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George Maccdonald published a model of superinfection in 19501. In presenting the model, Macdonald was trying to solve a problem: if super-infection was possible, then how long would it take until a person would clear all of the parasites to become uninfected again? Macdonald proposed a solution, but there was a problem with the mathematical formulation (An essay by Paul Fine, from 1975, is recommended reading)2: the model that Macdonald described didn’t match the function he used. In a model formulated as part of the Garki project (Dietz K, et al., 1974)3, the model allowed for superinfection and proposed a useful approximating model. One solution came from taking a hybrid modeling approach (Nåsell I, 1985)4, which we have discussed in a vignette about the Multiplicity of Infection (MoI).

The model Macdonald described was also formulted in queuing theory, where the model is called M/M/. In this model, each new infection increases the MoI by one; the transition rate to a higher MoI is the force of infection (FoI), denoted h. Each parasite can clear at some rate, r.

The following diagram shows how the fraction of the population with MoI =i, denoted ζi, changes over time.


ζ0hrζ1h2rζ2h3rζ3h4r


While the infinite system of differential equations can be formulated and solved numerically, Nåsell showed that the system can be described by a simple equation. Let m denote the mean MoI.

dmdt=hrm

He also showed that the distribution of the MoI would converge to a Poisson distribution:

M(t)fM(ζ;m)=Pois(m(t))