Mixed-pathogen infections of vectors rarely are considered in the epidemiological literature, although they may occur in nature. A review of published reports shows that many Anopheles species are capable of carrying sporozoites of > 1 plasmodium species, of doing so simultaneously in field conditions, and of acquiring and transmitting these in experimental situations. Mixed-species infections in mosquito populations occur at frequencies greater than or equal to the product of the constituent species prevalences, whereas human populations have apparent mixed-species infections at frequencies less than or equal to their corresponding expected values. We present a model for the accumulation of parasite infections over the lifespan of a mosquito that explains this surplus of mixedspecies infections. However, the expected frequencies of mixed infections on the basis of our model are greater than those found in nature, indicating that the sampling by mosquitoes of Plasmodium species from human malaria infections may not be random.
KeywordsAnopheles spp; Plasmodium spp; malaria transmission; parasite ecologyThe Pioneering studies of parasite community dynamics have focused on helminths (e.g., Schad 1963, Kennedy 1975, Holmes 1983, Esch et al. 1990. Recently Dobson (1985Dobson ( , 1990 and Dobson and Roberts (1994) have stressed the role of species life-histories in structuring helminth communities and have extended the general ecological principle that as a constituent species becomes more aggregated in its distribution, the relative importance of interspecific competition declines in its population regulation. Although data on the aggregation of Plasmodium species in Anopheles is essentially anecdotal, its variability indicates that Plasmodium species interactions may warrant more concerted investigation.In mixed-species malaria infections of humans, one of the constituent Plasmodium species typically dominates (e.g., Mayne and Young 1938, Molineaux and Grammicia 1980, Looareesuwan et al. 1987, Fox and Strickland 1989. Cohen (1973) analyzed the epidemiological literature and reported a general deficit of detectable mixed-species infections, associated this deficit with splenomegaly, and inferred an underlying heterologous immunity. Hence, the excess of mixed-species infections found during the Carlo project was unexpected, but it also was interpreted as representing species interactions of predominantly cooperative (Cohen and Singer 1979) or predominantly competitive (Molineaux and Grammicia 1980) character. Richie (1988) emphasized competitive interactions in an evolutionary context, suggesting that co-infections promote antigenic divergence, but also proposed a role for mutual facilitation in the ecological succession of Plasmodium species within a mammalian host.Neither the interactions of pathogen species within vectors or the pathogen sampling processes embodied in human-vector contacts have received much attention in the infectious-disease NIH Public Access (Turell et al. 1984, Paulson et al. 1992, V...