{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T02:48:32Z","timestamp":1790218112413,"version":"4.1.0"},"update-to":[{"DOI":"10.1371\/journal.ppat.1007144","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2018,6,27]],"date-time":"2018-06-27T00:00:00Z","timestamp":1530057600000}}],"reference-count":102,"publisher":"Public Library of Science (PLoS)","issue":"6","license":[{"start":{"date-parts":[[2018,6,15]],"date-time":"2018-06-15T00:00:00Z","timestamp":1529020800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["www.plospathogens.org"],"crossmark-restriction":false},"short-container-title":["PLoS Pathog"],"DOI":"10.1371\/journal.ppat.1007144","type":"journal-article","created":{"date-parts":[[2018,6,15]],"date-time":"2018-06-15T17:49:57Z","timestamp":1529084997000},"page":"e1007144","update-policy":"https:\/\/doi.org\/10.1371\/journal.ppat.corrections_policy","source":"Crossref","is-referenced-by-count":140,"title":["Cryptococcus neoformans urease affects the outcome of intracellular pathogenesis by modulating phagolysosomal pH"],"prefix":"10.1371","volume":"14","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5376-7243","authenticated-orcid":true,"given":"Man Shun","family":"Fu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carolina","family":"Coelho","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5669-3035","authenticated-orcid":true,"given":"Carlos M.","family":"De Leon-Rodriguez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Diego C. P.","family":"Rossi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8591-8176","authenticated-orcid":true,"given":"Emma","family":"Camacho","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2193-4411","authenticated-orcid":true,"given":"Eric H.","family":"Jung","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6889-5985","authenticated-orcid":true,"given":"Madhura","family":"Kulkarni","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Arturo","family":"Casadevall","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2018,6,15]]},"reference":[{"key":"ref1","article-title":"Global burden of disease of HIV-associated cryptococcal meningitis: an updated analysis","volume":"0","author":"R Rajasingham","year":"2017","journal-title":"Lancet Infect Dis"},{"key":"ref2","doi-asserted-by":"crossref","first-page":"685","DOI":"10.1128\/JB.62.6.685-690.1951","article-title":"ISOLATION OF CRYPTOCOCCUS NEOFORMANS FROM SOIL","volume":"62","author":"CW Emmons","year":"1951","journal-title":"J Bacteriol"},{"key":"ref3","first-page":"227","article-title":"Saprophytic sources of Cryptococcus neoformans associated with the pigeon (Columba livia)","volume":"62","author":"CW Emmons","year":"1955","journal-title":"Am J Hyg"},{"key":"ref4","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1080\/mmy.38.5.379.383","article-title":"Possible primary ecological niche of Cryptococcus neoformans","volume":"38","author":"MS Lazera","year":"2000","journal-title":"Med Mycol"},{"key":"ref5","first-page":"97","article-title":"First isolation of Cryptococcus neoformans genotype VNI MAT-alpha from wood inside hollow trunks of Hymenaea courbaril","volume":"54","author":"DM Castro e Silva","year":"2016","journal-title":"Med Mycol"},{"key":"ref6","doi-asserted-by":"crossref","first-page":"992","DOI":"10.2460\/javma.1979.175.09.992","article-title":"Cryptococcosis in a male Beccari\u2019s crowned pigeon","volume":"175","author":"PK Ensley","year":"1979","journal-title":"J Am Vet Med Assoc"},{"key":"ref7","first-page":"207","article-title":"Presence of pathogenic yeasts in the feces of the semi-domesticated pigeon (Columba livia, Gmelin 1789, urban type) from the city of Turin","volume":"31","author":"MG Gallo","year":"1989","journal-title":"Parassitologia"},{"key":"ref8","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1099\/00222615-11-2-215","article-title":"Carriage of Cryptococcus neoformans in the crops of pigeons","volume":"11","author":"ZU Khan","year":"1978","journal-title":"J Med Microbiol"},{"key":"ref9","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1023\/A:1006863705759","article-title":"Isolation of Cryptococcus neoformans from pigeon (Columba livia) droppings in northern Iran","volume":"139","author":"AR Khosravi","year":"1997","journal-title":"Mycopathologia"},{"key":"ref10","doi-asserted-by":"crossref","first-page":"642","DOI":"10.11150\/kansenshogakuzasshi1970.69.642","article-title":"Isolation of Cryptococcus neoformans from environments (pigeon excreta) in Nagasaki","volume":"69","author":"Y Yamamoto","year":"1995","journal-title":"Kansenshogaku Zasshi"},{"key":"ref11","first-page":"56","article-title":"Isolation of Cryptococcus neoformans and other opportunistic fungi from pigeon droppings","volume":"18","author":"M Soltani","year":"2013","journal-title":"J Res Med Sci Off J Isfahan Univ Med Sci"},{"key":"ref12","doi-asserted-by":"crossref","first-page":"932","DOI":"10.2353\/ajpath.2009.080673","article-title":"Cryptococcal Urease Promotes the Accumulation of Immature Dendritic Cells and a Non-Protective T2 Immune Response within the Lung","volume":"174","author":"JJ Osterholzer","year":"2009","journal-title":"Am J Pathol"},{"key":"ref13","doi-asserted-by":"crossref","first-page":"3244","DOI":"10.4049\/jimmunol.175.5.3244","article-title":"An Innate Immune System Cell Is a Major Determinant of Species-Related Susceptibility Differences to Fungal Pneumonia","volume":"175","author":"X Shao","year":"2005","journal-title":"J Immunol"},{"key":"ref14","doi-asserted-by":"crossref","first-page":"885","DOI":"10.1128\/IAI.67.2.885-890.1999","article-title":"Cryptococcus neoformans Resides in an Acidic Phagolysosome of Human Macrophages","volume":"67","author":"SM Levitz","year":"1999","journal-title":"Infect Immun"},{"key":"ref15","doi-asserted-by":"crossref","first-page":"4225","DOI":"10.1128\/IAI.68.7.4225-4237.2000","article-title":"Cryptococcus neoformans Is a Facultative Intracellular Pathogen in Murine Pulmonary Infection","volume":"68","author":"M Feldmesser","year":"2000","journal-title":"Infect Immun"},{"key":"ref16","doi-asserted-by":"crossref","first-page":"15945","DOI":"10.1073\/pnas.0607528103","article-title":"Recruitment of CD63 to Cryptococcus neoformans phagosomes requires acidification","volume":"103","author":"K Artavanis-Tsakonas","year":"2006","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref17","doi-asserted-by":"crossref","first-page":"2161","DOI":"10.1016\/j.cub.2006.09.061","article-title":"Phagosome Extrusion and Host-Cell Survival after Cryptococcus neoformans Phagocytosis by Macrophages","volume":"16","author":"M Alvarez","year":"2006","journal-title":"Curr Biol"},{"key":"ref18","doi-asserted-by":"crossref","first-page":"832","DOI":"10.1128\/IAI.68.2.832-838.2000","article-title":"Persistent Cryptococcus neoformans Pulmonary Infection in the Rat Is Associated with Intracellular Parasitism, Decreased Inducible Nitric Oxide Synthase Expression, and Altered Antibody Responsiveness to Cryptococcal Polysaccharide","volume":"68","author":"DL Goldman","year":"2000","journal-title":"Infect Immun"},{"key":"ref19","doi-asserted-by":"crossref","first-page":"522","DOI":"10.1086\/341564","article-title":"Pathogenesis of Cerebral Cryptococcus neoformans Infection after Fungemia","volume":"186","author":"F Chr\u00e9tien","year":"2002","journal-title":"J Infect Dis"},{"key":"ref20","doi-asserted-by":"crossref","first-page":"1550","DOI":"10.1128\/CVI.00242-07","article-title":"Serologic Evidence for Reactivation of Cryptococcosis in Solid-Organ Transplant Recipients","volume":"14","author":"DC Saha","year":"2007","journal-title":"Clin Vaccine Immunol CVI"},{"key":"ref21","first-page":"78","article-title":"Histopathology of cryptococcosis and other fungal infections in patients with acquired immunodeficiency syndrome","volume":"5","author":"K Shibuya","year":"2001","journal-title":"Int J Infect Dis IJID Off Publ Int Soc Infect Dis"},{"key":"ref22","doi-asserted-by":"crossref","first-page":"3204","DOI":"10.1128\/JCM.37.10.3204-3209.1999","article-title":"Epidemiological Evidence for Dormant Cryptococcus neoformans Infection","volume":"37","author":"D Garcia-Hermoso","year":"1999","journal-title":"J Clin Microbiol"},{"key":"ref23","doi-asserted-by":"crossref","first-page":"4792","DOI":"10.1128\/IAI.00587-07","article-title":"Depletion of Alveolar Macrophages Decreases the Dissemination of a Glucosylceramide-Deficient Mutant of Cryptococcus neoformans in Immunodeficient Mice","volume":"75","author":"TB Kechichian","year":"2007","journal-title":"Infect Immun"},{"key":"ref24","doi-asserted-by":"crossref","first-page":"2156","DOI":"10.1016\/j.cub.2006.09.032","article-title":"Expulsion of Live Pathogenic Yeast by Macrophages","volume":"16","author":"H Ma","year":"2006","journal-title":"Curr Biol"},{"key":"ref25","doi-asserted-by":"crossref","first-page":"1067","DOI":"10.1128\/EC.3.5.1067-1075.2004","article-title":"Role of Phagocytosis in the Virulence of Cryptococcus neoformans","volume":"3","author":"M Del Poeta","year":"2004","journal-title":"Eukaryot Cell"},{"key":"ref26","doi-asserted-by":"crossref","first-page":"120","DOI":"10.1128\/IAI.01065-08","article-title":"Evidence of a Role for Monocytes in Dissemination and Brain Invasion by Cryptococcus neoformans","volume":"77","author":"C Charlier","year":"2009","journal-title":"Infect Immun"},{"key":"ref27","doi-asserted-by":"crossref","first-page":"e1700898","DOI":"10.1126\/sciadv.1700898","article-title":"Vomocytosis of live pathogens from macrophages is regulated by the atypical MAP kinase ERK5","volume":"3","author":"AS Gilbert","year":"2017","journal-title":"Sci Adv"},{"key":"ref28","doi-asserted-by":"crossref","first-page":"15245","DOI":"10.1073\/pnas.261418798","article-title":"Cryptococcus neoformans interactions with amoebae suggest an explanation for its virulence and intracellular pathogenic strategy in macrophages","volume":"98","author":"JN Steenbergen","year":"2001","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref29","doi-asserted-by":"crossref","first-page":"667","DOI":"10.1016\/S1286-4579(03)00092-3","article-title":"The origin and maintenance of virulence for the human pathogenic fungus Cryptococcus neoformans","volume":"5","author":"JN Steenbergen","year":"2003","journal-title":"Microbes Infect"},{"key":"ref30","doi-asserted-by":"crossref","DOI":"10.3389\/fcimb.2018.00108","article-title":"Cryptococcus neoformans Escape From Dictyostelium Amoeba by Both WASH-Mediated Constitutive Exocytosis and Vomocytosis","volume":"8","author":"RA Watkins","year":"2018","journal-title":"Front Cell Infect Microbiol"},{"key":"ref31","doi-asserted-by":"crossref","first-page":"2604","DOI":"10.1128\/IAI.63.7.2604-2611.1995","article-title":"Ingestion of acapsular Cryptococcus neoformans occurs via mannose and beta-glucan receptors, resulting in cytokine production and increased phagocytosis of the encapsulated form","volume":"63","author":"CE Cross","year":"1995","journal-title":"Infect Immun"},{"key":"ref32","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1111\/cmi.12067","article-title":"Cryptococcus interactions with macrophages: evasion and manipulation of the phagosome by a fungal pathogen","volume":"15","author":"SA Johnston","year":"2013","journal-title":"Cell Microbiol"},{"key":"ref33","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1128\/IAI.16.1.99-106.1977","article-title":"Non-encapsulated variant of Cryptococcus neoformans. II. Surface receptors for cryptococcal polysaccharide and their role in inhibition of phagocytosis by polysaccharide","volume":"16","author":"TR Kozel","year":"1977","journal-title":"Infect Immun"},{"key":"ref34","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1080\/mmy.38.6.407.417","article-title":"Immunoregulation by capsular components of Cryptococcus neoformans","volume":"38","author":"A Vecchiarelli","year":"2000","journal-title":"Med Mycol"},{"key":"ref35","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1080\/j.1365-280X.1999.00218.x","article-title":"Melanin as a potential cryptococcal defence against microbicidal proteins","volume":"37","author":"TL Doering","year":"1999","journal-title":"Med Mycol"},{"key":"ref36","doi-asserted-by":"crossref","first-page":"2043","DOI":"10.1111\/j.1462-5822.2008.01186.x","article-title":"Capsule enlargement in Cryptococcus neoformans confers resistance to oxidative stress suggesting a mechanism for intracellular survival","volume":"10","author":"O Zaragoza","year":"2008","journal-title":"Cell Microbiol"},{"key":"ref37","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1046\/j.1365-2036.1996.22164006.x","article-title":"The role of Helicobacter pylori urease in the pathogenesis of gastritis and peptic ulceration","volume":"10","author":"H Mobley","year":"1996","journal-title":"Aliment Pharmacol Ther"},{"key":"ref38","doi-asserted-by":"crossref","first-page":"504","DOI":"10.1128\/IAI.74.1.504-515.2006","article-title":"Urease Produced by Coccidioides posadasii Contributes to the Virulence of This Respiratory Pathogen","volume":"74","author":"F Mirbod-Donovan","year":"2006","journal-title":"Infect Immun"},{"key":"ref39","doi-asserted-by":"crossref","first-page":"1406","DOI":"10.1111\/febs.13229","article-title":"Cryptococcus gattii urease as a virulence factor and the relevance of enzymatic activity in cryptococcosis pathogenesis","volume":"282","author":"V Feder","year":"2015","journal-title":"FEBS J"},{"key":"ref40","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1128\/IAI.68.2.443-448.2000","article-title":"Urease as a Virulence Factor in Experimental Cryptococcosis","volume":"68","author":"GM Cox","year":"2000","journal-title":"Infect Immun"},{"key":"ref41","doi-asserted-by":"crossref","first-page":"787","DOI":"10.1080\/07391102.2000.10506568","article-title":"Molecular Mechanics Evaluation of the Proposed Mechanisms for the Degradation of Urea by Urease","volume":"17","author":"M Zimmer","year":"2000","journal-title":"J Biomol Struct Dyn"},{"key":"ref42","doi-asserted-by":"crossref","first-page":"10828","DOI":"10.1021\/ja0525399","article-title":"The Burden Borne by Urease","volume":"127","author":"BP Callahan","year":"2005","journal-title":"J Am Chem Soc"},{"key":"ref43","doi-asserted-by":"crossref","first-page":"e1004062","DOI":"10.1371\/journal.ppat.1004062","article-title":"The Emerging Role of Urease as a General Microbial Virulence Factor","volume":"10","author":"JC Rutherford","year":"2014","journal-title":"PLOS Pathog"},{"key":"ref44","doi-asserted-by":"crossref","first-page":"1683","DOI":"10.1172\/JCI41963","article-title":"Real-time imaging of trapping and urease-dependent transmigration of Cryptococcus neoformans in mouse brain","volume":"120","author":"M Shi","year":"2010","journal-title":"J Clin Invest"},{"key":"ref45","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.00220-13","article-title":"Factors Required for Activation of Urease as a Virulence Determinant in Cryptococcus neoformans","volume":"4","author":"A Singh","year":"2013","journal-title":"mBio"},{"key":"ref46","doi-asserted-by":"crossref","first-page":"1761","DOI":"10.1016\/S0002-9440(10)63734-0","article-title":"Urease Expression by Cryptococcus neoformans Promotes Microvascular Sequestration, Thereby Enhancing Central Nervous System Invasion","volume":"164","author":"MA Olszewski","year":"2004","journal-title":"Am J Pathol"},{"key":"ref47","doi-asserted-by":"crossref","first-page":"2470","DOI":"10.1128\/IAI.59.7.2470-2475.1991","article-title":"Essential role of urease in pathogenesis of gastritis induced by Helicobacter pylori in gnotobiotic piglets","volume":"59","author":"KA Eaton","year":"1991","journal-title":"Infect Immun"},{"key":"ref48","doi-asserted-by":"crossref","first-page":"3604","DOI":"10.1128\/IAI.62.9.3604-3607.1994","article-title":"Effect of gastric pH on urease-dependent colonization of gnotobiotic piglets by Helicobacter pylori","volume":"62","author":"KA Eaton","year":"1994","journal-title":"Infect Immun"},{"key":"ref49","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1086\/517814","article-title":"Highly Significant Role of Helicobacter pylori Urease in Phagocytosis and Production of Oxygen Metabolites by Human Granulocytes","volume":"177","author":"A Makristathis","year":"1998","journal-title":"J Infect Dis"},{"key":"ref50","doi-asserted-by":"crossref","first-page":"1214","DOI":"10.1189\/jlb.0106030","article-title":"Role of urease in megasome formation and Helicobacter pylori survival in macrophages","volume":"79","author":"JT Schwartz","year":"2006","journal-title":"J Leukoc Biol"},{"key":"ref51","doi-asserted-by":"crossref","first-page":"6002","DOI":"10.4049\/jimmunol.168.12.6002","article-title":"Cutting Edge: Urease Release by Helicobacter pylori Stimulates Macrophage Inducible Nitric Oxide Synthase","volume":"168","author":"AP Gobert","year":"2002","journal-title":"J Immunol"},{"key":"ref52","doi-asserted-by":"crossref","first-page":"932","DOI":"10.2353\/ajpath.2009.080673","article-title":"Cryptococcal Urease Promotes the Accumulation of Immature Dendritic Cells and a Non-Protective T2 Immune Response within the Lung","volume":"174","author":"JJ Osterholzer","year":"2009","journal-title":"Am J Pathol"},{"key":"ref53","doi-asserted-by":"crossref","first-page":"e1002047","DOI":"10.1371\/journal.ppat.1002047","article-title":"Phospholipids Trigger Cryptococcus neoformans Capsular Enlargement during Interactions with Amoebae and Macrophages","volume":"7","author":"CJ Chrisman","year":"2011","journal-title":"PLOS Pathog"},{"key":"ref54","first-page":"1","article-title":"Recent Advances on Model Hosts","author":"A Casadevall","year":"2012"},{"key":"ref55","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1080\/13693786310001616500","article-title":"Cryptococcus neoformans var. gattii can exploit Acanthamoeba castellanii for growth","volume":"42","author":"SD Malliaris","year":"2004","journal-title":"Med Mycol"},{"key":"ref56","doi-asserted-by":"crossref","first-page":"2059","DOI":"10.1128\/IAI.01503-14","article-title":"Temporal Kinetics and Quantitative Analysis of Cryptococcus neoformans Nonlytic Exocytosis","volume":"82","author":"SA Stukes","year":"2014","journal-title":"Infect Immun"},{"key":"ref57","doi-asserted-by":"crossref","first-page":"e872827","DOI":"10.1155\/2014\/872827","article-title":"Establishing Standards for Studying Renal Function in Mice through Measurements of Body Size-Adjusted Creatinine and Urea Levels","volume":"2014","author":"WF Rodrigues","year":"2014","journal-title":"BioMed Res Int"},{"key":"ref58","doi-asserted-by":"crossref","first-page":"e00945","DOI":"10.1128\/mBio.00945-14","article-title":"Capsule Growth in Cryptococcus neoformans Is Coordinated with Cell Cycle Progression","volume":"5","author":"R Garc\u00eda-Rodas","year":"2014","journal-title":"mBio"},{"key":"ref59","first-page":"2219","article-title":"Cryptococcus neoformans-induced macrophage lysosome damage crucially contributes to fungal virulence","volume":"194","author":"MJ Davis","year":"2015","journal-title":"J Immunol Baltim Md 1950"},{"key":"ref60","article-title":"The Outcome of the Cryptococcus neoformans\u2013Macrophage Interaction Depends on Phagolysosomal Membrane Integrity","author":"CM De Leon-Rodriguez","year":"2018","journal-title":"J Immunol"},{"key":"ref61","first-page":"w13042","article-title":"Cathepsins and their involvement in immune responses","volume":"140","author":"S Conus","year":"2010","journal-title":"Swiss Med Wkly"},{"key":"ref62","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.00167-11","article-title":"Nonlytic Exocytosis of Cryptococcus neoformans from Macrophages Occurs In Vivo and Is Influenced by Phagosomal pH","volume":"2","author":"AM Nicola","year":"2011","journal-title":"mBio"},{"key":"ref63","doi-asserted-by":"crossref","first-page":"1640","DOI":"10.1172\/JCI119688","article-title":"Chloroquine induces human mononuclear phagocytes to inhibit and kill Cryptococcus neoformans by a mechanism independent of iron deprivation","volume":"100","author":"SM Levitz","year":"1997","journal-title":"J Clin Invest"},{"key":"ref64","doi-asserted-by":"crossref","first-page":"413","DOI":"10.1111\/j.1600-0854.2005.00284.x","article-title":"The Kinetics of Phagosome Maturation as a Function of Phagosome\/Lysosome Fusion and Acquisition of Hydrolytic Activity","volume":"6","author":"RM Yates","year":"2005","journal-title":"Traffic"},{"key":"ref65","doi-asserted-by":"crossref","first-page":"677","DOI":"10.1083\/jcb.201507112","article-title":"The position of lysosomes within the cell determines their luminal pH","volume":"212","author":"DE Johnson","year":"2016","journal-title":"J Cell Biol"},{"key":"ref66","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1086\/315649","article-title":"Conditional lethality of the diprotic weak bases chloroquine and quinacrine against Cryptococcus neoformans","volume":"182","author":"TS Harrison","year":"2000","journal-title":"J Infect Dis"},{"key":"ref67","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1111\/j.1574-695X.1996.tb00220.x","article-title":"Helicobacter pylori utilises urea for amino acid synthesis","volume":"13","author":"C l. Williams","year":"1996","journal-title":"FEMS Immunol Med Microbiol"},{"key":"ref68","doi-asserted-by":"crossref","first-page":"2370","DOI":"10.1128\/AEM.02737-07","article-title":"Role of Ureolytic Activity in Bacillus cereus Nitrogen Metabolism and Acid Survival","volume":"74","author":"M Mols","year":"2008","journal-title":"Appl Environ Microbiol"},{"key":"ref69","doi-asserted-by":"crossref","first-page":"504","DOI":"10.1128\/IAI.67.2.504-512.1999","article-title":"Genetic and Physiologic Characterization of Urease of Actinomyces naeslundii","volume":"67","author":"E Morou-Bermudez","year":"1999","journal-title":"Infect Immun"},{"key":"ref70","doi-asserted-by":"crossref","first-page":"e02183","DOI":"10.1128\/mBio.02183-16","article-title":"Trojan Horse Transit Contributes to Blood-Brain Barrier Crossing of a Eukaryotic Pathogen","volume":"8","author":"FH Santiago-Tirado","year":"2017","journal-title":"mBio"},{"key":"ref71","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1016\/j.micinf.2015.08.017","article-title":"Cryptococcal transmigration across a model brain blood-barrier: evidence of the Trojan horse mechanism and differences between Cryptococcus neoformans var. grubii strain H99 and Cryptococcus gattii strain R265","volume":"18","author":"TC Sorrell","year":"2016","journal-title":"Microbes Infect"},{"key":"ref72","doi-asserted-by":"crossref","first-page":"4985","DOI":"10.1128\/IAI.72.9.4985-4995.2004","article-title":"Cryptococcal Yeast Cells Invade the Central Nervous System via Transcellular Penetration of the Blood-Brain Barrier","volume":"72","author":"YC Chang","year":"2004","journal-title":"Infect Immun"},{"key":"ref73","doi-asserted-by":"crossref","first-page":"e35455","DOI":"10.1371\/journal.pone.0035455","article-title":"Capsule Independent Uptake of the Fungal Pathogen Cryptococcus neoformans into Brain Microvascular Endothelial Cells","volume":"7","author":"W Sabiiti","year":"2012","journal-title":"PLOS ONE"},{"key":"ref74","doi-asserted-by":"crossref","first-page":"34761","DOI":"10.1074\/jbc.M111.219378","article-title":"Invasion of Cryptococcus neoformans into human brain microvascular endothelial cells is mediated through the lipid rafts-endocytic pathway via the dual specificity tyrosine phosphorylation-regulated kinase 3 (DYRK3)","volume":"286","author":"S-H Huang","year":"2011","journal-title":"J Biol Chem"},{"key":"ref75","article-title":"Brain Inositol Is a Novel Stimulator for Promoting Cryptococcus Penetration of the Blood-Brain Barrier","volume":"9","author":"T-B Liu","year":"2013","journal-title":"PLoS Pathog"},{"key":"ref76","doi-asserted-by":"crossref","first-page":"3139","DOI":"10.1128\/IAI.00554-13","article-title":"Cryptococcus neoformans Promotes Its Transmigration into the Central Nervous System by Inducing Molecular and Cellular Changes in Brain Endothelial Cells","volume":"81","author":"K Vu","year":"2013","journal-title":"Infect Immun"},{"key":"ref77","article-title":"Invasion of the Central Nervous System by Cryptococcus neoformans Requires a Secreted Fungal Metalloprotease","volume":"5","author":"K Vu","year":"2014","journal-title":"mBio"},{"key":"ref78","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.01425-15","article-title":"Live Imaging of Host-Parasite Interactions in a Zebrafish Infection Model Reveals Cryptococcal Determinants of Virulence and Central Nervous System Invasion","volume":"6","author":"JL Tenor","year":"2015","journal-title":"mBio"},{"key":"ref79","doi-asserted-by":"crossref","first-page":"2729","DOI":"10.1128\/IAI.00094-07","article-title":"The Relative Susceptibility of Mouse Strains to Pulmonary Cryptococcus neoformans Infection Is Associated with Pleiotropic Differences in the Immune Response","volume":"75","author":"O Zaragoza","year":"2007","journal-title":"Infect Immun"},{"key":"ref80","doi-asserted-by":"crossref","first-page":"397","DOI":"10.3171\/jns.2001.94.3.0397","article-title":"Intracerebral microdialysis in neurointensive care: the use of urea as an endogenous reference compound","volume":"94","author":"E Ronne-Engstr\u00f6m","year":"2001","journal-title":"J Neurosurg"},{"key":"ref81","doi-asserted-by":"crossref","first-page":"909","DOI":"10.2215\/CJN.00320106","article-title":"Age-Related Increase in Plasma Urea Level and Decrease in Fractional Urea Excretion: Clinical Application in the Syndrome of Inappropriate Secretion of Antidiuretic Hormone","volume":"1","author":"W Musch","year":"2006","journal-title":"Clin J Am Soc Nephrol"},{"key":"ref82","doi-asserted-by":"crossref","first-page":"1490","DOI":"10.1111\/j.1600-0854.2011.01266.x","article-title":"A Weak Base-Generating System Suitable for Selective Manipulation of Lysosomal pH","volume":"12","author":"LR Carraro-Lacroix","year":"2011","journal-title":"Traffic"},{"key":"ref83","doi-asserted-by":"crossref","first-page":"2771","DOI":"10.1128\/IAI.06195-11","article-title":"Urease Activity Represents an Alternative Pathway for Mycobacterium tuberculosis Nitrogen Metabolism","volume":"80","author":"W Lin","year":"2012","journal-title":"Infect Immun"},{"key":"ref84","doi-asserted-by":"crossref","first-page":"e1003402","DOI":"10.1371\/journal.ppat.1003402","article-title":"Tripping on acid: trans-kingdom perspectives on biological acids in immunity and pathogenesis","volume":"9","author":"MF Criscitiello","year":"2013","journal-title":"PLoS Pathog"},{"key":"ref85","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1007\/s004080000012","article-title":"Bactericidal Activity of Alveolar Macrophages is Suppressed by V-ATPase Inhibition","volume":"178","author":"A Bidani","year":"2000","journal-title":"Lung"},{"key":"ref86","doi-asserted-by":"crossref","first-page":"7039","DOI":"10.1128\/IAI.68.12.7039-7048.2000","article-title":"Phagosome Acidification Has Opposite Effects on Intracellular Survival of Bordetella pertussis andB. bronchiseptica","volume":"68","author":"B Schneider","year":"2000","journal-title":"Infect Immun"},{"key":"ref87","doi-asserted-by":"crossref","first-page":"891","DOI":"10.1128\/IAI.71.2.891-903.2003","article-title":"Phagocytosis and Intracellular Fate of Aspergillus fumigatus Conidia in Alveolar Macrophages","volume":"71","author":"O Ibrahim-Granet","year":"2003","journal-title":"Infect Immun"},{"key":"ref88","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.02580-14","article-title":"Cryptococcus neoformans Host Adaptation: Toward Biological Evidence of Dormancy","volume":"6","author":"A Alanio","year":"2015","journal-title":"mBio"},{"key":"ref89","doi-asserted-by":"crossref","first-page":"593","DOI":"10.1016\/j.chom.2009.05.014","article-title":"Leading a sheltered life: Intracellular pathogens and maintenance of vacuolar compartments","volume":"5","author":"Y Kumar","year":"2009","journal-title":"Cell Host Microbe"},{"key":"ref90","doi-asserted-by":"crossref","first-page":"1088","DOI":"10.1111\/j.1365-2958.2011.07632.x","article-title":"SEC14 is a specific requirement for secretion of phospholipase B1 and pathogenicity of Cryptococcus neoformans","volume":"80","author":"M Chayakulkeeree","year":"2011","journal-title":"Mol Microbiol"},{"key":"ref91","first-page":"1252","article-title":"The Membrane Phospholipid Binding Protein Annexin A2 Promotes Phagocytosis and Non-lytic Exocytosis of Cryptococcus neoformans and Impacts Survival in Fungal Infection","volume":"197","author":"S Stukes","year":"2016","journal-title":"J Immunol Baltim Md 1950"},{"key":"ref92","doi-asserted-by":"crossref","first-page":"e1001041","DOI":"10.1371\/journal.ppat.1001041","article-title":"The Human Fungal Pathogen Cryptococcus neoformans Escapes Macrophages by a Phagosome Emptying Mechanism That Is Inhibited by Arp2\/3 Complex-Mediated Actin Polymerisation","volume":"6","author":"SA Johnston","year":"2010","journal-title":"PLOS Pathog"},{"key":"ref93","doi-asserted-by":"crossref","first-page":"973","DOI":"10.1016\/S0006-3495(89)82896-6","article-title":"Fusion of synaptic vesicle membranes with planar bilayer membranes","volume":"55","author":"MS Perin","year":"1989","journal-title":"Biophys J"},{"key":"ref94","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1128\/IAI.64.1.319-325.1996","article-title":"Intracellular fate of Mycobacterium avium: use of dual-label spectrofluorometry to investigate the influence of bacterial viability and opsonization on phagosomal pH and phagosome-lysosome interaction","volume":"64","author":"YK Oh","year":"1996","journal-title":"Infect Immun"},{"key":"ref95","doi-asserted-by":"crossref","first-page":"827","DOI":"10.1189\/jlb.0513251","article-title":"Hv1 proton channels differentially regulate the pH of neutrophil and macrophage phagosomes by sustaining the production of phagosomal ROS that inhibit the delivery of vacuolar ATPases","volume":"95","author":"AE Chemaly","year":"2014","journal-title":"J Leukoc Biol"},{"key":"ref96","first-page":"e53402","article-title":"Measuring Phagosome pH by Ratiometric Fluorescence Microscopy","author":"P Nunes","year":"2015","journal-title":"JoVE J Vis Exp"},{"key":"ref97","doi-asserted-by":"crossref","first-page":"1309","DOI":"10.1016\/j.cell.2015.08.027","article-title":"Pathogen Cell-to-cell Variability Drives Heterogeneity In Host Immune Responses","volume":"162","author":"R Avraham","year":"2015","journal-title":"Cell"},{"key":"ref98","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0163505","article-title":"Hyperspectral Imaging Using Intracellular Spies: Quantitative Real-Time Measurement of Intracellular Parameters In Vivo during Interaction of the Pathogenic Fungus Aspergillus fumigatus with Human Monocytes","volume":"11","author":"S Mohebbi","year":"2016","journal-title":"PLoS ONE"},{"key":"ref99","doi-asserted-by":"crossref","first-page":"1605","DOI":"10.1084\/jem.177.6.1605","article-title":"Histoplasma capsulatum modulates the acidification of phagolysosomes","volume":"177","author":"LG Eissenberg","year":"1993","journal-title":"J Exp Med"},{"key":"ref100","doi-asserted-by":"crossref","first-page":"584","DOI":"10.1128\/jcm.7.6.584-588.1978","article-title":"Rapid urea broth test for yeasts","volume":"7","author":"GD Roberts","year":"1978","journal-title":"J Clin Microbiol"},{"key":"ref101","doi-asserted-by":"crossref","first-page":"1751","DOI":"10.1128\/IAI.55.8.1751-1754.1987","article-title":"Urease inhibition by EDTA in the two varieties of Cryptococcus neoformans","volume":"55","author":"KJ Kwon-Chung","year":"1987","journal-title":"Infect Immun"},{"key":"ref102","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1038\/laban0511-155","article-title":"Retro-orbital injections in mice","volume":"40","author":"T Yardeni","year":"2011","journal-title":"Lab Anim"}],"updated-by":[{"DOI":"10.1371\/journal.ppat.1007144","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2018,6,27]],"date-time":"2018-06-27T00:00:00Z","timestamp":1530057600000}}],"container-title":["PLOS Pathogens"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/dx.plos.org\/10.1371\/journal.ppat.1007144","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,7,5]],"date-time":"2025-07-05T04:29:25Z","timestamp":1751689765000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.ppat.1007144"}},"subtitle":[],"editor":[{"given":"Robin Charles","family":"May","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2018,6,15]]},"references-count":102,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2018,6,15]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.ppat.1007144","relation":{},"ISSN":["1553-7374"],"issn-type":[{"value":"1553-7374","type":"electronic"}],"subject":[],"published":{"date-parts":[[2018,6,15]]}}}