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Subsequent analyses included\n                      <jats:italic>ERG11<\/jats:italic>\n                      alignment to determine the extent and distribution of missense substitutions, Ka\/Ks calculations to test for positive selection on\n                      <jats:italic>ERG11<\/jats:italic>\n                      , and detailed\n                      <jats:italic>CDR1<\/jats:italic>\n                      and\n                      <jats:italic>CDR2<\/jats:italic>\n                      mutational analysis across the coding sequence. Efflux function was assessed by measuring the fold reduction in the minimum inhibitory concentration (MIC) of azole drugs in the presence of milbemycin.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      A total of 27 azole-resistant\n                      <jats:italic>C. albicans<\/jats:italic>\n                      isolates from ICU patients with\n                      <jats:italic>Candida<\/jats:italic>\n                      BSIs in Linkou Chang Gung Memorial Hospital in Taiwan and HIV patients\u2019 oropharyngeal colonization were identified and analyzed. The in-depth, core analyses were performed on seven representative\n                      <jats:italic>C. albicans<\/jats:italic>\n                      isolates. The\n                      <jats:italic>C. albicans<\/jats:italic>\n                      isolates of HIV-infected patients had notably higher azole resistance (fluconazole MICs\u2009&gt;\u2009256\u00a0mg\/L) when compared with those of the ICU patients (fluconazole MICs 16\u201364\u00a0mg\/L), suggesting the involvement of additional mechanisms. The central role of\n                      <jats:italic>ERG11<\/jats:italic>\n                      mutations was supported by the presence of\n                      <jats:italic>ERG11<\/jats:italic>\n                      missense mutations in all azole-resistant\n                      <jats:italic>C. albicans<\/jats:italic>\n                      isolates. The\n                      <jats:italic>ERG11<\/jats:italic>\n                      coding regions analyses showed no evidence of positive selection (Ka\/Ks\u2009&lt;\u20091.0) and no specific mutation unique to the\n                      <jats:italic>C. albicans<\/jats:italic>\n                      isolates of the HIV patients, but unique missense mutations only in\n                      <jats:italic>CDR1<\/jats:italic>\n                      \/\n                      <jats:italic>CDR2<\/jats:italic>\n                      were noted. Milbemycin substantially decreased the azole MICs in all\n                      <jats:italic>C. albicans<\/jats:italic>\n                      isolates and confirmed the efflux pump involvement. The effects of milbemycin were much lower in\n                      <jats:italic>C. albicans<\/jats:italic>\n                      of the HIV patients than those from the ICU patients. Furthermore, unique\n                      <jats:italic>ERG11<\/jats:italic>\n                      promoter variants, including changes at the Hap43p\/Hap5p sites, were noted in the\n                      <jats:italic>C. albicans<\/jats:italic>\n                      isolates of the HIV patients.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>\n                      While\n                      <jats:italic>ERG11<\/jats:italic>\n                      structural mutations are foundational, the elevated azole resistance in the\n                      <jats:italic>C. albicans<\/jats:italic>\n                      isolates of HIV-infected patients is associated with specific\n                      <jats:italic>CDR1<\/jats:italic>\n                      \/\n                      <jats:italic>CDR2<\/jats:italic>\n                      mutations and distinct\n                      <jats:italic>ERG11<\/jats:italic>\n                      promoter variants, highlighting genomic features that warrant further functional validation.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1186\/s12929-026-01231-4","type":"journal-article","created":{"date-parts":[[2026,2,22]],"date-time":"2026-02-22T07:41:36Z","timestamp":1771746096000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Genetic variation and mutational determinants of azole resistance in Candida albicans strains of oropharyngeal colonization in HIV patients and bloodstream infections"],"prefix":"10.1186","volume":"33","author":[{"given":"Ming-Horng","family":"Tsai","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chih-Hung","family":"Hsieh","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"I.-An","family":"Tsai","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ching-Min","family":"Chang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ting-Wen","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jen-Fu","family":"Hsu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shih-Ming","family":"Chu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hsuan-Rong","family":"Huang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shao-Hung","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jang-Jih","family":"Lu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,2,22]]},"reference":[{"key":"1231_CR1","doi-asserted-by":"publisher","DOI":"10.1128\/spectrum.02453-22","volume":"10","author":"P Badiee","year":"2022","unstructured":"Badiee P, Boekhout T, Haddadi P, Mohammadi R, Ghadimi-Moghadam A, Soltani J, et al. 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