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Genes of the major histocompatibility complex (MHC) have been studied for several decades because of their pronounced allelic polymorphism. Structural allelic polymorphism is, however, not the only source of variability subjected to natural selection. Genetic variation may also exist in gene expression patterns. Here, we show that in a natural population of three-spined sticklebacks ( Gasterosteus aculeatus) the expression of MHC class IIB genes was positively correlated with parasite load, which indicates increased immune activation of the MHC when infections are frequent. To experimentally study MHC expression, we used laboratory-bred sticklebacks that were exposed to three naturally occurring species of parasite. We found strong differences in MHC class IIB expression patterns among fish families, which were consistent over two generations, thus demonstrating a genetic component. The average number of MHC class IIB sequence variants within families was negatively correlated to the MHC expression level suggesting compensatory up-regulation in fish with a low (i.e. suboptimal) MHC sequence variability. The observed differences among families and the negative correlation with individual sequence diversity imply that MHC expression is evolutionary relevant for the onset and control of the immune response in natural populations. [ABSTRACT FROM AUTHOR]

How complex life cycles of parasites are maintained is still a fascinating and unresolved topic. Complex life cycles using three host species, free-living stages, asexual and sexual reproduction are widespread in parasitic helminths. For such life cycles, we propose here that maintaining a second intermediate host in the life cycle can be advantageous for the individual parasite to increase the intermixture of different clones and therefore decrease the risk of matings between genetically identical individuals in the definitive host. Using microsatellite markers, we show that clone mixing occurs from the first to the second intermediate host in natural populations of the eye-fluke Diplostomum pseudospathaceum. Most individuals released by the first intermediate host belonged to one clone. In contrast, the second intermediate host was infected with a diverse array of mostly unique parasite genotypes. The proposed advantage of increased parasite clone intermixture may be a novel selection pressure favouring the maintenance of complex life cycles. [ABSTRACT FROM AUTHOR]

Abstract Parasite mediated selection may result in arms races between host defence and parasite virulence. In particular, simultaneous infections from multiple parasite species should cause diversification (i.e. balancing selection) in resistance genes both at the population and the individual level. Here, we tested these ideas in highly polymorphic major histocompatibility complex (MHC) genes from three-spined sticklebacks (Gasterosteus aculeatus L.). In eight natural populations, parasite diversity (15 different species), and MHC class IIB diversity varied strongly between habitat types (lakes vs. rivers vs. estuaries) with lowest values in rivers. Partial correlation analysis revealed an influence of parasite diversity on MHC class IIB variation whereas general genetic diversity assessed at seven microsatellite loci was not significantly correlated with parasite diversity. Within individual fish, intermediate, rather than maximal allele numbers were associated with minimal parasite load, supporting theoretical models of self-reactive T-cell elimination. The optimal individual diversity matched those values female fish try to achieve in their offspring by mate choice. We thus present correlative evidence supporting the ‘allele counting’ strategy for optimizing the immunocompetence in stickleback offspring. [ABSTRACT FROM AUTHOR]

Two ciliates and 16 metazoan parasites were identified in 434 0+ year three-spined sticklebacksGasterosteus aculeatus collected from two small rivers and four lakes located in Schleswig-Holstein, Germany. By repeated sampling and analysis of dispersion patterns of six frequently occurring parasites no consistent evidence was found for mortality induced by a single parasite species. Linear log-variance to log-mean abundance ratios with slopes ofc . 2 indicated negative binomial distributions for five of the six parasites. The numbers of these six parasites were combined as multiples of s.d. of each parasite species over all samples to form an ‘ individual parasitation index ’ (IPI), which showed that only in one locality a slight decrease in parasite burden occurred between September and April. In two of the lake populations, however, there was a distinct decline in the degree of dispersion in spring samples. This indicates that a combination of different species might cause parasite-induced host mortality, undetectable by patterns obtained from single species. There were differences in parasite diversity and intensity of infection among river compared to lake populations suggesting a role for parasites as selective agents in the ecological divergence of three-spined sticklebacks. Copyright 2002 The Fisheries Society of the British Isles. Published by Elsevier Science Ltd. All rights reserved. [Copyright &y& Elsevier]