The temporal dynamics of slightly deleterious mutations in Escherichia coli and Shigella spp

The Shigella are recently emerged clones of Escherichia coli, which have independently adopted an intracellular pathogenic lifestyle. We examined the molecular evolutionary consequences of this niche specialization by comparing the normalized, directional frequency profiles of unique polymorphisms within 2,098 orthologues representing the intersection of five E. coli and four Shigella genomes. We note a surfeit of AT-enriching changes (GC-->AT), transversions, and nonsynonymous changes in the Shigella genomes. By examining these differences within a temporal framework, we conclude that our results are consistent with relaxed or inefficient selection in Shigella owing to a reduced effective population size. Alternative interpretations, and the interesting exception of Shigella sonnei, are discussed. Finally, this analysis lends support to the view that nucleotide composition typically does not lie at mutational equilibrium but that selection plays a role in maintaining a higher GC content than would result solely from mutation bias. This argument sheds light on the enrichment of adenine and thymine in the genomes of bacterial endosymbionts where purifying selection is very weak.