The role of sigma factor competition in bacterial adaptation under prolonged starvation

The study of adaptive microbial evolution in the laboratory can illuminate the genetic mechanisms of gaining fitness under a pre- defined set of selection factors. Laboratory evolution of bacteria under long- term starvation has gained importance in recent years because of its ability to uncover adaptive strategies that overcome prolonged nutrient limitation, a condition often encountered by natural microbes. In this evolutionary paradigm, bacteria are maintained in an energy- restricted environment in a growth phase called long- term stationary phase (LTSP). This phase is characterized by a stable, viable population size and highly dynamic genetic changes. Multiple independent iterations of LTSP evolution experiments have given rise to mutants that are slow- growing compared to the ancestor. Although the antagonistic regulation between rapid growth and the stress response is well- known in bacteria (especially Escherichia coli), the growth deficit of many LTSP- adapted mutants has not been explored in detail. In this review, I pinpoint the trade- off between growth and stress response as a dominant driver of evolutionary strategies under prolonged starvation. Focusing on mainly E. coli- based research, I discuss the various affectors and regulators of the competition between sigma factors to occupy their targets on the genome, and assess its effect on growth advantage in stationary phase (GASP). Finally, I comment on some crucial issues that hinder the progress of the field, including identification of novel metabolites in nutrient- depleted media, and the importance of using multidisciplinary research to resolve them.