Bacterial transcription is controlled by sigma factors, the RNA polymerase
subunits that act as initiation factors. Although a single housekeeping sigma
factor enables transcription from thousands of promoters, environmentally
induced sigma factors redirect gene expression toward small regulons to carry
out focused responses. Using structural and functional analyses, we determined
the molecular basis of -10 promoter element recognition by Escherichia coli
σ(E), which revealed an unprecedented way to achieve promoter melting. Group IV
sigma factors induced strand separation at the -10 element by flipping out a
single nucleotide from the nontemplate-strand DNA base stack. Unambiguous
selection of this critical base was driven by a dynamic protein loop, which can
be substituted to modify specificity of promoter recognition. This mechanism of
promoter melting explains the increased promoter-selection stringency of
environmentally induced sigma factors.