How to win a tug-of-war: the adaptive evolution of Phytophthora effectors.

The 'zigzag' model formulates some of the fundamental principles underpinning the dynamic interactions between pathogen effectors and plant immunity. As key virulence factors, effectors often exhibit a pattern of rapid evolution, presumably as a result of the host-pathogen arms race. Here, we summarize the current knowledge of mechanisms that may accelerate effector evolution in the highly successful Phytophthora pathogens. Recent findings on epigenetic regulation of effector genes that allows evasion of host recognition and maintenance of cost/benefit balance, and a conserved structural unit in effector proteins that may promote the evolution of virulence activities are highlighted.