Sensing Metabolic Signals with Nascent RNA Transcripts: The T Box and S Box Riboswitches as Paradigms

Recent studies in a variety of bacterial systems have revealed a number of regulatory systems in which the 5' region of a gene plays a key role in regulation of the downstream coding sequences. These RNA regions act in cis to determine if the full-length transcript will be synthesized or if the coding sequence(s) will be translated. Each class of system includes an RNA element whose structure is modulated in response to a specific regulatory signal, and the signals measured can include small molecules, small RNAs (including tRNA), and physical parameters such as temperature. Multiple sets of genes can be regulated by a particular mechanism, and multiple systems of this type, each of which responds to a specific signal, can be present in a single organism. In addition, different classes of RNA elements can be found that respond to a particular signal, indicating the existence of multiple alternate solutions to the same regulatory problem. The T box and S box systems, which respond to uncharged tRNA and S-adenosylmethionine (SAM), respectively, provide paradigms of two systems of this type.