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Genetic robustness, the preservation of an optimal phenotype in the face ofmutations, is critical to the understanding of evolution as phenotypicallyexpressed genetic variation is the fuel of natural selection. The origin ofgenetic robustness, whether it evolves directly by natural selection or it is acorrelated byproduct of other phenotypic traits, is, however, unresolved.Examining microRNA (miRNA) genes of several eukaryotic species, Borenstein andRuppin (Borenstein et al. 2006, PNAS 103: 6593), showed that the structure ofmiRNA precursor stem-loops exhibits significantly increased mutationalrobustness in comparison with a sample of random RNA sequences with the samestem-loop structure. The observed robustness was found to be uncorrelated withtraditional measures of environmental robustness -- implying that miRNAsequences show evidence of the direct evolution of genetic robustness. Thesefindings are surprising as theoretical results indicate that the directevolution of robustness requires high mutation rates and/or large effectivepopulation sizes only found among RNA viruses, not multicellular eukaryotes.Introducing a novel measure of environmental robustness based on theequilibrium thermodynamic ensemble of secondary structures of the miRNAprecursor sequences we demonstrate that the biophysics of RNA folding, inducesa high level of correlation between genetic (mutational) and environmental(thermodynamic) robustness, as expected from the theory of plastogeneticcongruence introduced by Ancel and Fontana (Ancel et al. 2000, J. Exp. Zool.288: 242). In light of theoretical considerations we believe that thiscorrelation strongly suggests that genetic robustness observed in miRNAsequences is the byproduct of selection for environmental robustness.

Congruent evolution of genetic and environmental robustness in microRNA