On the determinants of population structure in antigenically diverse pathogens

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Abstract
Many pathogens exhibit antigenic diversity and elicit strain–specific immune responses. This potential for cross–immunity structure in the host resource motivates the development of mathematical models, stressing competition for susceptible hosts in driving pathogen population dynamics and genetics. Here we establish that certain model formulations exhibit characteristics of prototype pattern–forming systems, with pathogen population structure emerging as three possible patterns: (i) incidence is steady and homogeneous; (ii) incidence is steady but heterogeneous; and (iii) incidence shows oscillatory dynamics, with travelling waves in strain–space. Results are robust to strain number, but sensitive to the mechanism of cumulative immunity.