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Originally published as Genetics Published Articles Ahead of Print on February 19, 2006.
Genetics, Vol. 173, 461-472, May 2006, Copyright © 2006
doi:10.1534/genetics.105.049445
The Influence of Hitchhiking and Deleterious Mutation Upon Asexual Mutation Rates
Michael E. Palmer1 and Marc Lipsitch
Department of Epidemiology, Harvard School of Public Health, Boston, Massachusetts 02115
1 Corresponding author: 3841 18th St., San Francisco, CA 94114.
E-mail: genetics{at}mepalmer.net
The question of how natural selection affects asexual mutation rates has been considered since the 1930s, yet our understanding continues to deepen. The distribution of mutation rates observed in natural bacteria remains unexplained. It is well known that environmental constancy can favor minimal mutation rates. In contrast, environmental fluctuation (e.g., at period T) can create indirect selective pressure for stronger mutators: genes modifying mutation rate may "hitchhike" to greater frequency along with environmentally favored mutations they produce. This article extends a well-known model of Leigh to consider fitness genes with multiple mutable sites (call the number of such sites 
). The phenotypic effect of such a gene is enabled if all sites are in a certain state and disabled otherwise. The effects of multiple deleterious loci are also included (call the number of such loci 
). The analysis calculates the indirect selective effects experienced by a gene inducing various mutation rates for given values of , , and T. Finite-population simulations validate these results and let us examine the interaction of drift with hitchhiking selection. We close by commenting on the importance of other factors, such as spatiotemporal variation, and on the origin of variation in mutation rates.
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