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The pathology associated with infection depends on bacterial delivery of effector proteins into the host via the membrane spanning Dot\/Icm type IV secretion system (T4SS). We have determined sub-3.0 \u00c5 resolution maps of the Dot\/Icm T4SS core complex by single particle cryo-EM. The high-resolution structural analysis has allowed us to identify proteins encoded outside the Dot\/Icm genetic locus that contribute to the core T4SS structure. We can also now define two distinct areas of symmetry mismatch, one that connects the C18 periplasmic ring (PR) and the C13 outer membrane cap (OMC) and one that connects the C13 OMC with a 16-fold symmetric dome. Unexpectedly, the connection between the PR and OMC is DotH, with five copies sandwiched between the OMC and PR to accommodate the symmetry mismatch. Finally, we observe multiple conformations in the reconstructions that indicate flexibility within the structure.\n                  <\/jats:p>","DOI":"10.7554\/elife.70427","type":"journal-article","created":{"date-parts":[[2021,9,14]],"date-time":"2021-09-14T08:00:33Z","timestamp":1631606433000},"update-policy":"https:\/\/doi.org\/10.7554\/elife.70427","source":"Crossref","is-referenced-by-count":44,"title":["Cryo-EM reveals new species-specific proteins and symmetry elements in the Legionella pneumophila Dot\/Icm T4SS"],"prefix":"10.7554","volume":"10","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3185-1727","authenticated-orcid":true,"given":"Michael J","family":"Sheedlo","sequence":"first","affiliation":[{"name":"Department of Pharmacology, University of Minnesota","place":["Minneapolis, United States"]},{"name":"Department of Pathology, Microbiology, and Immunology, Vanderbilt University Medical Center","place":["Nashville, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4027-4386","authenticated-orcid":true,"given":"Clarissa L","family":"Durie","sequence":"first","affiliation":[{"name":"Life Sciences Institute, University of Michigan","place":["Ann Arbor, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4285-8764","authenticated-orcid":true,"given":"Jeong Min","family":"Chung","sequence":"additional","affiliation":[{"name":"Life Sciences Institute, University of Michigan","place":["Ann Arbor, United States"]},{"name":"Department of Biotechnology, The Catholic University of Korea","place":["Gyeonggi, Republic of Korea"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Louise","family":"Chang","sequence":"additional","affiliation":[{"name":"Life Sciences Institute, University of Michigan","place":["Ann Arbor, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jacquelyn","family":"Roberts","sequence":"additional","affiliation":[{"name":"Life Sciences Institute, University of Michigan","place":["Ann Arbor, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2542-0266","authenticated-orcid":true,"given":"Michele","family":"Swanson","sequence":"additional","affiliation":[{"name":"Department of Microbiology and Immunology, University Of Michigan","place":["Ann Arbor, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2273-8121","authenticated-orcid":true,"given":"Dana Borden","family":"Lacy","sequence":"additional","affiliation":[{"name":"Department of Pathology, Microbiology, and Immunology, Vanderbilt University Medical Center","place":["Nashville, United States"]},{"name":"The Veterans Affairs Tennessee Valley Healthcare System","place":["Nasvhille, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1750-4793","authenticated-orcid":true,"given":"Melanie D","family":"Ohi","sequence":"additional","affiliation":[{"name":"Life Sciences Institute, University of Michigan","place":["Ann Arbor, United States"]},{"name":"Department of Cell and Developmental Biology, University of Michigan","place":["Ann Arbor, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"4374","published-online":{"date-parts":[[2021,9,14]]},"reference":[{"key":"bib1","doi-asserted-by":"publisher","first-page":"531","DOI":"10.1107\/S2059798318006551","article-title":"Real-space refinement in PHENIX for cryo-em and crystallography","volume":"74","author":"Afonine","year":"2018","journal-title":"Acta Crystallographica. 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