Epidemiological models with parametric heterogeneity: Deterministic theory for closed populations

We present a unified mathematical approach to epidemiological models with parametric heterogeneity, i.e., to the models that describe individuals in the population as having specific parameter (trait) values that vary from one individuals to another. This is a natural framework to model, e.g., heterogeneity in susceptibility or infectivity of individuals. We review, along with the necessary theory, the results obtained using the discussed approach. In particular, we formulate and analyze an SIR model with distributed susceptibility and infectivity, showing that the epidemiological models for closed populations are well suited to the suggested framework. A number of known results from the literature is derived, including the final epidemic size equation for an SIR model with distributed susceptibility. It is proved that the bottom up approach of the theory of heterogeneous populations with parametric heterogeneity allows to infer the population level description, which was previously used without a firm mechanistic basis; in particular, the power law transmission function is shown to be a consequence of the initial gamma distributed susceptibility and infectivity. We discuss how the general theory can be applied to the modeling goals to include the heterogeneous contact population structure and provide analysis of an SI model with heterogeneous contacts. We conclude with a number of open questions and promising directions, where the theory of heterogeneous populations can lead to important simplifications and generalizations.Comment: 26 pages, 6 figures, submitted to Mathematical Modelling of Natural Phenomen

Adaptive Fitness Landscape for Replicator Systems: To Maximize or not to Maximize

Sewall Wright's adaptive landscape metaphor penetrates a significant part of evolutionary thinking. Supplemented with Fisher's fundamental theorem of natural selection and Kimura's maximum principle, it provides a unifying and intuitive representation of the evolutionary process under the influence of natural selection as the hill climbing on the surface of mean population fitness. On the other hand, it is also well known that for many more or less realistic mathematical models this picture is a sever misrepresentation of what actually occurs. Therefore, we are faced with two questions. First, it is important to identify the cases in which adaptive landscape metaphor actually holds exactly in the models, that is, to identify the conditions under which system's dynamics coincides with the process of searching for a (local) fitness maximum. Second, even if the mean fitness is not maximized in the process of evolution, it is still important to understand the structure of the mean fitness manifold and see the implications of this structure on the system's dynamics. Using as a basic model the classical replicator equation, in this note we attempt to answer these two questions and illustrate our results with simple well studied systems.Comment: 13 pages, 4 figure