Breaking the barriers:: Microbial effector molecules subvert plant immunity

Adaptation to specialized environments allows microorganisms to inhabit an enormous variety of ecological niches. Growth inside plant tissues is a niche offering a constant nutrient supply, but to access this niche, plant defense mechanisms ranging from passive barriers to induced defense reactions have to be overcome. Pathogens have to break several, if not all, of these barriers. For this purpose, they secrete effector molecules into plant cells to interfere with individual defense responses. Plant defense is organized in multiple layers, and therefore the action of effectors likely follows this same order, leading to a hierarchy in effector orchestration. In this review we summarize the latest findings regarding the level at which effectors manipulate plant immunity. Particular attention is given to those effectors whose mechanism of action is known. Additionally, we compare methods to identify and characterize effector molecules.