白的番茄系中分别诱导不同水平的坏死或 HR。与作为防御性毒力因子的 Avr4 不同,M. fijiensis Ecp2 可能通过与导致宿主细胞坏死的推定宿主靶标相互作用来促进毒力,而 Cf-Ecp2 可能保护 Ecp2 的毒力靶标并触发 Cf-Ecp2 介导的 HR。总的来说,我们的数据表明,Avr4 和 Ecp2 代表核心效应子,它们被单个同源 Cf 蛋白共同识别。将这些 Cf 基因转移到受含有这些同源核心效应子的真菌攻击的植物物种中,为培育抗病作物提供了独特的方法。
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