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Type VI secretion systems (T6SS) are macromolecular machines of the cell envelope of Gram-negative bacteria responsible for bacterial killing and/or virulence towards different host cells. Here, we characterized the regulatory mechanism underlying expression of the enteroagregative  Escherichia coli sci1  T6SS gene cluster. We identified Fur as the main regulator of the  sci1  cluster. A detailed analysis of the promoter region showed the presence of three GATC motifs, which are target of the DNA adenine methylase Dam. Using a combination of reporter fusion, gel shift, and  in vivo  and  in vitro  Dam methylation assays, we dissected the regulatory role of Fur and Dam-dependent methylation. We showed that the  sci1  gene cluster expression is under the control of an epigenetic switch depending on methylation: fur binding prevents methylation of a GATC motif, whereas methylation at this specific site decreases the affinity of Fur for its binding box. A model is proposed in which the  sci1  promoter is regulated by iron availability, adenine methylation, and DNA replication.

An Epigenetic Switch Involving Overlapping Fur and DNA Methylation Optimizes Expression of a Type VI Secretion Gene Cluster