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Efflux pumps from various superfamilies with broad substrate preferences sequester and pump drugs across the inner membrane to supply the highly polyspecific and powerful tripartite resistance\u2013nodulation\u2013cell division (RND) efflux pumps with compounds to be extruded across the outer membrane barrier. In\n                    <jats:italic>Escherichia coli<\/jats:italic>\n                    , the tripartite efflux system AcrAB\u2013TolC is the archetype RND multiple drug efflux pump complex. The homotrimeric inner membrane component acriflavine resistance B (AcrB) is the drug specificity and energy transduction center for the drug\/proton antiport process. Drugs are bound and expelled via a cycle of mainly three consecutive states in every protomer, constituting a flexible alternating access channel system. This review recapitulates the molecular basis of drug and inhibitor binding, including mechanistic insights into drug efflux by AcrB. It also summarizes 17 years of mutational analysis of the gene\n                    <jats:italic>acrB<\/jats:italic>\n                    , reporting the effect of every substitution on the ability of\n                    <jats:italic>E. coli<\/jats:italic>\n                    to confer resistance toward antibiotics (\n                    <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" xlink:href=\"http:\/\/goethe.link\/AcrBsubstitutions\">http:\/\/goethe.link\/AcrBsubstitutions<\/jats:ext-link>\n                    ). We emphasize the functional robustness of AcrB toward single\u2010site substitutions and highlight regions that are more sensitive to perturbation.\n                  <\/jats:p>","DOI":"10.1111\/nyas.14239","type":"journal-article","created":{"date-parts":[[2019,10,7]],"date-time":"2019-10-07T01:40:30Z","timestamp":1570412430000},"page":"38-68","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":175,"title":["AcrB: a mean, keen, drug efflux machine"],"prefix":"10.1111","volume":"1459","author":[{"given":"Jessica","family":"Kobylka","sequence":"first","affiliation":[{"name":"Institute of Biochemistry Goethe\u2010University Frankfurt  Frankfurt am Main Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Miriam S.","family":"Kuth","sequence":"additional","affiliation":[{"name":"Institute of Biochemistry Goethe\u2010University Frankfurt  Frankfurt am Main Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Reinke T.","family":"M\u00fcller","sequence":"additional","affiliation":[{"name":"Institute of Biochemistry Goethe\u2010University Frankfurt  Frankfurt am Main Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2789-5444","authenticated-orcid":false,"given":"Eric R.","family":"Geertsma","sequence":"additional","affiliation":[{"name":"Institute of Biochemistry Goethe\u2010University Frankfurt  Frankfurt am Main Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9035-3827","authenticated-orcid":false,"given":"Klaas M.","family":"Pos","sequence":"additional","affiliation":[{"name":"Institute of Biochemistry Goethe\u2010University Frankfurt  Frankfurt am Main Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2019,10,6]]},"reference":[{"key":"e_1_2_28_2_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature10388"},{"key":"e_1_2_28_3_1","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms13803"},{"key":"e_1_2_28_4_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.drup.2004.02.003"},{"key":"e_1_2_28_5_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature21377"},{"key":"e_1_2_28_6_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature22335"},{"key":"e_1_2_28_7_1","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a025197"},{"key":"e_1_2_28_8_1","volume-title":"Microbiology An Evolving Science","author":"Slonczewski 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