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Sequence and structural analysis of T6SS reveals a resemblance to the T4 bacteriophage tail, in which an outer sheath structure contracts an internal tube for injecting nucleic acid into bacterial cells. However, the molecular details of how this phage tail-like T6SS structure is assembled in vivo and executed for exoprotein or effector secretion remain largely unknown.
Here, we used a systematic approach to identify T6SS machinery and secreted components and investigate the interaction among the putative sheath and tube components of Agrobacterium tumefaciens. Interestingly, Atu and the putative tube components, Hcp and VgrG, are mainly localized in the cytoplasm but also detected on the bacterial surface. Further co-immunoprecipitation and pulldown assays revealed these subcomplex es in A.
Protein secretion systems play central roles in export or import of macromolecules across the cell envelope in bacteria. T6SS is tightly regulated and has a variety of biological functions such as promoting or limiting virulence and cytotoxicity in eukaryotic or bacterial hosts  — . A unique feature of Hcp and VgrG is that they are both secreted exoproteins but are also part of a secretion apparatus  ,  — .
Growing evidence of structural and sequence analogy of several T6SS components to the T4 bacteriophage tail baseplate, tail tube and sheath suggests that the T6SS machinery and phage tail are evolutionarily conserved. Hcp forms a hexameric ring structure with a 4. In addition, VgrG structurally resembles the phage tail spike gp5 3 — gp27 3 complex, and Hcp possesses sequence homology to the phage tube gp19 and is structurally similar to gp5 and the tandem tube domain of gp27  ,  , .
Thus, the Hcp and VgrG exoproteins may assemble into a phage tail-like structure. Furthermore, Mekalanos and colleagues discovered that TssB—TssC tubule structures are highly dynamic and cycle between an extended and contracted conformation  , a phenomenon also recently reported by other groups  , .
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