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Enterobacter cloacae  has emerged as an opportunistic pathogen in healthcare-associated infections. Analysis of the genomic sequences of several  E. cloacae  strains revealed the presence of genes that code for expression of at least one type VI secretion system (T6SS). Here, we report that  E. cloacae  strain ATCC 13047 codes for two functional T6SS named T6SS-1 and T6SS-2. T6SS-1 and T6SS-2 were preferentially expressed in tryptic soy broth and tissue culture medium (DMEM), respectively. Mutants in T6SS-1-associated genes  clpV1  and  hcp1  significantly affected their ability of inter- and intra-bacterial killing indicating that T6SS-1 is required for bacterial competition. In addition, the Hcp effector protein was detected in supernatants of  E. cloacae  cultures and a functional T6SS-1 was required for the secretion of this protein. A  clpV2  mutant was impaired in both biofilm formation and adherence to epithelial cells, supporting the notion that these phenotypes are T6SS-2 dependent.  In vivo  data strongly suggest that both T6SSs are required for intestinal colonization because single and double mutants in  clpV1  and  clpV2  genes were defective in gut colonization in mice. We conclude that the two T6SSs are involved in the pathogenesis scheme of  E. cloacae  with specialized functions in the interaction with other bacteria and with host cells.

Two Type VI Secretion Systems of Enterobacter cloacae Are Required for Bacterial Competition, Cell Adherence, and Intestinal Colonization