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Significance Since their discovery more than 50 years ago, dinucleoside tetraphosphates have been found in all realms of life, at levels correlated with a variety of intriguing physiological changes. However, their molecular mechanism of action has largely remained an enigma. Here we present evidence that in bacteria they are incorporated as an alternative chain-initiating nucleotide during transcription by RNA polymerase, generating RNAs that bear a novel nucleoside tetraphosphate cap at the 5′ end. Their assimilation into nascent transcripts is remarkably efficient, up to nine times the efficiency of nucleoside triphosphates. These and other findings suggest that dinucleoside tetraphosphates influence bacterial physiology not by signaling via a hypothetical receptor, but rather by modifying cellular RNAs and impacting their function.

Np 4 A alarmones function in bacteria as precursors to RNA caps

Np ₄ A alarmones function in bacteria as precursors to RNA caps