Tuesday, July 7, 2009

Selection Mosaics or environmental interactions


Vale and Little (2009) published recent work on parasite infection variation across a temperature gradient. Specific parasite infections are often the result of genetic interactions of both the host and parasite, sometimes referred to as genotype by genotype interactions (GxG). The authors of this paper used an ideal interaction between Daphnia magna and a bacterial parasite, Pasteuria ramose. The experiment was such that they could test multiple levels on interactions. They isolated multiple host clonal lines (n = 4) as well as parasite lines (n = 4) and compared infection rates as well as parasite growth rates across three different temperatures. The paper details the experiment very well, so I'll spare details here, but a good model for future studies.

The authors found significant GxG interactions for most of the traits measured in the infection process, including both early (probability of infection) and later (parasite growth rate). However differences in genotype by environment (GxE) interactions showed up for different places in the infection timeline. The probability of infection showed a host genotype by temperature interaction, but this was a weak affect and the authors make the important point that the relative rank order wasn't changed. The reason this is key is that it is often emphasized that GxE interactions are a mechanism of the maintenance of different genotypes. If each genotype has high fitness in only some environments, and the environment varies, then there can be some period of time where polymorphism is maintained. In terms of interactions of the parasite genotype and the environment, there were initially some interactions with transmission potential and growth rate, however rank differences were again absent. The paper makes one further step and examines the combined transmission potential (spore production and infectivity). This isn't quite a measure of R0 because of complications with the effect of dose on infection rate and the interaction between parasite genotype and temperature disappears.

The study failed to find evidence of a GxGxE interaction, but the authors were correct to point out, that this is only the case for the environmental variable measured (temperature). Given that temperature is an important component of the environment for this interaction, I was surprised by this result. Perhaps, it would have been different if the difference were not just in constant temperature, but in some sort of variable environment. In the very last paragraph, Vale and Little (2009) emphasize that the lack of GxGxE interactions mean that the specificity of the interactions are robust to environmental noise. However, it is just such noise that others have proposed as important in maintaining variation. These interactions are the selection mosaics in the Geographic Mosaic Theory of Coevolution (Thompson 1999, 2005).

References

Thompson, J. N. 1999. Specific hypotheses on the geographic mosaic of coevolution. American Naturalist 153:S1-S14.

Thompson, J. N. 2005.
The Geographic Mosaic of Coevolution. University of Chicago Press, Chicago.

Vale, P. F., and T. J. Little. 2009. Measuring parasite fitness under genetic and thermal variation. Heredity online early.

Paper read


Vale, P., & Little, T. (2009). Measuring parasite fitness under genetic and thermal variation Heredity DOI: 10.1038/hdy.2009.54