{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T04:43:51Z","timestamp":1787028231957,"version":"build-2736575974"},"reference-count":43,"publisher":"MDPI AG","issue":"3","license":[{"start":{"date-parts":[[2022,2,28]],"date-time":"2022-02-28T00:00:00Z","timestamp":1646006400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["31860035"],"award-info":[{"award-number":["31860035"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004761","name":"Natural Science Foundation of Hainan Province","doi-asserted-by":"publisher","award":["2019RC227"],"award-info":[{"award-number":["2019RC227"]}],"id":[{"id":"10.13039\/501100004761","id-type":"DOI","asserted-by":"publisher"}]},{"name":"College Student Innovation Project","award":["201911810006"],"award-info":[{"award-number":["201911810006"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Genes"],"abstract":"<jats:p>Yeasts play important roles in both the environment and in human welfare. While some environmental yeasts positively contribute to nutrient cycling and food production, a significant number of yeast species are opportunistic human pathogens, including several that are tolerant\/resistant to commonly used antifungal drugs. At present, most of our understanding of environmental yeasts has come from a few terrestrial environments in selected geographic regions. Relatively little is known about yeast diversity in tropical environments and their potential impacts on human health. Here, we characterize culturable yeasts in 968 environmental samples from eight regions in tropical China. Among the 516 soil, 273 freshwater, and 179 seawater samples, 71.5%, 85.7%, and 43.6% contained yeasts, respectively. A total of 984 yeast isolates were analyzed for their DNA barcode sequences and their susceptibilities to fluconazole. DNA sequence comparisons revealed that the 984 yeast isolates likely belonged to 144 species, including 106 known species and 38 putative novel species. About 38% of the 984 isolates belonged to known human pathogens and the most common species was Candida tropicalis, accounting for 21% (207\/984) of all isolates. Further analyses based on multi-locus sequence typing revealed that some of these environmental C. tropicalis shared identical genotypes with clinical isolates previously reported from tropical China and elsewhere. Importantly, 374 of the 984 (38%) yeast isolates showed intermediate susceptibility or resistance to fluconazole. Our results suggest that these environmental yeasts could have significant negative impacts on human health.<\/jats:p>","DOI":"10.3390\/genes13030444","type":"journal-article","created":{"date-parts":[[2022,2,28]],"date-time":"2022-02-28T20:09:57Z","timestamp":1646078997000},"page":"444","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":14,"title":["Extensive Diversity and Prevalent Fluconazole Resistance among Environmental Yeasts from Tropical China"],"prefix":"10.3390","volume":"13","author":[{"given":"Yiwei","family":"Liu","sequence":"first","affiliation":[{"name":"Public Laboratory, Hainan Medical University, Haikou 571199, China"},{"name":"Key Laboratory of Tropical Translational Medicine of Ministry of Education, Hainan Medical University, Haikou 571199, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhongyao","family":"Chen","sequence":"additional","affiliation":[{"name":"Public Laboratory, Hainan Medical University, Haikou 571199, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jingyuan","family":"Li","sequence":"additional","affiliation":[{"name":"Public Laboratory, Hainan Medical University, Haikou 571199, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhiqing","family":"Zhu","sequence":"additional","affiliation":[{"name":"Public Laboratory, Hainan Medical University, Haikou 571199, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sibei","family":"Pang","sequence":"additional","affiliation":[{"name":"Public Laboratory, Hainan Medical University, Haikou 571199, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2915-2780","authenticated-orcid":false,"given":"Jianping","family":"Xu","sequence":"additional","affiliation":[{"name":"Public Laboratory, Hainan Medical University, Haikou 571199, China"},{"name":"Department of Biology, McMaster University, Hamilton, ON L8S 4K1, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3045-1323","authenticated-orcid":false,"given":"Jinyan","family":"Wu","sequence":"additional","affiliation":[{"name":"Public Laboratory, Hainan Medical University, Haikou 571199, China"},{"name":"Key Laboratory of Tropical Translational Medicine of Ministry of Education, Hainan Medical University, Haikou 571199, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2022,2,28]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1016\/S0580-9517(01)30052-1","article-title":"Collection and identification of marine yeasts","volume":"30","author":"Fell","year":"2001","journal-title":"Methods Microbiol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"103098","DOI":"10.1016\/j.isci.2021.103098","article-title":"Global patterns in culturable soil yeast diversity","volume":"24","author":"Samarasinghe","year":"2021","journal-title":"IScience"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"91","DOI":"10.2166\/wh.2020.270","article-title":"Aquatic yeasts: Diversity, characteristics and potential health implications","volume":"18","author":"Monapathi","year":"2020","journal-title":"J. Water Health"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1002\/yea.1599","article-title":"Marine yeasts\u2014A review","volume":"25","author":"Kutty","year":"2008","journal-title":"Yeast"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"2101","DOI":"10.1111\/brv.12550","article-title":"Fungal evolution: Diversity, taxonomy and phylogeny of the Fungi","volume":"94","year":"2019","journal-title":"Biol. Rev. Camb. Philos. Soc."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"26","DOI":"10.1016\/j.ijfoodmicro.2016.07.018","article-title":"Yeast diversity of sourdoughs and associated metabolic properties and functionalities","volume":"239","author":"Vuyst","year":"2016","journal-title":"Int. J. Food Microbiol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"529","DOI":"10.1111\/1567-1364.12145","article-title":"Genome structure and dynamics of the yeast pathogen Candida glabrata","volume":"14","author":"Ahmad","year":"2014","journal-title":"FEMS Yeast Res."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1038\/nature02579","article-title":"Genome evolution in yeasts","volume":"430","author":"Dujon","year":"2004","journal-title":"Nature"},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Timmermans, B., De Las Pe\u00f1as, A., Casta\u00f1o, I., and Van Dijck, P. (2018). Adhesins in Candida glabrata. J. Fungi, 4.","DOI":"10.3390\/jof4020060"},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Douglass, A.P., Offei, B., Braun-Galleani, S., Coughlan, A.Y., Martos, A.A., Ortiz-Merino, R.A., Byrne, K.P., and Wolfe, K.H. (2018). Population genomics shows no distinction between pathogenic Candida krusei and environmental Pichia kudriavzevii: One species, four names. PLoS Pathog., 14.","DOI":"10.1371\/journal.ppat.1007138"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"e1007138","DOI":"10.1128\/mBio.03181-20","article-title":"Environmental isolation of Candida auris from the coastal wetlands of Andaman Islands, India","volume":"12","author":"Arora","year":"2021","journal-title":"MBio"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"2279","DOI":"10.2147\/IDR.S316368","article-title":"Species Diversity and Antifungal Susceptibilities of Oral Yeasts from Patients with Head and Neck Cancer","volume":"14","author":"Wu","year":"2021","journal-title":"Infect. Drug Resist."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1007\/s11046-014-9747-3","article-title":"Prevalent drug resistance among oral yeasts from asymptomatic patients in Hainan, China","volume":"177","author":"Wu","year":"2014","journal-title":"Mycopathologia"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Chowdhary, A., Kathuria, S., Xu, J., Sharma, C., Sundar, G., Singh, P.K., Gaur, S.N., Hagen, F., Klaassen, C.H., and Meis, J.F. (2012). Clonal expansion and emergence of environmental multiple-triazole-resistant Aspergillus fumigatus strains carrying the TR34\/L98H mutations in the cyp51A gene in India. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0052871"},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Chowdhary, A., Kathuria, S., Xu, J., and Meis, J.F. (2013). Emergence of azole resistant Aspergillus fumigatus strains due to agricultural azole use creates an increasing threat to human health. PLoS Pathog., 9.","DOI":"10.1371\/annotation\/4ffcf1da-b180-4149-834c-9c723c5dbf9b"},{"key":"ref_16","unstructured":"National Bureau of Statistics, PRC (2020). China Statistical Yearbook."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1452","DOI":"10.1073\/pnas.85.5.1452","article-title":"A Candida albicans dispersed, repeated gene family and its epidemiologic applications","volume":"85","author":"Scherer","year":"1988","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"fiz122","DOI":"10.1093\/femsec\/fiz122","article-title":"Fantastic yeasts and where to find them: The discovery of a predominantly clonal Cryptococcus deneoformans population in Saudi Arabian soils","volume":"95","author":"Samarasinghe","year":"2019","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1016\/j.simyco.2016.11.007","article-title":"DNA barcoding analysis of more than 9000 yeast isolates contributes to quantitative thresholds for yeast species and genera delimitation","volume":"85","author":"Vu","year":"2016","journal-title":"Stud. Mycol."},{"key":"ref_20","unstructured":"Institute Cals (2009). M44-A2 Reference Method for Antifungaldisk Diffusion Susceptibility Testing of Yeasts, Clinical and Laboratory Standards Institute. [2nd ed.]."},{"key":"ref_21","unstructured":"(2011). NEO-SENSITABS User\u2019s Guide in Antifungal Susceptibility Testing of Yeast, Rosco Diagno\u2019stica A\/S."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"42537","DOI":"10.1038\/srep42537","article-title":"Multilocus sequence analyses reveal extensive diversity and multiple origins of fluconazole resistance in Candida tropicalis from tropical China","volume":"7","author":"Wu","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"2725","DOI":"10.1093\/molbev\/mst197","article-title":"MEGA6: Molecular evolutionary genetics analysis version 6.0","volume":"30","author":"Tamura","year":"2013","journal-title":"Mol. Biol. Evol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"2537","DOI":"10.1093\/bioinformatics\/bts460","article-title":"GenAlEx 6.5: Genetic analysis in Excel. Population genetic software for teaching and research\u2014An update","volume":"28","author":"Peakall","year":"2012","journal-title":"Bioinformatics"},{"key":"ref_25","first-page":"5","article-title":"Package \u201cVegan\u201d","volume":"2","author":"Oksanen","year":"2020","journal-title":"Community Ecol. Package Version"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1312","DOI":"10.1093\/bioinformatics\/btu033","article-title":"RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies","volume":"30","author":"Stamatakis","year":"2014","journal-title":"Bioinformatics"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"foz032","DOI":"10.1093\/femsyr\/foz032","article-title":"Pathogenic budding yeasts isolated outside of clinical settings","volume":"19","author":"Opulente","year":"2019","journal-title":"FEMS Yeast Res."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"707","DOI":"10.3389\/fmicb.2018.00707","article-title":"Fungal biodiversity and their role in soil health","volume":"9","author":"Hannula","year":"2018","journal-title":"Front. Microbiol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"436","DOI":"10.1007\/s12257-018-0477-0","article-title":"Invasive fungal infections and their epidemiology: Measures in the clinical scenario","volume":"24","author":"Bajpai","year":"2019","journal-title":"Biotechnol. Bioproc. E"},{"key":"ref_30","unstructured":"Castano, G., Yarrarapu, S.N., and Mada, P.K. (2021). Trichosporonosis. StatPearls, StatPearls Publishing."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"e180566","DOI":"10.1590\/0074-02760180566","article-title":"Occurrence, antifungal susceptibility, and virulence factors of opportunistic yeasts isolated from Brazilian beaches","volume":"114","author":"Maciel","year":"2019","journal-title":"Mem. Inst. Oswaldo. Cruz."},{"key":"ref_32","first-page":"e62816","article-title":"Frequency of uncommon clinical yeast species confirmed by ITS-Sequencing","volume":"14","author":"Ruosta","year":"2019","journal-title":"Arch. Clin. Infect. Dis."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"91","DOI":"10.5578\/mb.20188","article-title":"Nadir Etken Bir Maya: \u00dc\u00e7\u00fcnc\u00fc Basamak \u00dcniversite Hastanesinde \u0130zlenen Rhodotorula mucilaginosa Enfeksiyonu Olgular\u0131","volume":"55","author":"Ayaz","year":"2021","journal-title":"Mikrobiyol. Bul."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"O83","DOI":"10.1111\/1469-0691.12337","article-title":"Epidemiological study of a large cluster of fungaemia cases due to Kodamaea ohmeri in an Indian tertiary care centre","volume":"20","author":"Chakrabarti","year":"2014","journal-title":"Clin. Microbiol. Infect."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"866","DOI":"10.1017\/ice.2015.103","article-title":"Infection control challenges of infrequent and rare fungal pathogens: Lessons from disseminated Fusarium and Kodamaea ohmeri infections","volume":"36","author":"Das","year":"2015","journal-title":"Infect. Control. Hosp. Epidemiol."},{"key":"ref_36","first-page":"21","article-title":"Molecular identification and antifungal susceptibility pattern of Candida species isolated from HIV infected Patients with candidiasis","volume":"5","author":"Paul","year":"2019","journal-title":"Curr. Med. Mycol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"636","DOI":"10.1111\/myc.13094","article-title":"Kodamaea ohmeri infections in humans: A systematic review","volume":"63","author":"Ioannou","year":"2020","journal-title":"Mycoses"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1111\/myc.12875","article-title":"Rhodotorula infection in haematological patient: Risk factors and outcome","volume":"62","author":"Potenza","year":"2019","journal-title":"Mycoses"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"792","DOI":"10.1016\/j.cmi.2019.03.028","article-title":"Resistance of Candida to azoles and echinocandins worldwide","volume":"25","author":"Pristov","year":"2019","journal-title":"Clin. Microbiol. Infect."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1819","DOI":"10.1590\/0001-3765201520130418","article-title":"Prevalence of Candida tropicalis and Candida krusei in onychomycosis in Jo\u00e3o Pessoa, Paraiba, Brazil from 1999 to 2010","volume":"87","author":"Arrua","year":"2015","journal-title":"An. Acad. Bras. Cienc."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"659","DOI":"10.1080\/21505594.2018.1438026","article-title":"A fungus among us: The emerging opportunistic pathogen Candida tropicalis and PKA signaling","volume":"9","author":"Kumar","year":"2018","journal-title":"Virulence"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"900","DOI":"10.3389\/fmicb.2019.00900","article-title":"Analyses of the global multilocus genotypes of the human pathogenic yeast Candida tropicalis","volume":"10","author":"Wu","year":"2019","journal-title":"Front. Microbiol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1139\/gen-2020-0022","article-title":"Fungal species concepts in the genomics era","volume":"63","author":"Xu","year":"2020","journal-title":"Genome"}],"container-title":["Genes"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4425\/13\/3\/444\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T22:29:16Z","timestamp":1760135356000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4425\/13\/3\/444"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,2,28]]},"references-count":43,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2022,3]]}},"alternative-id":["genes13030444"],"URL":"https:\/\/doi.org\/10.3390\/genes13030444","relation":{},"ISSN":["2073-4425"],"issn-type":[{"value":"2073-4425","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,2,28]]}}}