Transcription attenuation in bacteria: theme and variations

Premature termination of transcription, or attenuation, is an efficient RNA-based regulatory strategy that is commonly used in bacterial organisms. Attenuators are generally located in the 5' untranslated regions of genes or operons and combine a Rho-independent terminator, controlling transcription, with an RNA element that senses specific environmental signals. A striking diversity of sensing elements enable regulation of gene expression in response to multiple environmental conditions, including temperature changes, availability of small metabolites (such as ions, amino acids, nucleobases or vitamins), or availability of macromolecules such as tRNAs and regulatory proteins. The wide distribution of attenuators suggests an early emergence among bacteria. However, attenuators also display a great mobility and lability, illustrated by a multiplicity of recent horizontal transfers and duplications. For these reasons, attenuation systems are of high interest both from a fundamental evolutionary perspective and for possible biotechnological applications.