PT - JOURNAL ARTICLE AU - Kevin S. Franco AU - Zhe Sun AU - Yixiong Chen AU - Cedric Cagliero AU - Yuhong Zuo AU - Yan Ning Zhou AU - Mikhail Kashlev AU - Ding Jun Jin AU - Thomas D. Schneider TI - <em>Escherichia coli σ</em><sup>38</sup> promoters use two UP elements instead of a −35 element: resolution of a paradox and discovery that <em>σ</em><sup>38</sup> transcribes ribosomal promoters AID - 10.1101/2020.02.05.936344 DP - 2020 Jan 01 TA - bioRxiv PG - 2020.02.05.936344 4099 - http://biorxiv.org/content/early/2020/02/06/2020.02.05.936344.short 4100 - http://biorxiv.org/content/early/2020/02/06/2020.02.05.936344.full AB - In E. coli, one RNA polymerase (RNAP) transcribes all RNA species, and different regulons are transcribed by employing different sigma (σ) factors. RNAP containing σ38 (σS) activates genes responding to stress conditions such as stationary phase. The structure of σ38 promoters has been controversial for more than two decades. To construct a model of σ38 promoters using information theory, we aligned proven transcriptional start sites to maximize the sequence information, in bits, and identified a −10 element similar to σ70 promoters. We could not align any −35 sequence logo; instead we found two patterns upstream of the −35 region. These patterns have dyad symmetry sequences and correspond to the location of UP elements in ribosomal RNA (rRNA) promoters. Additionally the UP element dyad symmetry suggests that the two polymerase α subunits, which bind to the UPs, should have two-fold dyad axis of symmetry on the polymerase and this is indeed observed in an X-ray crystal structure. Curiously the αCTDs should compete for overlapping UP elements. In vitro experiments confirm that σ38 recognizes the rrnB P1 promoter, requires a −10, UP elements and no −35. This clarifies the long-standing paradox of how σ38 promoters differ from those of σ70.