{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,17]],"date-time":"2026-10-17T02:39:31Z","timestamp":1792204771279,"version":"4.4.0"},"reference-count":51,"publisher":"MDPI AG","issue":"10","license":[{"start":{"date-parts":[[2022,5,17]],"date-time":"2022-05-17T00:00:00Z","timestamp":1652745600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"NSFC","doi-asserted-by":"publisher","award":["31770123"],"award-info":[{"award-number":["31770123"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"NSFC","doi-asserted-by":"publisher","award":["RSP-2022\/205"],"award-info":[{"award-number":["RSP-2022\/205"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"name":"King Saud University, Riyadh, Saudi Arabia","award":["31770123"],"award-info":[{"award-number":["31770123"]}]},{"name":"King Saud University, Riyadh, Saudi Arabia","award":["RSP-2022\/205"],"award-info":[{"award-number":["RSP-2022\/205"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Arsenic (As), distributed widely in the natural environment, is a toxic substance which can severely impair the normal functions in living cells. Research on the genetic determinants conferring functions in arsenic resistance and metabolism is of great importance for remediating arsenic-contaminated environments. Many organisms, including bacteria, have developed various strategies to tolerate arsenic, by either detoxifying this harmful element or utilizing it for energy generation. More and more new arsenic resistance (ars) determinants have been identified to be conferring resistance to diverse arsenic compounds and encoded in ars operons. There is a hazard in mobilizing arsenic during gold-mining activities due to gold- and arsenic-bearing minerals coexisting. In this study, we isolated 8 gold enrichment strains from the Zijin gold and copper mine (Longyan, Fujian Province, China) wastewater treatment site soil, at an altitude of 192 m. We identified two Brevundimonas nasdae strains, Au-Bre29 and Au-Bre30, among these eight strains, having a high minimum inhibitory concentration (MIC) for As(III). These two strains contained the same ars operons but displayed differences regarding secretion of extra-polymeric substances (EPS) upon arsenite (As(III)) stress. B. nasdae Au-Bre29 contained one extra plasmid but without harboring any additional ars genes compared to B. nasdae Au-Bre30. We optimized the growth conditions for strains Au-Bre29 and Au-Bre30. Au-Bre30 was able to tolerate both a lower pH and slightly higher concentrations of NaCl. We also identified folE, a folate synthesis gene, in the ars operon of these two strains. In most organisms, folate synthesis begins with a FolE (GTP-Cyclohydrolase I)-type enzyme, and the corresponding gene is typically designated folE (in bacteria) or gch1 (in mammals). Heterologous expression of folE, cloned from B. nasdae Au-Bre30, in the arsenic-hypersensitive strain Escherichia coli AW3110, conferred resistance to As(III), arsenate (As(V)), trivalent roxarsone (Rox(III)), pentavalent roxarsone (Rox(V)), trivalent antimonite (Sb(III)), and pentavalent antimonate (Sb(V)), indicating that folate biosynthesis is a target of arsenite toxicity and increased production of folate confers increased resistance to oxyanions. Genes encoding Acr3 and ArsH were shown to confer resistance to As(III), Rox(III), Sb(III), and Sb(V), and ArsH also conferred resistance to As(V). Acr3 did not confer resistance to As(V) and Rox(V), while ArsH did not confer resistance to Rox(V).<\/jats:p>","DOI":"10.3390\/ijms23105619","type":"journal-article","created":{"date-parts":[[2022,5,18]],"date-time":"2022-05-18T03:20:43Z","timestamp":1652844043000},"page":"5619","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":17,"title":["Characterization of Two Highly Arsenic-Resistant Caulobacteraceae Strains of Brevundimonas nasdae: Discovery of a New Arsenic Resistance Determinant"],"prefix":"10.3390","volume":"23","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7617-9751","authenticated-orcid":false,"given":"Xiaojun","family":"Yang","sequence":"first","affiliation":[{"name":"College of Life Sciences, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuanping","family":"Li","sequence":"additional","affiliation":[{"name":"Institute of Environmental Microbiology, College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Renwei","family":"Feng","sequence":"additional","affiliation":[{"name":"Institute of Environmental Microbiology, College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jian","family":"Chen","sequence":"additional","affiliation":[{"name":"Department of Cellular Biology and Pharmacology, Herbert Wertheim College of Medicine, Florida International University, Miami, FL 33199, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hend A.","family":"Alwathnani","sequence":"additional","affiliation":[{"name":"Department of Botany and Microbiology, College of Science, King Saud University, Riyadh 11451, Saudi Arabia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Weifeng","family":"Xu","sequence":"additional","affiliation":[{"name":"Joint International Research Laboratory of Water and Nutrient in Crop, College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christopher","family":"Rensing","sequence":"additional","affiliation":[{"name":"Institute of Environmental Microbiology, College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2022,5,17]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1016\/j.envint.2008.10.004","article-title":"Arsenic in groundwater: A threat to sustainable agriculture in South and South-east Asia","volume":"35","author":"Brammer","year":"2009","journal-title":"Environ. Int."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1016\/S0378-4274(02)00085-1","article-title":"Epidemiologic evidence of diabetogenic effect of arsenic","volume":"133","author":"Tseng","year":"2002","journal-title":"Toxicol. Lett."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"888","DOI":"10.1038\/bjc.1992.380","article-title":"Cancer potential in liver, lung, bladder and kidney due to ingested inorganic arsenic in drinking water","volume":"66","author":"Chen","year":"1992","journal-title":"Br. J. Cancer"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1146\/annurev-earth-060313-054942","article-title":"Earth Abides Arsenic Biotransformations","volume":"42","author":"Zhu","year":"2014","journal-title":"Annu. Rev. Earth Planet. Sci."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1146\/annurev.micro.60.080805.142053","article-title":"Arsenic and selenium in microbial metabolism","volume":"60","author":"Stolz","year":"2006","journal-title":"Annu. Rev. Microbiol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"730","DOI":"10.1007\/s002530051457","article-title":"Microbial heavy-metal resistance","volume":"51","author":"Nies","year":"1999","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1016\/S0012-8252(01)00070-8","article-title":"Antimony in the environment: A review focused on natural waters: I. Occurrence","volume":"57","author":"Chen","year":"2002","journal-title":"Earth Sci. Rev."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"43","DOI":"10.3389\/fenvs.2020.00043","article-title":"The Arsenic Methylation Cycle: How Microbial Communities Adapted Methylarsenicals for Use as Weapons in the Continuing War for Dominance","volume":"8","author":"Chen","year":"2020","journal-title":"Front. Environ. Sci."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"5482","DOI":"10.1128\/AEM.01375-16","article-title":"Microbial Antimony Biogeochemistry: Enzymes, Regulation, and Related Metabolic Pathways","volume":"82","author":"Li","year":"2016","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1016\/B978-0-12-394390-3.00012-4","article-title":"Pathways of arsenic uptake and efflux","volume":"69","author":"Yang","year":"2012","journal-title":"Curr. Top. Membr."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"e00246-19","DOI":"10.1128\/mBio.00246-19","article-title":"A Parasitic Arsenic Cycle That Shuttles Energy from Phytoplankton to Heterotrophic Bacterioplankton","volume":"10","author":"Giovannoni","year":"2019","journal-title":"Mbio"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"6182","DOI":"10.1021\/acs.est.9b01187","article-title":"Role of ArsEFG in Roxarsone and Nitarsone Detoxification and Resistance","volume":"53","author":"Chen","year":"2019","journal-title":"Environ. Sci. Technol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"e00502-21","DOI":"10.1128\/Spectrum.00502-21","article-title":"Identification of the Biosynthetic Gene Cluster for the Organoarsenical Antibiotic Arsinothricin","volume":"9","author":"Paul","year":"2021","journal-title":"Microbiol. Spectr."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"7550","DOI":"10.1111\/1462-2920.15817","article-title":"ArsV and ArsW provide synergistic resistance to the antibiotic methylarsenite","volume":"23","author":"Chen","year":"2021","journal-title":"Environ. Microbiol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"2075","DOI":"10.1073\/pnas.0506836103","article-title":"Arsenic detoxification and evolution of trimethylarsine gas by a microbial arsenite S-adenosylmethionine methyltransferase","volume":"103","author":"Qin","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1111\/j.1574-6941.2008.00613.x","article-title":"Identification of genes conferring arsenic resistance to Escherichia coli from an effluent treatment plant sludge metagenomic library","volume":"67","author":"Chauhan","year":"2009","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"762","DOI":"10.1111\/1462-2920.15599","article-title":"Organoarsenical tolerance in Sphingobacterium wenxiniae, a bacterium isolated from activated sludge","volume":"24","author":"Chen","year":"2022","journal-title":"Environ. Microbiol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1042","DOI":"10.1111\/mmi.12988","article-title":"ArsH is an organoarsenical oxidase that confers resistance to trivalent forms of the herbicide monosodium methylarsenate and the poultry growth promoter roxarsone","volume":"96","author":"Chen","year":"2015","journal-title":"Mol. Microbiol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"945","DOI":"10.1111\/mmi.13371","article-title":"Synergistic interaction of glyceraldehydes-3-phosphate dehydrogenase and ArsJ, a novel organoarsenical efflux permease, confers arsenate resistance","volume":"100","author":"Chen","year":"2016","journal-title":"Mol. Microbiol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"e01842-18","DOI":"10.1128\/AEM.01842-18","article-title":"Efflux Transporter ArsK Is Responsible for Bacterial Resistance to Arsenite, Antimonite, Trivalent Roxarsone, and Methylarsenite","volume":"84","author":"Shi","year":"2018","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"625","DOI":"10.1111\/mmi.13145","article-title":"ArsP: A methylarsenite efflux permease","volume":"98","author":"Chen","year":"2015","journal-title":"Mol. Microbiol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1289\/ehp.02110s5745","article-title":"Arsenate reductases in prokaryotes and eukaryotes","volume":"110","author":"Mukhopadhyay","year":"2002","journal-title":"Environ. Health Perspect."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"3534","DOI":"10.1128\/JB.01798-08","article-title":"The glutathione\/glutaredoxin system is essential for arsenate reduction in Synechocystis sp. strain PCC 6803","volume":"191","author":"Reyes","year":"2009","journal-title":"J. Bacteriol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"3738","DOI":"10.1128\/AEM.72.5.3738-3742.2006","article-title":"arsRBOCT arsenic resistance system encoded by linear plasmid pHZ227 in Streptomyces sp. strain FR-008","volume":"72","author":"Wang","year":"2006","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"794315","DOI":"10.3389\/fmicb.2021.794315","article-title":"Complete Genome Sequencing and Comparative Genomics of Three Potential Probiotic Strains, Lacticaseibacillus casei FBL6, Lacticaseibacillus chiayiensis FBL7, and Lacticaseibacillus zeae FBL8","volume":"12","author":"Kim","year":"2022","journal-title":"Front. Microbiol."},{"key":"ref_26","unstructured":"Su, J. (2020). Isolation, Identification and Arsenic Resistant Mechanism Analysis of an Arsenic Tolerant Bacteria Paenibacillus taichungensis NC1 (In Chinese). [Master Thesis, Fujian Agriculture and Forestry University]."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1016\/j.jes.2016.10.004","article-title":"Comparative cytotoxicity of fourteen trivalent and pentavalent arsenic species determined using real-time cell sensing","volume":"49","author":"Moe","year":"2016","journal-title":"J. Environ. Sci."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1169","DOI":"10.1016\/j.envpol.2009.10.045","article-title":"The chemistry and behaviour of antimony in the soil environment with comparisons to arsenic: A critical review","volume":"158","author":"Wilson","year":"2010","journal-title":"Environ. Pollut."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"2230","DOI":"10.1111\/1462-2920.14991","article-title":"ArsH protects Pseudomonas putida from oxidative damage caused by exposure to arsenic","volume":"22","author":"Nikel","year":"2020","journal-title":"Environ. Microbiol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"141174","DOI":"10.1016\/j.scitotenv.2020.141174","article-title":"Comprehensive genomic and proteomic profiling reveal Acinetobacter johnsonii JH7 responses to Sb(III) toxicity","volume":"748","author":"Gu","year":"2020","journal-title":"Sci. Total Environ."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"133453","DOI":"10.1016\/j.chemosphere.2021.133453","article-title":"Sb(III) resistance mechanism and oxidation characteristics of Klebsiella aerogenes X","volume":"293","author":"Rong","year":"2022","journal-title":"Chemosphere"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"705","DOI":"10.1006\/bbrc.1995.2026","article-title":"Homology cloning of GTP-cyclohydrolase I from various unrelated eukaryotes by reverse-transcription polymerase chain reaction using a general set of degenerate primers","volume":"212","author":"Maier","year":"1995","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"51534","DOI":"10.1074\/jbc.M409440200","article-title":"Structural basis of biopterin-induced inhibition of GTP cyclohydrolase I by GFRP, its feedback regulatory protein","volume":"279","author":"Maita","year":"2004","journal-title":"J. Biol. Chem."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"13567","DOI":"10.1073\/pnas.240463497","article-title":"Zinc plays a key role in human and bacterial GTP cyclohydrolase I","volume":"97","author":"Auerbach","year":"2000","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"449","DOI":"10.2135\/cropsci2005.0449","article-title":"Folate synthesis and metabolism in plants and prospects for biofortification","volume":"45","author":"Basset","year":"2005","journal-title":"Crop Sci."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1111\/j.1753-4887.2008.00043.x","article-title":"Can folate intake reduce arsenic toxicity?","volume":"66","author":"Kile","year":"2008","journal-title":"Nutr. Rev."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"380","DOI":"10.1093\/ajcn\/nqy148","article-title":"Folic acid supplementation enhances arsenic methylation: Results from a folic acid and creatine supplementation randomized controlled trial in Bangladesh","volume":"109","author":"Bozack","year":"2019","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"5158","DOI":"10.1073\/pnas.0401342101","article-title":"Enhancement of folates in plants through metabolic engineering","volume":"101","author":"Hossain","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"3069","DOI":"10.1128\/AEM.69.6.3069-3076.2003","article-title":"Increased production of folate by metabolic engineering of Lactococcus lactis","volume":"69","author":"Sybesma","year":"2003","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"612","DOI":"10.1111\/j.1742-4658.2006.05627.x","article-title":"The bile\/arsenite\/riboflavin transporter (BART) superfamily","volume":"274","author":"Mansour","year":"2007","journal-title":"FEBS J."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"19887","DOI":"10.1074\/jbc.M109.011882","article-title":"Properties of arsenite efflux permeases (Acr3) from Alkaliphilus metalliredigens and Corynebacterium glutamicum","volume":"284","author":"Fu","year":"2009","journal-title":"J. Biol. Chem."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"30061","DOI":"10.1074\/jbc.272.48.30061","article-title":"The Saccharomyces cerevisiae ACR3 gene encodes a putative membrane protein involved in arsenite transport","volume":"272","author":"Wysocki","year":"1997","journal-title":"J. Biol. Chem."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"585","DOI":"10.1016\/j.envint.2019.02.058","article-title":"Pathways of arsenic uptake and efflux","volume":"126","author":"Garbinski","year":"2019","journal-title":"Environ. Int."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1007\/s10661-021-09022-0","article-title":"Variability factors of heavy metals in soils and transfer to pasture plants of Mayabeque in Cuba","volume":"193","author":"Lima","year":"2021","journal-title":"Environ. Monit. Assess."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"9825","DOI":"10.1007\/s00253-013-5294-y","article-title":"Design and application of specific 16S rDNA-targeted primers for assessing endophytic diversity in Dendrobium officinale using nested PCR-DGGE","volume":"97","author":"Yu","year":"2013","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1093\/bib\/bbn017","article-title":"MEGA: A biologist-centric software for evolutionary analysis of DNA and protein sequences","volume":"9","author":"Kumar","year":"2008","journal-title":"Brief. Bioinform."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1016\/j.jhazmat.2018.04.007","article-title":"A new model for simulating microbial cyanide production and optimizing the medium parameters for recovering precious metals from waste printed circuit boards","volume":"353","author":"Yuan","year":"2018","journal-title":"J. Hazard. Mater."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"111229","DOI":"10.1016\/j.ecoenv.2020.111229","article-title":"Improved grain yield and lowered arsenic accumulation in rice plants by inoculation with arsenite-oxidizing Achromobacter xylosoxidans GD03","volume":"206","author":"Wang","year":"2020","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_49","first-page":"188","article-title":"R factor transfer in Rhizobium leguminosarum","volume":"84","author":"Beringer","year":"1974","journal-title":"J. Gen. Microbiol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"6614","DOI":"10.1093\/nar\/gkw569","article-title":"NCBI prokaryotic genome annotation pipeline","volume":"44","author":"Tatusova","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"981","DOI":"10.1128\/jb.177.4.981-986.1995","article-title":"The ars operon of Escherichia coli confers arsenical and antimonial resistance","volume":"177","author":"Carlin","year":"1995","journal-title":"J. Bacteriol."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/23\/10\/5619\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T23:13:41Z","timestamp":1760138021000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/23\/10\/5619"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,5,17]]},"references-count":51,"journal-issue":{"issue":"10","published-online":{"date-parts":[[2022,5]]}},"alternative-id":["ijms23105619"],"URL":"https:\/\/doi.org\/10.3390\/ijms23105619","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,5,17]]}}}