Determinants of Density- and Frequency-Dependent Fitness in Competing Plant Pathogens

Using mathematical models, we investigated how infection and sporulation characteristics of competing plant pathogens determine the density and frequency dependence of relative fitness. Two models, one for the infection stage and one for the sporulation stage of a pathogen's life cycle, describe reproductive output of pathogen strains in mixture as a function of the strains' population densities. Model parameters include infection and sporulation efficiencies, carrying capacities on leaves for sporulating lesions and spore production, and coefficients of interstrain competitive effects in both life cycle stages. Although the models were originally developed for rust fungi, they are generally applicable to any organism with distinct colonization (e.g., infection) and propagative (e.g., sporulation) life cycle stages. In this work, paired hypothetical strains were assigned equal baseline parameter values. Parameters were then altered one at a time for one or both strains, and relative fitness was calculated over a range of densities and strain frequencies. Except for infection efficiency, the fitness benefit conferred by an advantage in a single parameter was always density dependent. Relative fitness was frequency dependent whenever inter- and intrastrain competitive effects were not equal. These results suggest that the fitness of pathogens in nature is rarely fixed, but, rather, may typically be highly dependent on the densities and frequencies of all coexisting strains in a habitat.