{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T16:56:46Z","timestamp":1791824206661,"version":"4.3.4"},"reference-count":112,"publisher":"Public Library of Science (PLoS)","issue":"8","license":[{"start":{"date-parts":[[2021,8,26]],"date-time":"2021-08-26T00:00:00Z","timestamp":1629936000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000060","name":"National Institute of Allergy and Infectious Diseases","doi-asserted-by":"publisher","award":["R01AI130128"],"award-info":[{"award-number":["R01AI130128"]}],"id":[{"id":"10.13039\/100000060","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000060","name":"National Institute of Allergy and Infectious Diseases","doi-asserted-by":"publisher","award":["R01AI146121"],"award-info":[{"award-number":["R01AI146121"]}],"id":[{"id":"10.13039\/100000060","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000897","name":"Cystic Fibrosis Foundation","doi-asserted-by":"publisher","award":["STANTO15RO"],"award-info":[{"award-number":["STANTO15RO"]}],"id":[{"id":"10.13039\/100000897","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000897","name":"Cystic Fibrosis Foundation","doi-asserted-by":"publisher","award":["CRAMERGO19"],"award-info":[{"award-number":["CRAMERGO19"]}],"id":[{"id":"10.13039\/100000897","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000057","name":"National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["P20-GM113132"],"award-info":[{"award-number":["P20-GM113132"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000050","name":"National Heart, Lung, and Blood Institute","doi-asserted-by":"publisher","award":["5T32HL134598"],"award-info":[{"award-number":["5T32HL134598"]}],"id":[{"id":"10.13039\/100000050","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000060","name":"National Institute of Allergy and Infectious Diseases","doi-asserted-by":"publisher","award":["T32AI007519"],"award-info":[{"award-number":["T32AI007519"]}],"id":[{"id":"10.13039\/100000060","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plospathogens.org"],"crossmark-restriction":false},"short-container-title":["PLoS Pathog"],"abstract":"<jats:p>\n                    <jats:italic>Aspergillus fumigatus<\/jats:italic>\n                    is a saprophytic, filamentous fungus found in soils and compost and the causative agent of several pulmonary diseases in humans, birds, and other mammals.\n                    <jats:italic>A<\/jats:italic>\n                    .\n                    <jats:italic>fumigatus<\/jats:italic>\n                    and other filamentous fungi grow as networks of filamentous hyphae that have characteristics of a classic microbial biofilm. These characteristics include production of an extracellular matrix (ECM), surface adhesion, multicellularity, and increased antimicrobial drug resistance.\n                    <jats:italic>A<\/jats:italic>\n                    .\n                    <jats:italic>fumigatus<\/jats:italic>\n                    biofilm growth occurs in vivo at sites of infection, highlighting the importance of defining mechanisms underlying biofilm development and associated emergent properties. We propose that there are 3 distinct phases in the development of\n                    <jats:italic>A<\/jats:italic>\n                    .\n                    <jats:italic>fumigatus<\/jats:italic>\n                    biofilms: biofilm initiation, immature biofilm, and mature biofilm. These stages are defined both temporally and by unique genetic and structural changes over the course of development. Here, we review known mechanisms within each of these stages that contribute to biofilm structure, ECM production, and increased resistance to contemporary antifungal drugs. We highlight gaps in our understanding of biofilm development and function that when addressed are expected to aid in the development of novel antifungal therapies capable of killing filamentous fungal biofilms.\n                  <\/jats:p>","DOI":"10.1371\/journal.ppat.1009794","type":"journal-article","created":{"date-parts":[[2021,8,26]],"date-time":"2021-08-26T13:25:58Z","timestamp":1629984358000},"page":"e1009794","update-policy":"https:\/\/doi.org\/10.1371\/journal.ppat.corrections_policy","source":"Crossref","is-referenced-by-count":95,"title":["Aspergillus fumigatus biofilms: Toward understanding how growth as a multicellular network increases antifungal resistance and disease progression"],"prefix":"10.1371","volume":"17","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9637-030X","authenticated-orcid":true,"given":"Kaesi A.","family":"Morelli","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6869-7404","authenticated-orcid":true,"given":"Joshua D.","family":"Kerkaert","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5503-5006","authenticated-orcid":true,"given":"Robert A.","family":"Cramer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2021,8,26]]},"reference":[{"key":"ppat.1009794.ref001","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1084\/jem.3.1.53","article-title":"Hospi and. A CASE OF PSEUDO-LUPUS VULGARIS CAUSED BY A BLASTOMYCES","volume":"3","author":"BT Caspar Gilchrist","year":"1898","journal-title":"J Exp Med"},{"issue":"9","key":"ppat.1009794.ref002","doi-asserted-by":"crossref","first-page":"1205","DOI":"10.1099\/jmm.0.47247-0","article-title":"Development of a simple model for studying the effects of antifungal agents on multicellular communities of Aspergillus fumigatus","volume":"56","author":"E Mowat","year":"2007","journal-title":"J Med Microbiol"},{"issue":"3","key":"ppat.1009794.ref003","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1111\/j.1472-765X.2006.01960.x","article-title":"Production of cellulase by Aspergillus niger biofilms developed on polyester cloth","volume":"43","author":"GK Villena","year":"2006","journal-title":"Lett Appl Microbiol"},{"issue":"6","key":"ppat.1009794.ref004","doi-asserted-by":"crossref","first-page":"1588","DOI":"10.1111\/j.1462-5822.2007.00895.x","article-title":"An extracellular matrix glues together the aerial-grown hyphae of Aspergillus fumigatus","volume":"9","author":"A Beauvais","year":"2007","journal-title":"Cell Microbiol"},{"issue":"11","key":"ppat.1009794.ref005","doi-asserted-by":"crossref","first-page":"4130","DOI":"10.1128\/AAC.00234-08","article-title":"Aspergillus fumigatus forms biofilms with reduced antifungal drug susceptibility on bronchial epithelial cells","volume":"52","author":"MJ Seidler","year":"2008","journal-title":"Antimicrob Agents Chemother"},{"issue":"11","key":"ppat.1009794.ref006","doi-asserted-by":"crossref","first-page":"475","DOI":"10.1016\/j.tim.2009.08.007","article-title":"Can filamentous fungi form biofilms?","volume":"17","author":"MW Harding","year":"2009","journal-title":"Trends Microbiol"},{"issue":"1","key":"ppat.1009794.ref007","doi-asserted-by":"crossref","first-page":"19","DOI":"10.2217\/17460913.3.1.19","article-title":"Do Aspergillus species produce biofilm?","volume":"3","author":"PH Chandrasekar","year":"2008","journal-title":"Future Microbiol"},{"key":"ppat.1009794.ref008","doi-asserted-by":"crossref","DOI":"10.1080\/13693780802238834","article-title":"The characteristics of Aspergillus fumigatus mycetoma development: Is this a biofilm?","volume":"47","author":"E Mowat","year":"2009","journal-title":"Med Mycol"},{"key":"ppat.1009794.ref009","first-page":"1235","volume-title":"Recent advances on filamentous fungal biofilms for industrial uses. In: Applied Biochemistry and Biotechnology","author":"M Guti\u00e9rrez-Correa","year":"2012"},{"issue":"Suppl. I","key":"ppat.1009794.ref010","doi-asserted-by":"crossref","first-page":"S96","DOI":"10.3109\/13693786.2010.502190","article-title":"Aspergillus fumigatus biofilms in the clinical setting","volume":"49","author":"FMC M\u00fcller","year":"2011","journal-title":"Med Mycol"},{"issue":"4","key":"ppat.1009794.ref011","doi-asserted-by":"crossref","DOI":"10.1128\/microbiolspec.MB-0017-2015","article-title":"Aspergillus Biofilm In Vitro and In Vivo.","volume":"3","author":"A Beauvais","year":"2015","journal-title":"Microbiology Spectrum"},{"issue":"3","key":"ppat.1009794.ref012","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1111\/j.1462-5822.2009.01409.x","article-title":"In vivo biofilm composition of Aspergillus fumigatus","volume":"12","author":"C Loussert","year":"2010","journal-title":"Cell Microbiol"},{"key":"ppat.1009794.ref013","first-page":"1073","article-title":"Antifungal Resistance in Pathogenic Fungi.","volume":"35","author":"S Perea","year":"2002","journal-title":"Antifungal Resistance."},{"issue":"36","key":"ppat.1009794.ref014","doi-asserted-by":"crossref","first-page":"22473","DOI":"10.1073\/pnas.2003700117","article-title":"Fungal biofilm architecture produces hypoxic microenvironments that drive antifungal resistance.","volume":"117","author":"CH Kowalski","year":"2020","journal-title":"PNAS."},{"issue":"6","key":"ppat.1009794.ref015","doi-asserted-by":"crossref","first-page":"1281","DOI":"10.1093\/jac\/dkn402","article-title":"Phase-dependent antifungal activity against Aspergillus fumigatus developing multicellular filamentous biofilms","volume":"62","author":"E Mowat","year":"2008","journal-title":"J Antimicrob Chemother"},{"issue":"2","key":"ppat.1009794.ref016","article-title":"c-di-GMP and its Effects on Biofilm Formation and Dispersion: a Pseudomonas Aeruginosa Review","volume":"3","author":"D-G Ha","year":"2015","journal-title":"Microbiology Spectrum"},{"issue":"1","key":"ppat.1009794.ref017","article-title":"Molecular Mechanisms of Conidial Germination in Aspergillus spp","volume":"84","author":"TJH Baltussen","year":"2019","journal-title":"Microbiol Mol Biol Rev"},{"key":"ppat.1009794.ref018","doi-asserted-by":"crossref","first-page":"3279","DOI":"10.1099\/00221287-146-12-3279","article-title":"Landmarks in the early duplication cycles of Aspergillus fumigatus and Aspergillus nidulans: polarity, germ tube emergence and septation","volume":"146","author":"M Momany","year":"2000","journal-title":"Microbiology"},{"key":"ppat.1009794.ref019","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1139\/m72-045","article-title":"The effects of elevated temperatures on spore swelling and germination in Aspergillus niger","volume":"18","author":"JG Anderson","year":"1972","journal-title":"Can J Microbiol"},{"key":"ppat.1009794.ref020","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1093\/genetics\/155.2.647","article-title":"Conidial Germination in Aspergillus nidulans Requires RAS Signaling and Protein Synthesis","volume":"155","author":"N Osherov","year":"2000","journal-title":"Genetics"},{"key":"ppat.1009794.ref021","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1099\/00222615-41-5-311","article-title":"Oxygen requirements of Aspergillus species","volume":"41","author":"LA Hall","year":"1994","journal-title":"J Med Microbiol"},{"issue":"6","key":"ppat.1009794.ref022","doi-asserted-by":"crossref","first-page":"1814","DOI":"10.1111\/mmi.14395","article-title":"Two hybrid histidine kinases, TcsB and the phytochrome FphA, are involved in temperature sensing in Aspergillus nidulans","volume":"112","author":"Z Yu","year":"2019","journal-title":"Mol Microbiol"},{"issue":"1","key":"ppat.1009794.ref023","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1534\/genetics.105.040584","article-title":"The heterotrimeric G-protein GanB(\u03b1)-SfaD(\u03b2)-GpgA(\u03b3) is a carbon source sensor involved in early cAMP-dependent germination in Aspergillus nidulans.","volume":"171","author":"A Lafon","year":"2005","journal-title":"Genetics"},{"issue":"4","key":"ppat.1009794.ref024","doi-asserted-by":"crossref","first-page":"1001","DOI":"10.1046\/j.1365-2958.2002.02933.x","article-title":"cAMP and ras signalling independently control spore germination in the filamentous fungus Aspergillus nidulans","volume":"44","author":"S Fillinger","year":"2002","journal-title":"Mol Microbiol"},{"issue":"10","key":"ppat.1009794.ref025","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1008419","article-title":"GPCR-mediated glucose sensing system regulates light-dependent fungal development and mycotoxin production","volume":"15","author":"TF dos Reis","year":"2019","journal-title":"PLoS Genet"},{"issue":"9","key":"ppat.1009794.ref026","article-title":"Characterization of gprK encoding a putative hybrid G-protein-coupled receptor in Aspergillus fumigatus","volume":"11","author":"MG Jung","year":"2016","journal-title":"PLoS ONE"},{"issue":"6","key":"ppat.1009794.ref027","doi-asserted-by":"crossref","first-page":"631","DOI":"10.1007\/s00294-009-0276-4","article-title":"G\u03b2\u03b3-mediated growth and developmental control in Aspergillus fumigatus","volume":"55","author":"KS Shin","year":"2009","journal-title":"Curr Genet"},{"issue":"3","key":"ppat.1009794.ref028","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1016\/j.funbio.2020.11.004","article-title":"Minimal nutrient requirements for induction of germination of Aspergillus niger conidia.","volume":"125","author":"M Ijadpanahsaravi","year":"2020","journal-title":"Fungal Biol."},{"issue":"2","key":"ppat.1009794.ref029","doi-asserted-by":"crossref","first-page":"557","DOI":"10.1128\/EC.3.2.557-560.2004","article-title":"A mitogen-activated protein kinase that senses nitrogen regulates conidial germination and growth in Aspergillus fumigatus","volume":"3","author":"T Xue","year":"2004","journal-title":"Eukaryot Cell"},{"issue":"5","key":"ppat.1009794.ref030","doi-asserted-by":"crossref","first-page":"354","DOI":"10.1007\/s00284-006-0413-y","article-title":"Role of respiration in the germination process of the pathogenic mold Aspergillus fumigatus","volume":"54","author":"A Taubitz","year":"2007","journal-title":"Curr Microbiol"},{"issue":"11","key":"ppat.1009794.ref031","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1000200","article-title":"A sterol-regulatory element binding protein is required for cell polarity, hypoxia adaptation, azole drug resistance, and virulence in Aspergillus fumigatus.","volume":"4","author":"SD Willger","year":"2008","journal-title":"PLoS Pathog."},{"issue":"4","key":"ppat.1009794.ref032","doi-asserted-by":"crossref","first-page":"1045","DOI":"10.1111\/j.1365-2958.2010.07509.x","article-title":"Divergent Protein Kinase A isoforms co-ordinately regulate conidial germination, carbohydrate metabolism and virulence in Aspergillus fumigatus","volume":"79","author":"KK Fuller","year":"2011","journal-title":"Mol Microbiol"},{"key":"ppat.1009794.ref033","doi-asserted-by":"crossref","first-page":"796","DOI":"10.1111\/mpp.12342","article-title":"The Magnaporthe grisea species complex and plant pathogenesis","volume":"17","author":"H Zhang","year":"2016","journal-title":"Mol Plant Pathol"},{"key":"ppat.1009794.ref034","first-page":"11","article-title":"Magnaporthe grisea Pth11p Is a Novel Plasma Membrane Protein That Mediates Appressorium Differentiation in Response to Inductive Substrate Cues","author":"TM Dezwaan","year":"2013","journal-title":"Plant Cell"},{"key":"ppat.1009794.ref035","article-title":"Transcriptomic analysis of the exit from dormancy of Aspergillus fumigatus conidia","author":"C Lamarre","year":"2008","journal-title":"BMC Genomics"},{"issue":"3","key":"ppat.1009794.ref036","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1008052","article-title":"The p38\/HOG stress-activated protein kinase network couples growth to division in Candida albicans.","volume":"15","author":"A Sellam","year":"2019","journal-title":"PLoS Genet."},{"key":"ppat.1009794.ref037","doi-asserted-by":"crossref","first-page":"1014","DOI":"10.1016\/j.cub.2004.11.027","article-title":"How cells coordinate growth and division","volume":"14","author":"P Jorgensen","year":"2004","journal-title":"Curr Biol"},{"issue":"1","key":"ppat.1009794.ref038","doi-asserted-by":"crossref","DOI":"10.1186\/s12864-016-2689-z","article-title":"Comparative transcriptome analysis revealing dormant conidia and germination associated genes in Aspergillus species: An essential role for AtfA in conidial dormancy","volume":"17","author":"D Hagiwara","year":"2016","journal-title":"BMC Genomics"},{"issue":"1","key":"ppat.1009794.ref039","doi-asserted-by":"crossref","DOI":"10.1186\/s12866-016-0859-4","article-title":"Analysis and description of the stages of Aspergillus fumigatus biofilm formation using scanning electron microscopy","volume":"16","author":"AI Gonz\u00e1lez-Ram\u00edrez","year":"2016","journal-title":"BMC Microbiol"},{"issue":"8","key":"ppat.1009794.ref040","first-page":"5333","article-title":"Developmental Decisions in Aspergillus nidulans Are Modulated by Ras Activity","volume":"14","author":"T Som","year":"1994","journal-title":"Mol Cell Biol"},{"issue":"7","key":"ppat.1009794.ref041","doi-asserted-by":"crossref","first-page":"1085","DOI":"10.1128\/EC.00086-08","article-title":"Calcineurin target CrzA regulates conidial germination, hyphal growth, and pathogenesis of Aspergillus fumigatus","volume":"7","author":"RA Cramer","year":"2008","journal-title":"Eukaryot Cell"},{"key":"ppat.1009794.ref042","doi-asserted-by":"crossref","first-page":"1127","DOI":"10.1126\/science.8493556","article-title":"Expression of Pseudomonas aeruginosa Virulence Genes Requires Cell-to-Cell Communication","volume":"260","author":"L Passador","year":"1993","journal-title":"Science"},{"issue":"11","key":"ppat.1009794.ref043","doi-asserted-by":"crossref","first-page":"3620","DOI":"10.1128\/JB.187.11.3620-3629.2005","article-title":"Vibrio fischeri uses two quorum-sensing systems for the regulation of early and late colonization factors","volume":"187","author":"C Lupp","year":"2005","journal-title":"J Bacteriol"},{"key":"ppat.1009794.ref044","doi-asserted-by":"crossref","first-page":"1688","DOI":"10.1086\/317606","article-title":"Impact of the agr Quorum-Sensing System on Adherence to Polystyrene in Staphylococcus aureus","volume":"182","author":"C Vuong","year":"2000","journal-title":"J Infect Dis"},{"issue":"3","key":"ppat.1009794.ref045","doi-asserted-by":"crossref","first-page":"1356","DOI":"10.1128\/AEM.70.3.1356-1359.2004","article-title":"Inoculum Size Effect in Dimorphic Fungi: Extracellular Control of Yeast-Mycelium Dimorphism in Ceratocystis ulmi","volume":"70","author":"JM Hornby","year":"2004","journal-title":"Appl Environ Microbiol"},{"key":"ppat.1009794.ref046","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1016\/S0924-8579(02)00189-9","article-title":"Effect of increasing inoculum sizes of Aspergillus hyphae on MICs and MFCs of antifungal agents by broth microdilution method.","volume":"21","author":"C Lass-Fl\u00f6 Rl","year":"2003","journal-title":"Int J Antimicrob Agents"},{"issue":"7","key":"ppat.1009794.ref047","doi-asserted-by":"crossref","first-page":"2982","DOI":"10.1128\/AEM.67.7.2982-2992.2001","article-title":"Quorum Sensing in the Dimorphic Fungus Candida albicans Is Mediated by Farnesol","volume":"67","author":"JM Hornby","year":"2001","journal-title":"Appl Environ Microbiol"},{"issue":"3","key":"ppat.1009794.ref048","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1111\/j.1365-2958.2007.05666.x","article-title":"TUP1 disruption in Cryptococcus neoformans uncovers a peptide-mediated density-dependent growth phenomenon that mimics quorum sensing","volume":"64","author":"H Lee","year":"2007","journal-title":"Mol Microbiol"},{"key":"ppat.1009794.ref049","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1038\/s41579-018-0121-1","article-title":"Fungal secondary metabolism: regulation, function and drug discovery","volume":"17","author":"NP Keller","year":"2019","journal-title":"Nat Rev Microbiol"},{"issue":"5","key":"ppat.1009794.ref050","doi-asserted-by":"crossref","first-page":"1908","DOI":"10.1128\/AAC.48.5.1908-1911.2004","article-title":"Novel Inhalational Murine Model of Invasive Pulmonary Aspergillosis","volume":"48","author":"DC Sheppard","year":"2004","journal-title":"Antimicrob Agents Chemother"},{"issue":"4","key":"ppat.1009794.ref051","doi-asserted-by":"crossref","first-page":"219","DOI":"10.2500\/ajra.2011.25.3622","article-title":"Aspergillus fumigatus biofilm on primary human sinonasal epithelial culture","volume":"25","author":"D Singhal","year":"2011","journal-title":"Am J Rhinol Allergy"},{"issue":"3","key":"ppat.1009794.ref052","first-page":"0232","article-title":"Aspergillus fumigatus triggers inflammatory responses by stage-specific \u03b2-glucan display.","volume":"1","author":"TM Hohl","year":"2005","journal-title":"PLoS Pathog"},{"issue":"3","key":"ppat.1009794.ref053","doi-asserted-by":"crossref","DOI":"10.1111\/cmi.12802","article-title":"Aspergillus fumigatus adhesion factors in dormant conidia revealed through comparative phenotypic and transcriptomic analyses","volume":"20","author":"A Takahashi-Nakaguchi","year":"2018","journal-title":"Cell Microbiol"},{"issue":"4","key":"ppat.1009794.ref054","doi-asserted-by":"crossref","first-page":"1146","DOI":"10.1128\/iai.64.4.1146-1153.1996","article-title":"Binding of human fibronectin to Aspergillus fumigatus conidia","volume":"64","author":"MC Penalver","year":"1996","journal-title":"Infect Immun"},{"issue":"6","key":"ppat.1009794.ref055","doi-asserted-by":"crossref","first-page":"714","DOI":"10.1099\/jmm.0.005991-0","article-title":"Identification and characterization of a laminin-binding protein of Aspergillus fumigatus: Extracellular thaumatin domain protein (AfCalAp).","volume":"58","author":"SK Upadhyay","year":"2009","journal-title":"J Med Microbiol"},{"key":"ppat.1009794.ref056","first-page":"2","article-title":"Aspergillus fumigatus CalA binds to integrin \u03b15\u03b21 and mediates host cell invasion","volume":"14","author":"H Liu","year":"2016","journal-title":"Nat Microbiol"},{"issue":"10","key":"ppat.1009794.ref057","doi-asserted-by":"crossref","first-page":"4380","DOI":"10.1128\/iai.62.10.4380-4388.1994","article-title":"Rodletless Mutants of Aspergillus fumigatus","volume":"62","author":"N Thau","year":"1994","journal-title":"Infect Immun"},{"issue":"6","key":"ppat.1009794.ref058","doi-asserted-by":"crossref","first-page":"1021","DOI":"10.1007\/s10156-013-0614-9","article-title":"Molecular characterization of AfuFleA, an l-fucose-specific lectin from Aspergillus fumigatus.","volume":"19","author":"S Kuboi","year":"2013","journal-title":"J Infect Chemother"},{"issue":"12","key":"ppat.1009794.ref059","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0083077","article-title":"A soluble fucose-specific lectin from Aspergillus fumigatus conidia\u2014Structure, specificity and possible role in fungal pathogenicity.","volume":"8","author":"J Houser","year":"2013","journal-title":"PLoS ONE."},{"issue":"12","key":"ppat.1009794.ref060","doi-asserted-by":"crossref","first-page":"6891","DOI":"10.1128\/IAI.70.12.6891-6895.2002","article-title":"Purification and partial characterization of a 32-kilodalton sialic acid-specific lectin from Aspergillus fumigatus","volume":"70","author":"G Tronchin","year":"2002","journal-title":"Infect Immun"},{"issue":"1","key":"ppat.1009794.ref061","doi-asserted-by":"crossref","first-page":"26","DOI":"10.1128\/EC.05270-11","article-title":"\u03b11,3 glucans are dispensable in Aspergillus fumigatus","volume":"11","author":"C Henry","year":"2012","journal-title":"Eukaryot Cell"},{"issue":"8","key":"ppat.1009794.ref062","doi-asserted-by":"crossref","first-page":"707","DOI":"10.1016\/j.fgb.2010.04.006","article-title":"Cell wall \u03b11-3glucans induce the aggregation of germinating conidia of Aspergillus fumigatus","volume":"47","author":"T Fontaine","year":"2010","journal-title":"Fungal Genet Biol"},{"key":"ppat.1009794.ref063","doi-asserted-by":"crossref","first-page":"S239","DOI":"10.1093\/mmy\/myy140","article-title":"Integration of electron microscopy and solid-state NMR analysis for new views and compositional parameters of Aspergillus fumigatus biofilms","volume":"57","author":"C Reichhardt","year":"2019","journal-title":"Med Mycol"},{"issue":"10","key":"ppat.1009794.ref064","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pbio.2006872","article-title":"Candida albicans biofilm\u2013induced vesicles confer drug resistance through matrix biogenesis","volume":"16","author":"R Zarnowski","year":"2018","journal-title":"PLoS Biol"},{"key":"ppat.1009794.ref065","article-title":"Characterization of Aspergillus fumigatus Extracellular Vesicles and Their Effects on Macrophages and Neutrophils Functions.","volume":"10","author":"JAM Souza","year":"2019","journal-title":"Front Microbiol."},{"issue":"8","key":"ppat.1009794.ref066","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1003575","article-title":"Aspergillus Galactosaminogalactan Mediates Adherence to Host Constituents and Conceals Hyphal \u03b2-Glucan from the Immune System.","volume":"9","author":"FN Gravelat","year":"2013","journal-title":"PLoS Pathog."},{"issue":"4","key":"ppat.1009794.ref067","doi-asserted-by":"crossref","first-page":"473","DOI":"10.1111\/j.1462-5822.2009.01408.x","article-title":"Aspergillus fumigatus MedA governs adherence, host cell interactions and virulence","volume":"12","author":"FN Gravelat","year":"2010","journal-title":"Cell Microbiol"},{"issue":"6","key":"ppat.1009794.ref068","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.02329-20","article-title":"The Transcription Factor SomA Synchronously Regulates Biofilm Formation and Cell Wall Homeostasis in Aspergillus fumigatus.","volume":"11","author":"Y Chen","year":"2020","journal-title":"MBio"},{"issue":"11","key":"ppat.1009794.ref069","article-title":"Transcription Factor SomA Is Required for Adhesion, Development and Virulence of the Human Pathogen Aspergillus fumigatus","volume":"11","author":"CJ Lin","year":"2015","journal-title":"PLoS Pathogens"},{"issue":"1","key":"ppat.1009794.ref070","doi-asserted-by":"crossref","DOI":"10.1038\/s41467-019-14191-1","article-title":"The negative cofactor 2 complex is a key regulator of drug resistance in Aspergillus fumigatus.","volume":"11","author":"T Furukawa","year":"2020","journal-title":"Nat Commun"},{"key":"ppat.1009794.ref071","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1016\/j.tcsw.2018.03.002","article-title":"Mitogen activated protein kinases (MAPK) and protein phosphatases are involved in Aspergillus fumigatus adhesion and biofilm formation.","volume":"1","author":"AO Manfiolli","year":"2018","journal-title":"Cell Surface"},{"issue":"8","key":"ppat.1009794.ref072","doi-asserted-by":"crossref","first-page":"728","DOI":"10.1128\/EC.00008-15","article-title":"The Aspergillus fumigatus sitA phosphatase homologue is important for adhesion, cell wall integrity, biofilm formation, and virulence","volume":"14","author":"VLP Bom","year":"2015","journal-title":"Eukaryot Cell"},{"key":"ppat.1009794.ref073","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1016\/j.fgb.2014.10.011","article-title":"The role of AtfA and HOG MAPK pathway in stress tolerance in conidia of Aspergillus fumigatus","volume":"73","author":"D Hagiwara","year":"2014","journal-title":"Fungal Genet Biol"},{"issue":"15","key":"ppat.1009794.ref074","doi-asserted-by":"crossref","first-page":"5576","DOI":"10.1073\/pnas.0407097102","article-title":"A contact-activated kinase signals Candida albicans invasive growth and biofilm development","volume":"102","author":"CA Kumamoto","year":"2005","journal-title":"PNAS"},{"issue":"5","key":"ppat.1009794.ref075","doi-asserted-by":"crossref","first-page":"2092","DOI":"10.1128\/AAC.01189-10","article-title":"Azole resistance of Aspergillus fumigatus biofilms is partly associated with efflux pump activity","volume":"55","author":"R Rajendran","year":"2011","journal-title":"Antimicrob Agents Chemother"},{"issue":"47","key":"ppat.1009794.ref076","doi-asserted-by":"crossref","first-page":"45688","DOI":"10.1074\/jbc.M206479200","article-title":"A novel electron paramagnetic resonance approach to determine the mechanism of drug transport by P-glycoprotein","volume":"277","author":"H Omote","year":"2002","journal-title":"J Biol Chem"},{"issue":"SUPPL.1","key":"ppat.1009794.ref077","article-title":"The ergosterol biosynthesis pathway, transporter genes, and azole resistance in Aspergillus fumigatus","volume":"43","author":"ME da Silva Ferreira","year":"2005","journal-title":"Med Mycol"},{"key":"ppat.1009794.ref078","article-title":"Fungal biofilm resistance","author":"G Ramage","year":"2012","journal-title":"International Journal of Microbiology"},{"issue":"1","key":"ppat.1009794.ref079","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1128\/EC.05102-11","article-title":"Global transcriptome changes underlying colony growth in the opportunistic human pathogen Aspergillus fumigatus","volume":"11","author":"JG Gibbons","year":"2012","journal-title":"Eukaryot Cell"},{"issue":"5","key":"ppat.1009794.ref080","doi-asserted-by":"crossref","first-page":"1719","DOI":"10.1128\/AAC.47.5.1719-1726.2003","article-title":"Multiple resistance mechanisms among Aspergillus fumigatus mutants with high-level resistance to itraconazole","volume":"47","author":"AM Nascimento","year":"2003","journal-title":"Antimicrob Agents Chemother"},{"issue":"1","key":"ppat.1009794.ref081","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1006096","article-title":"A Novel Zn2-Cys6Transcription Factor AtrR Plays a Key Role in an Azole Resistance Mechanism of Aspergillus fumigatus by Co-regulating cyp51A and cdr1B Expressions.","volume":"13","author":"D Hagiwara","year":"2017","journal-title":"PLoS Pathog"},{"issue":"8","key":"ppat.1009794.ref082","doi-asserted-by":"crossref","first-page":"3505","DOI":"10.1128\/AAC.00227-10","article-title":"Role of Fks1p and matrix glucan in Candida albicans biofilm resistance to an echinocandin, pyrimidine, and polyene","volume":"54","author":"JE Nett","year":"2010","journal-title":"Antimicrob Agents Chemother"},{"issue":"12","key":"ppat.1009794.ref083","doi-asserted-by":"crossref","first-page":"2430","DOI":"10.1038\/s41564-019-0558-7","article-title":"Fungal biofilm morphology impacts hypoxia fitness and disease progression.","volume":"4","author":"CH Kowalski","year":"2019","journal-title":"Nat Microbiol"},{"issue":"27","key":"ppat.1009794.ref084","doi-asserted-by":"crossref","first-page":"7124","DOI":"10.1073\/pnas.1702798114","article-title":"Microbial glycoside hydrolases as antibiofilm agents with cross-kingdom activity.","volume":"114","author":"BD Snarr","year":"2017","journal-title":"PNAS"},{"issue":"6","key":"ppat.1009794.ref085","doi-asserted-by":"crossref","DOI":"10.1128\/AAC.00241-18","article-title":"Insight into the significance of Aspergillus fumigatus cyp51A polymorphisms","volume":"62","author":"R Garcia-Rubio","year":"2018","journal-title":"Antimicrob Agents Chemother"},{"issue":"12","key":"ppat.1009794.ref086","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1002374","article-title":"SREBP coordinates iron and ergosterol homeostasis to mediate triazole drug and hypoxia responses in the human fungal pathogen Aspergillus fumigatus.","volume":"7","author":"M Blatzer","year":"2011","journal-title":"PLoS Genet."},{"key":"ppat.1009794.ref087","doi-asserted-by":"crossref","DOI":"10.3389\/fmicb.2016.00682","article-title":"Dynamics of actin cables in polarized growth of the filamentous fungus Aspergillus nidulans.","volume":"7","author":"A Bergs","year":"2016","journal-title":"Front Microbiol."},{"issue":"5","key":"ppat.1009794.ref088","doi-asserted-by":"crossref","first-page":"634","DOI":"10.1091\/mbc.e16-10-0750","article-title":"Microtubules are reversibly depolymerized in response to changing gaseous microenvironments within Aspergillus nidulans biofilms","volume":"28","author":"N Shukla","year":"2017","journal-title":"Mol Biol Cell"},{"issue":"3","key":"ppat.1009794.ref089","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1002\/jobm.201900550","article-title":"Developmental stages of biofilm and characterization of extracellular matrix of manglicolous fungus Aspergillus niger BSC-1","volume":"60","author":"S Chatterjee","year":"2020","journal-title":"J Basic Microbiol"},{"issue":"9","key":"ppat.1009794.ref090","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1005884","article-title":"The Extracellular Matrix of Candida albicans Biofilms Impairs Formation of Neutrophil Extracellular Traps.","volume":"12","author":"CJ Johnson","year":"2016","journal-title":"PLoS Pathog"},{"issue":"10","key":"ppat.1009794.ref091","article-title":"The Fungal Exopolysaccharide Galactosaminogalactan Mediates Virulence by Enhancing Resistance to Neutrophil Extracellular Traps.","volume":"11","author":"MJ Lee","year":"2015","journal-title":"PLoS Pathog."},{"issue":"3","key":"ppat.1009794.ref092","doi-asserted-by":"crossref","first-page":"420","DOI":"10.1128\/EC.00287-12","article-title":"Extracellular DNA release acts as an antifungal resistance mechanism in mature Aspergillus fumigatus biofilms","volume":"12","author":"R Rajendran","year":"2013","journal-title":"Eukaryot Cell"},{"issue":"7","key":"ppat.1009794.ref093","doi-asserted-by":"crossref","first-page":"2083","DOI":"10.1099\/mic.0.2007\/006031-0","article-title":"Role of autolysin-mediated DNA release in biofilm formation of Staphylococcus epidermidis","volume":"153","author":"Z Qin","year":"2007","journal-title":"Microbiology"},{"issue":"2\u20133","key":"ppat.1009794.ref094","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1007\/s12010-008-8174-7","article-title":"Regulation of autolysis in Aspergillus nidulans","volume":"151","author":"T Emri","year":"2008","journal-title":"Appl Biochem Biotechnol"},{"issue":"1","key":"ppat.1009794.ref095","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.03579-20","article-title":"A Heterogeneously Expressed Gene Family Modulates the Biofilm Architecture and Hypoxic Growth of Aspergillus fumigatus.","volume":"12","author":"CH Kowalski","year":"2021","journal-title":"MBio"},{"issue":"9","key":"ppat.1009794.ref096","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1000154","article-title":"Sub-telomere directed gene expression during initiation of invasive aspergillosis","volume":"4","author":"A McDonagh","year":"2008","journal-title":"PLoS Pathog"},{"issue":"5","key":"ppat.1009794.ref097","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1007787","article-title":"Circuit diversification in a biofilm regulatory network.","volume":"15","author":"MY Huang","year":"2019","journal-title":"PLoS Pathog"},{"issue":"5","key":"ppat.1009794.ref098","doi-asserted-by":"crossref","first-page":"399","DOI":"10.2144\/000113399","article-title":"Laser nephelometry applied in an automated microplate system to study filamentous fungus growth","volume":"48","author":"A Joubert","year":"2010","journal-title":"Biotechniques"},{"key":"ppat.1009794.ref099","doi-asserted-by":"crossref","first-page":"676","DOI":"10.1038\/nmeth.2019","article-title":"Fiji: An open-source platform for biological-image analysis.","volume":"9","author":"J Schindelin","journal-title":"Nat Methods"},{"issue":"2","key":"ppat.1009794.ref100","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1038\/s41564-020-00817-4","article-title":"Quantitative image analysis of microbial communities with BiofilmQ.","volume":"6","author":"R Hartmann","year":"2021","journal-title":"Nat Microbiol"},{"issue":"1","key":"ppat.1009794.ref101","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.02752-19","article-title":"Three-dimensional light sheet fluorescence microscopy of lungs to dissect local host immune-aspergillus fumigatus interactions.","volume":"11","author":"J Amich","year":"2020","journal-title":"MBio."},{"issue":"1","key":"ppat.1009794.ref102","doi-asserted-by":"crossref","DOI":"10.1038\/s41467-021-21965-z","article-title":"Antibody-guided in vivo imaging of Aspergillus fumigatus lung infections during antifungal azole treatment","volume":"12","author":"S Henneberg","year":"2021","journal-title":"Nat Commun"},{"key":"ppat.1009794.ref103","doi-asserted-by":"crossref","first-page":"S50","DOI":"10.1016\/j.jinf.2016.04.022","article-title":"Aspergillus infections in cystic fibrosis.","volume":"72","author":"J King","year":"2016","journal-title":"J Infect."},{"issue":"3","key":"ppat.1009794.ref104","doi-asserted-by":"crossref","DOI":"10.1128\/CMR.00138-18","article-title":"Cystic Fibrosis and Pseudomonas aeruginosa: the Host-Microbe Interface.","volume":"32","author":"S Malhotra","year":"2019","journal-title":"Clin Microbiol Rev"},{"issue":"O1","key":"ppat.1009794.ref105","doi-asserted-by":"crossref","DOI":"10.3109\/13693786.2010.503665","article-title":"Characteristics and consequences of airway colonization by filamentous fungi in 201 adult patients with cystic fibrosis in France","volume":"48","author":"A Paugam","year":"2010","journal-title":"Med Mycol"},{"issue":"1","key":"ppat.1009794.ref106","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1007\/s11046-017-0190-0","article-title":"Aspergillus Bronchitis in Patients with Cystic Fibrosis","volume":"183","author":"C Brandt","year":"2018","journal-title":"Mycopathologia"},{"key":"ppat.1009794.ref107","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1111\/j.1574-6968.2010.02130.x","article-title":"Pseudomonas aeruginosa and their small diffusible extracellular molecules inhibit Aspergillus fumigatus biofilm formation","volume":"313","author":"E Mowat","year":"2010","journal-title":"FEMS Microbiol Lett"},{"issue":"11","key":"ppat.1009794.ref108","doi-asserted-by":"crossref","first-page":"1211","DOI":"10.1164\/rccm.200603-423OC","article-title":"Effect of allergic bronchopulmonary aspergillosis on lung function in children with cystic fibrosis","volume":"174","author":"R Kraemer","year":"2006","journal-title":"Am J Respir Crit Care Med"},{"key":"ppat.1009794.ref109","doi-asserted-by":"crossref","first-page":"449","DOI":"10.1007\/s12275-018-8114-3","article-title":"Interdependence between iron acquisition and biofilm formation in Pseudomonas aeruginosa","volume":"56","author":"D Kang","year":"2018","journal-title":"J Microbiol"},{"issue":"14","key":"ppat.1009794.ref110","doi-asserted-by":"crossref","first-page":"4959","DOI":"10.1128\/AEM.00182-11","article-title":"SidL, an Aspergillus fumigatus transacetylase involved in biosynthesis of the siderophores ferricrocin and hydroxyferricrocin","volume":"77","author":"M Blatzer","year":"2011","journal-title":"Appl Environ Microbiol"},{"issue":"5","key":"ppat.1009794.ref111","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0216085","article-title":"Intermicrobial interaction: Aspergillus fumigatus siderophores protect against competition by Pseudomonas aeruginosa.","volume":"14","author":"G Sass","year":"2019","journal-title":"PLoS ONE"},{"issue":"23","key":"ppat.1009794.ref112","doi-asserted-by":"crossref","DOI":"10.21769\/BioProtoc.1667","article-title":"Establishing a Biofilm Co-culture of Pseudomonas and Aspergillus for Metabolite Extraction.","volume":"5","author":"H Zheng","year":"2015","journal-title":"Bio-Protocol."}],"container-title":["PLOS Pathogens"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.ppat.1009794","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,8,26]],"date-time":"2021-08-26T13:27:25Z","timestamp":1629984445000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.ppat.1009794"}},"subtitle":[],"editor":[{"given":"Chaoyang","family":"Xue","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2021,8,26]]},"references-count":112,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2021,8,26]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.ppat.1009794","relation":{},"ISSN":["1553-7374"],"issn-type":[{"value":"1553-7374","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,8,26]]}}}