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Many infectious diseases exist in several pathogenic variants, or strains, which interact via cross-immunity. It is observed that strains tend to self-organise into groups, or clusters. The aim of this paper is to investigate cluster formation. Computations demonstrate that clustering is independent of the model used, and is an intrinsic feature of the strain system itself. We observe that an ordered strain system, if it is sufficiently complex, admits several cluster structures of different types. Appearance of a particular cluster structure depends on levels of cross-immunity and, in some cases, on initial conditions. Clusters, once formed, are stable, and behave remarkably regularly (in contrast to the generally chaotic behaviour of the strains themselves). In general, clustering is a type of self-organisation having many features in common with pattern formation.

Original publication




Journal article


J Theor Biol

Publication Date





75 - 83


Animals, Antigenic Variation, Communicable Diseases, Immunity, Innate, Models, Immunological, Species Specificity