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Broth stationary-phase cultures of most<jats:italic>dot\/icm<\/jats:italic>mutants are defective in entry and evasion of phagosome acidification. However, those virulence defects can be reversed by incubating broth cultures of<jats:italic>dot\/icm<\/jats:italic>mutants in water, termed water stress (WS). WS reversal requires the<jats:italic>lvh<\/jats:italic>T4ASS locus, suggesting an interaction between the two T4SSs in producing<jats:named-content xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" content-type=\"genus-species\" xlink:type=\"simple\">Legionella<\/jats:named-content>virulence phenotypes. In the current work, the loss of WS reversal in a<jats:italic>dotA<\/jats:italic>\u0394<jats:italic>lvh<\/jats:italic>mutant of strain JR32 was shown to be attributable to loss of the<jats:italic>lvh virD4<\/jats:italic>gene, encoding the putative coupling protein of the T4ASS. Transformation of a<jats:italic>dotA<\/jats:italic>\u0394<jats:italic>lvh<\/jats:italic>mutant with<jats:italic>virD4<\/jats:italic>also reversed entry and phagosome acidification defects in broth cultures. In addition, broth cultures of \u0394<jats:italic>lvh<\/jats:italic>and \u0394<jats:italic>virD4<\/jats:italic>mutants, which were<jats:italic>dot\/icm<\/jats:italic><jats:sup>+<\/jats:sup>, showed 5-fold and &gt;6-fold increases in translocation of the Dot\/Icm translocation substrates, proteins RalF and SidD, respectively. These data demonstrate that the Lvh T4ASS functions in both broth stationary-phase cultures conventionally used for infection and cultures exposed to WS treatment. Our studies in a<jats:italic>dotA<\/jats:italic>\u0394<jats:italic>lvh<\/jats:italic>mutant and in a<jats:italic>dot\/icm<\/jats:italic><jats:sup>+<\/jats:sup>background establish that VirD4 and the Lvh T4ASS contribute to virulence phenotypes and are consistent with independent functioning of Dot\/Icm and Lvh T4SSs or functional substitution of the Lvh VirD4 protein for a component(s) of the Dot\/Icm T4BSS.<\/jats:p>","DOI":"10.1128\/jb.00430-13","type":"journal-article","created":{"date-parts":[[2013,6,1]],"date-time":"2013-06-01T05:58:20Z","timestamp":1370066300000},"page":"3468-3475","update-policy":"https:\/\/doi.org\/10.1128\/asmj-crossmark-policy-page","source":"Crossref","is-referenced-by-count":18,"title":["Implication of the VirD4 Coupling Protein of the Lvh Type 4 Secretion System in Virulence Phenotypes of Legionella pneumophila"],"prefix":"10.1128","volume":"195","author":[{"given":"Purnima","family":"Bandyopadhyay","sequence":"first","affiliation":[{"name":"Department of Biochemistry, Albert Einstein College of Medicine, Bronx, New York, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Elza A. 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