{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,16]],"date-time":"2026-09-16T18:05:33Z","timestamp":1789581933526,"version":"build-2803163510"},"reference-count":113,"publisher":"American Association for the Advancement of Science (AAAS)","issue":"867","funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01 HL128156"],"award-info":[{"award-number":["R01 HL128156"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01 HL143998"],"award-info":[{"award-number":["R01 HL143998"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.science.org"],"crossmark-restriction":true},"short-container-title":["Sci. Transl. Med."],"published-print":{"date-parts":[[2026,9,16]]},"abstract":"<jats:p>\n                    Nearly all known cases of antimicrobial resistance have been linked to agents that directly target microbes. This paradigm overlooks other drivers of microbial evolution and resistance. Here, we report compelling evidence of resistance-associated mutations in\n                    <jats:italic toggle=\"yes\">Pneumocystis jirovecii<\/jats:italic>\n                    , a high-priority fungal pathogen designated by WHO as having unknown resistance. These mutations emerged under selective pressure from mycophenolic acid (MPA), a widely used immunosuppressant not used for\n                    <jats:italic toggle=\"yes\">Pneumocystis<\/jats:italic>\n                    pneumonia (PCP) treatment. A potent inhibitor of inosine monophosphate dehydrogenase (IMPDH), MPA has been linked to altered\n                    <jats:italic toggle=\"yes\">P. jirovecii<\/jats:italic>\n                    susceptibility in solid organ transplant (SOT) recipients. Genetic analysis of 163\n                    <jats:italic toggle=\"yes\">P. jirovecii<\/jats:italic>\n                    samples from PCP outbreaks in SOT recipients and nontransplant controls across six countries identified six\n                    <jats:italic toggle=\"yes\">impdh<\/jats:italic>\n                    mutations, four of which were previously unreported. These mutations showed strong associations with prior MPA exposure, geographic segregation, and temporal shifts. They arose independently in 11 distinct strains worldwide between 2005 and 2019, with evidence of switching within and between strains, suggesting ongoing adaptive evolution. Structural modeling predicted that these mutations reduced IMPDH stability and MPA binding; accordingly, all mutations conferred MPA resistance in enzyme inhibition assays. Our findings uncover the unanticipated global emergence of multiple resistance-associated mutations in\n                    <jats:italic toggle=\"yes\">P. jirovecii<\/jats:italic>\n                    driven by a human-targeted immunosuppressant, likely conferring a survival advantage in SOT recipients and facilitating transmission and outbreaks. These results challenge the current status quo on\n                    <jats:italic toggle=\"yes\">P. jirovecii<\/jats:italic>\n                    \u2019s unknown resistance, calling for reevaluating immunosuppressive strategies and infection control measures and highlighting the broader potential for other pathogens to evolve under selective pressures imposed by human-targeted therapies.\n                  <\/jats:p>","DOI":"10.1126\/scitranslmed.adz4471","type":"journal-article","created":{"date-parts":[[2026,9,16]],"date-time":"2026-09-16T17:58:11Z","timestamp":1789581491000},"update-policy":"https:\/\/doi.org\/10.34133\/aaas_crossmark","source":"Crossref","is-referenced-by-count":0,"title":["Unanticipated global emergence and evolution of\n                    <i>Pneumocystis jirovecii<\/i>\n                    mutations and resistance driven by a human-targeted drug"],"prefix":"10.1126","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6078-5730","authenticated-orcid":true,"given":"Liang","family":"Ma","sequence":"first","affiliation":[{"name":"Critical Care Medicine Department, National Institutes of Health Clinical Center, Bethesda, MD 20892, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ming","family":"Hao","sequence":"additional","affiliation":[{"name":"Laboratory of Human Retrovirology and Immunoinformatics, Frederick National Laboratory for Cancer Research, Frederick, MD 21702, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1413-2763","authenticated-orcid":true,"given":"Weizhong","family":"Chang","sequence":"additional","affiliation":[{"name":"Laboratory of Human Retrovirology and Immunoinformatics, Frederick National Laboratory for Cancer Research, Frederick, MD 21702, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-0197-5295","authenticated-orcid":true,"given":"Akanchhya","family":"Khanal","sequence":"additional","affiliation":[{"name":"Graduate Program in Biotechnology, Brandeis University, Waltham, MA 02453, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Steven M.","family":"Ewell","sequence":"additional","affiliation":[{"name":"Graduate Program in Biochemistry and Biophysics, Brandeis University, Waltham, MA 02453, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1085-5155","authenticated-orcid":true,"given":"Xilong","family":"Deng","sequence":"additional","affiliation":[{"name":"Critical Care Medicine Department, National Institutes of Health Clinical Center, Bethesda, MD 20892, USA."},{"name":"Department of Critical Care Medicine, Guangzhou Eighth People\u2019s Hospital, Guangzhou Medical University, Guangzhou 510030, Guangdong Province, China."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Junfeng","family":"Sun","sequence":"additional","affiliation":[{"name":"Critical Care Medicine Department, National Institutes of Health Clinical Center, Bethesda, MD 20892, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marwan M.","family":"Azar","sequence":"additional","affiliation":[{"name":"Department of Internal Medicine, Section of Infectious Diseases, Yale School of Medicine, and US Medical Affairs, GlaxoSmithKline Pharmaceuticals, New Haven, CT 06510, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5086-0293","authenticated-orcid":true,"given":"Alexia","family":"Cusini","sequence":"additional","affiliation":[{"name":"Department of Infectious Diseases, Inselspital, Bern University Hospital, University of Bern, 3010 Bern, Switzerland."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1668-1800","authenticated-orcid":true,"given":"Thomas","family":"Fehr","sequence":"additional","affiliation":[{"name":"Department of Internal Medicine, Cantonal Hospital Graubuenden, Chur 7000, Switzerland."},{"name":"Division of Nephrology, University Hospital Zurich, University of Zurich, 8091 Zurich, Switzerland."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9927-0849","authenticated-orcid":true,"given":"Sara","family":"Gianella","sequence":"additional","affiliation":[{"name":"Infectious Diseases and Global Public Health, University of California San Diego, La Jolla, CA 92037, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4931-3203","authenticated-orcid":true,"given":"Norihiko","family":"Goto","sequence":"additional","affiliation":[{"name":"Transplant Nephrology, Goto 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