A microparasite typically multiplies within an infected host on a much shorter time scale than between-host transmission. A useful first model classifies hosts by infection status. A macroparasite, such as an adult parasitic worm, instead requires a count of parasites per host: two infected hosts can carry very different within-host parasite burdens. These are modelling distinctions rather than a rule determined only by an organism's physical size.
For a microparasite with no lasting immunity, a homogeneous SIS model is
The basic reproduction number is , and the prevalence describes the infected-host fraction. A SIR model adds an immune class, and a SEIR model adds a latent class. Contact structure, age, susceptibility, recovery rate and infectiousness can require several such classes. Then a next-generation matrix or operator, rather than a single averaged contact rate, sets the invasion threshold. Stochastic epidemic models are relevant to early extinction and finite populations; network or spatial models represent nonuniform contact opportunities.
For a simple macroparasite burden model, let be the fraction of hosts carrying adult worms. Assume acquisitions at rate , independent adult-worm losses at rate , and a number of infective stages in a well-mixed environmental reservoir. The immigration-death worm-burden model has
This birth-death process needs the burden distribution or its probability generating function, not just an infected/not-infected partition. Its mean satisfies . With host number fixed and each adult producing infective stages at rate , one possible environmental closure is
For this asexual, unsaturated transmission approximation, one worm's basic reproduction number is . Growth above the invasion threshold requires additional density regulation if a finite endemic burden is sought. Sex-dependent mating would change this closure.
With constant external exposure , the equilibrium burden follows a Poisson distribution with mean , giving infected-host prevalence . If exposure rates vary across hosts as a gamma distribution, the Poisson-gamma mixture gives a negative binomial distribution with mean and aggregation parameter :
The same mean burden can therefore coexist with very different prevalence and highly infected subgroups. Aggregation of macroparasite burdens affects treatment coverage, burden-dependent host mortality, immunity and parasite fecundity. For sexually reproducing worms, hosts need both sexes for fertile output; a mean alone may not determine it. With independent equally likely sexes and Poisson burden, the probability of both sexes is , illustrating mating-limited macroparasite transmission and its low-burden bottleneck.
Both microparasite and macroparasite models need heterogeneous exposure and host responses. The special extra difficulty for macroparasites is translating a distributed worm burden, and possibly worm sex or developmental stage, into transmission and host harm. Conversely, complicated within-host microparasite processes can also require more than a binary host state when the simple fast-within-host approximation fails.
Use an assortatively mixed two-risk-group SIS model with equal per-person recovery rate . Let be infectious proportions in groups of sizes . Let be their fixed contact rates and the transmission probability per contact. Suppose a small fixed fraction of high-risk contacts is with the low-risk group. Contact reciprocity in a two-group epidemic requires
For sufficiently small , both groups make most contacts within their own group, and the total cross-group contacts are of order . Define , , and . The forces of infection give
Here and hold in the small- regime. High-risk per-person cross contact is order one but its group size is order ; low-risk per-person cross contact is order . These distinctions are needed to count each cross-group partnership consistently.
Linearize about the disease-free state. In infectious fractions the transmission matrix is
In infected-individual counts the next-generation matrix is instead
The count and fraction formulations are related by a diagonal change of coordinates, so their eigenvalues coincide. The basic reproduction number is the spectral radius , not an average of the two group reproduction numbers. The larger real eigenvalue is
This model makes a particular reciprocal mixing convention explicit; different defensible contact laws can change the first-order coefficients.