Plant pathogens threaten global food security. During early infection, oomycetes and fungi, including the fungal rice destroyer Magnaporthe oryzae, often grow in intimate contact with living host plant cells. During this growth stage the pathogen deploys secreted proteins (effectors) to suppress host defenses. In turn, the plant can recognize these effectors via intracellular resistance (R) proteins to trigger immunity. However, pathogens can lose or rapidly alter effectors to enable host jumps and cause new epidemics. Identifying pathogen cytoplasmic effectors and host targets can safeguard agriculture by informing which genomic features to deploy against which pathogen populations. Unfortunately, however, new fungal cytoplasmic effectors are difficult to predict from genomes because they are highly diverse and lack recognizable features. Furthermore, preventing disease beyond effector discovery and blockage requires mechanistic details, which are sparse. This proposal will address these knowledge gaps by leveraging recent key findings related to the role of mRNA translation in effector secretion and infection success, to clarify the rules for effector evolution, discovery and deployment that will be applicable across a broad class of pathogens. The experimental and educational objectives of this project will be integrated to inspire undergraduates to excel in research, enable graduate students to develop as mentors, and provide all students with the tools to succeed in STEM