sigma(54) is the promoter recognition subunit of the form of bacterial
RNA polymerase that transcribes from promoters with enhancer elements
. DNase footprinting experiments show that sigma(54) is attached selec
tively to the template strand, which must be single-stranded for trans
cription initiation. sigma(54) remains bound at the promoter after cor
e polymerase begins elongation, in contrast to the well-established si
gma(70)-holoenzyme transcription cycle. Permanganate footprinting expe
riments show that the bound sigma(54) and the elongating core RNA poly
merase downstream of it are each associated with a single-stranded DNA
region. Template commitment assays show that the promoter-bound sigma
(54) must be reconfigured before reinitiation of transcription can occ
ur. This unexpected pathway raises interesting possibilities for trans
criptional regulation, especially with regard to control at the level
of reinitiation.