A Rabies Model with Distributed Latent Period and Territorial and Diffusing Rabid Foxes

Rabies is an infectious viral disease. It is usually fatal if a victim reaches the rabid stage, which starts after the appearance of disease symptoms. The disease virus attacks the central nervous system, and then it migrates from peripheral nerves to the spinal cord and brain. At the time when the rabies virus reaches the brain, the incubation period is over and the symptoms of clinical disease appear on the victim. From the brain, the virus travels via nerves to the salivary glands and saliva. A mathematical model is developed for the spread of rabies in a spatially distributed fox population to model the spread of the rabies epizootic through middle Europe that occurred in the second half of the 20th century. The model considers both territorial and wandering rabid foxes and includes a latent period for the infection. Since the model assumes these two kinds of rabid foxes, it is a system of both partial differential and integral equations (with integration over space and, occasionally, also over time). To study the spreading speeds of the rabies epidemic, the model is reduced to a scalar Volterra-Hammerstein integral equation, and space-time Laplace transform of the integral equation is used to derive implicit formulas for the spreading speed. The spreading speeds are discussed and implicit formulas are given for latent periods of fixed length, exponentially distributed length, Gamma distributed length, and log-normally distributed length. A number of analytic and numerical results are shown pertaining to the spreading speeds. Further, a numerical algorithm is described for the simulation of the spread of rabies in a spatially distributed fox population on a bounded domain with Dirichlet boundary conditions. I propose the following methods for the numerical approximation of solutions. The partial differential and integral equations are discretized in the space variable by central differences of second order and by the composite trapezoidal rule. Next, the ordinary or delay differential equations that are obtained this way are i discretized in time by explicit continuous Runge-Kutta methods of fourth order for ordinary and delay differential systems. My particular interest is in how the partition of rabid foxes into territorial and diffusing rabid foxes influences the spreading speed, a question that can be answered by purely analytic means only for small basic reproduction numbers. I will restrict the numerical analysis to latent periods of fixed length and to exponentially distributed latent periods. The results of the numerical calculations are compared for latent periods of fixed and exponentially distributed length and for various proportions of territorial and wandering rabid foxes. The speeds of spread observed in the simulations are compared to spreading speeds obtained by numerically solving the analytic formulas and to observed speeds of epizootic frontlines in the European rabies outbreak 1940 to 1980. For instance, when I assume that all rabid foxes are territorial and the latent period has fixed length, the spreading speed c∗ is found to be about 28.3 [km/year], but when all rabid foxes diffuse with exponentially distributed length of the latent period, the spreading speed c∗ is about 105.2 [km/year]. These spreading speeds compare quite well with those found in nature and in the literature. In addition, the spreading speeds show that the epidemic model on a bounded domain Ω with Dirichlet boundary conditions shows a less severe epidemic outbreak than the epidemic model on R, and the spread of the disease modeled on Ω is not as fast as the spread of the disease modeled on R. Furthermore, the numerical results for c∗ confirm that the latent period with fixed length always gives the smallest spreading speeds among the latent periods with arbitrary length distribution and same mean length.