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Antimycins are an extended family of depsipeptides that are made by filamentous actinomycete bacteria and were first isolated more than 60 years ago. Recently, antimycins have attracted renewed interest because of their activities against the anti-apoptotic machineries inside human cells which could make them promising anti-cancer compounds. The biosynthetic pathway for antimycins was recently characterised but very little is known about the organisation and regulation of the antimycin ( ant ) gene cluster. Here we report that the  ant  gene cluster in  Streptomyces albus  is organized into four transcriptional units; the  antBA ,  antCDE ,  antGF  and  antHIJKLMNO  operons. Unusually for secondary metabolite clusters, the  antG  and  antH  promoters are regulated by an extracytoplasmic function (ECF) RNA polymerase sigma factor named  σ  AntA  which represents a new sub-family of ECF  σ  factors that is only found in antimycin producing strains. We show that  σ  AntA  controls production of the unusual precursor 3-aminosalicylate which is absolutely required for the production of antimycins.  σ  AntA  is highly conserved in antimycin producing strains and the −10 and −35 elements at the  σ  AntA  regulated  antG  and  antH  promoters are also highly conserved suggesting a common mechanism of regulation. We also demonstrate that altering the C-terminal Ala-Ala residues found in all  σ  AntA  proteins to Asp-Asp increases expression of the  antFG  and  antGHIJKLMNO  operons and we speculate that this Ala-Ala motif may be a signal for the protease ClpXP.

Regulation of antimycin biosynthesis by the orphan ECF RNA polymerase sigma factorσAntA

Regulation of antimycin biosynthesis by the orphan ECF RNA polymerase sigma factorσ^AntA