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preferences might change after infection). However, the infected hosts now effectively have the genotype of the pathogen, and transmission acts as a birth process converting susceptible hosts into infected individuals that can also be considered as “shared natural enemies” of uninfected hosts of all susceptible species. We can briefly examine a submatrix of food web interactions for specialist and generalized pathogens within a food web.


Specialist parasites and competing host species

Pathogens shared between competing host species

In these two matrices of species interaction, host species A and B compete with each other for resources such as food or space, and each host species has a pathogen associated with it (thus infected hosts of species A are characterized as “species” Ia). In the case of specialist parasites (upper matrix), infected hosts of species A cannot infect species B; the complementary case operates for the lower matrix, where both species of pathogen infect both species of host. The main consequence of host species sharing nonspecific parasite species is that several elements of the interactions matrix have to be converted (across the main diagonal) from “zeros” into “plus–minus” consumer–resource relationships. If we are concerned with the stability properties of the web, then May (1973) has shown that the dominant eigenvalues of this matrix have to be negative if there is to be any hope of web stability. In May’s initial formulation, increased species diversity and hence increased connectance should reduce the probability that the web is stable. However, although the net effect of shared pathogens is to increase the connectance of the food web, this occurs in a subtle and important way. Namely, the conversion of specific pathogens to generalized pathogens greatly increases the proportion



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