Saturday, November 14, 2009

Is the Red Queen showing her face? Evidence of negative frequency dependent selection by parasites

Recently Wolinska and Spaak (2009) provide a survey of Daphnia infections by genotype across a number of lakes in Italy and Switzerland. They present their results as empirical evidence of Red Queen dynamics in which coevolution with virulent parasites generates continued evolution. Although Van Valen (1973) originally presented a macroevolutionary argument where by reciprocal selection of hosts and their parasites generates conditions for continuous change, Bell (1982) narrowed the focus as a mechanistic explanation for the evolution or maintenance of sexual reproduction through cyclical changes in genotype frequencies. Wolinska and Spaak (2009) are not addressing the evolution of sex, but looking for evidence that parasites in Daphnia populations are generating negative frequency dependent selection such that a rare genotype has an advantage. Evidence consistent with the Red Queen has been found in other systems using spatially distributed samples (e.g. Dybdahl and Lively 1995) to look non-random infection rates as well as more directly looking at changes in frequencies of common genotypes (e.g. Dybdahl and Lively 1998).

Wolinska and Spaak (2009) propose three hypotheses to test with their data. The first is that common genotypes should be either over or under infected compared to a random sample. This prediction is based on stereotyped cyclical dynamics of genotypes of hosts and parasites (image two out of sync sine waves). At some points, the common clones will be targeted by the parasites and become overly infected. As a genotype becomes common, parasites haven't started attacking this genotype yet (i.e. time lagged), so it is under infected. In their survey, the found that indeed, some of the populations showed over infection (n = 1) and other showed under infection (n = 11), although the majority of cases did show no significant difference from random infection probabilities which is predicted as being a rare event. Their second hypothesis was that common genotypes should over the course of time decline if they are being tracked by parasites. The previous sample included only different lakes; where as the data needed to test this hypothesis are temporal samples from the same location. Their additional data is consistent with common genotypes declining over time (9 out of 10 cases). However, it is unclear to me how the general trend in this data of common genotypes decreasing over time, leads to the evidence supporting the first hypothesis. Shouldn't they find many more over infected common clones? A third hypothesis that they tested regarded host-parasite interactions maintaining diversity and an evenness of genotype frequencies which their data supported.

When discussing this paper, we were interested in what happens to predictions based on Red Queen dynamics when more than one parasite is involved. Previous empirical papers and theory seems to be generally focused on a host and a common parasite, but we know hosts are attacked by all kinds of parasites and pathogens. The system described by Wolinska and Spaak (2009) involves a host hybrid complex as well as four different parasites and questions about host specialization and hybrid maintenance were addressed in a previous paper (Wolinska et al. 2007). Where is the companion theoretical work to provide testable hypotheses?


Bell, G. 1982. The Masterpiece of Nature: The Evolution and Genetics of Sexuality. University of California Press, Berkeley.

Dybdahl, M. F., and C. M. Lively. 1995. Host-Parasite Interactions: Infection of Common Clones in Natural Populations of a Freshwater Snail (Potamopyrgus antipodarum). Proceedings of the Royal Society of London. Series B: Biological Sciences 260:99-103.

Dybdahl, M. F., and C. M. Lively. 1998. Host-parasite coevolution: Evidence for rare advantage and time-lagged selection in a natural population. Evolution 52:1057-1066.

Van Valen, L. 1973. A new evolutionary law. Evolutionary Theory 1:1-30.

Wolinska, J., B. Keller, M. Manca, and P. Spaak. 2007. Parasite survey of a Daphnia hybrid complex: host-specificity and environment determine infection. Journal of Animal Ecology 76:191-200.

Wolinska, J., and P. Spaak. 2009. The cost of being common: evidence from natural Daphnia populations. Evolution 63:1893-1901.

Wolinska, J., & Spaak, P. (2009). The cost of being common: evidence from natural Daphnia populations Evolution, 63 (7), 1893-1901 DOI: 10.1111/j.1558-5646.2009.00663.x

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