{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,9]],"date-time":"2026-04-09T09:38:20Z","timestamp":1775727500134,"version":"3.50.1"},"reference-count":54,"publisher":"MDPI AG","issue":"6","license":[{"start":{"date-parts":[[2021,6,20]],"date-time":"2021-06-20T00:00:00Z","timestamp":1624147200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["EXPL\/BIA-MIC\/2525\/2013"],"award-info":[{"award-number":["EXPL\/BIA-MIC\/2525\/2013"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["IF\/00124\/2015"],"award-info":[{"award-number":["IF\/00124\/2015"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"name":"COMPETE2020","award":["LISBOA-01-0145-FEDER-007660"],"award-info":[{"award-number":["LISBOA-01-0145-FEDER-007660"]}]},{"name":"European Union\u2019s Horizon 2020 research and innovation programme","award":["810856"],"award-info":[{"award-number":["810856"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Microorganisms"],"abstract":"<jats:p>In yeast, iron storage and detoxification depend on the Ccc1 transporter that mediates iron accumulation in vacuoles. While deletion of the CCC1 gene renders cells unable to survive under iron overload conditions, the deletion of its previously identified regulators only partially affects survival, indicating that the mechanisms controlling iron storage and detoxification in yeast are still far from well understood. This work reveals that CCC1 is equipped with a complex transcriptional structure comprising several regulatory regions. One of these is located inside the coding sequence of the gene and drives the expression of a short transcript encoding an N-terminally truncated protein, designated as s-Ccc1. s-Ccc1, though less efficiently than Ccc1, is able to promote metal accumulation in the vacuole, protecting cells against iron toxicity. While the expression of the s-Ccc1 appears to be repressed in the normal genomic context, our current data clearly demonstrates that it is functional and has the capacity to play a role under iron overload conditions.<\/jats:p>","DOI":"10.3390\/microorganisms9061337","type":"journal-article","created":{"date-parts":[[2021,6,20]],"date-time":"2021-06-20T21:50:15Z","timestamp":1624225815000},"page":"1337","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":4,"title":["An Internal Promoter Drives the Expression of a Truncated Form of CCC1 Capable of Protecting Yeast from Iron Toxicity"],"prefix":"10.3390","volume":"9","author":[{"given":"Catarina","family":"Amaral","sequence":"first","affiliation":[{"name":"ITQB NOVA, Instituto de Tecnologia Qu\u00edmica e Biol\u00f3gica Ant\u00f3nio Xavier, Universidade Nova de Lisboa, Av. da Rep\u00fablica, 2780-157 Oeiras, Portugal"}]},{"given":"Cristina Teixeira","family":"Vicente","sequence":"additional","affiliation":[{"name":"ITQB NOVA, Instituto de Tecnologia Qu\u00edmica e Biol\u00f3gica Ant\u00f3nio Xavier, Universidade Nova de Lisboa, Av. da Rep\u00fablica, 2780-157 Oeiras, Portugal"}]},{"given":"Soraia Marques","family":"Caetano","sequence":"additional","affiliation":[{"name":"ITQB NOVA, Instituto de Tecnologia Qu\u00edmica e Biol\u00f3gica Ant\u00f3nio Xavier, Universidade Nova de Lisboa, Av. da Rep\u00fablica, 2780-157 Oeiras, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0248-8965","authenticated-orcid":false,"given":"Ana","family":"Gaspar-Cordeiro","sequence":"additional","affiliation":[{"name":"ITQB NOVA, Instituto de Tecnologia Qu\u00edmica e Biol\u00f3gica Ant\u00f3nio Xavier, Universidade Nova de Lisboa, Av. da Rep\u00fablica, 2780-157 Oeiras, Portugal"}]},{"given":"Yang","family":"Yang","sequence":"additional","affiliation":[{"name":"ID16A Beamline, ESRF-The European Synchrotron, 38043 Grenoble, France"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4129-9091","authenticated-orcid":false,"given":"Peter","family":"Cloetens","sequence":"additional","affiliation":[{"name":"ID16A Beamline, ESRF-The European Synchrotron, 38043 Grenoble, France"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9474-2405","authenticated-orcid":false,"given":"C\u00e9lia V.","family":"Rom\u00e3o","sequence":"additional","affiliation":[{"name":"ITQB NOVA, Instituto de Tecnologia Qu\u00edmica e Biol\u00f3gica Ant\u00f3nio Xavier, Universidade Nova de Lisboa, Av. da Rep\u00fablica, 2780-157 Oeiras, Portugal"}]},{"given":"Claudina","family":"Rodrigues-Pousada","sequence":"additional","affiliation":[{"name":"ITQB NOVA, Instituto de Tecnologia Qu\u00edmica e Biol\u00f3gica Ant\u00f3nio Xavier, Universidade Nova de Lisboa, Av. da Rep\u00fablica, 2780-157 Oeiras, Portugal"}]},{"given":"Catarina","family":"Pimentel","sequence":"additional","affiliation":[{"name":"ITQB NOVA, Instituto de Tecnologia Qu\u00edmica e Biol\u00f3gica Ant\u00f3nio Xavier, Universidade Nova de Lisboa, Av. da Rep\u00fablica, 2780-157 Oeiras, Portugal"}]}],"member":"1968","published-online":{"date-parts":[[2021,6,20]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"783","DOI":"10.1016\/S0891-5849(97)00016-6","article-title":"Iron Homeostasis, Oxidative Stress, and DNA Damage","volume":"23","author":"Meneghini","year":"1997","journal-title":"Free. Radic. Biol. Med."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"4536","DOI":"10.1021\/cr9001676","article-title":"Iron Acquisition and Transcriptional Regulation","volume":"109","author":"Kaplan","year":"2009","journal-title":"Chem. Rev."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1007\/s11274-017-2215-8","article-title":"Mechanisms of iron sensing and regulation in the yeast Saccharomyces cerevisiae","volume":"33","author":"Puig","year":"2017","journal-title":"World J. Microbiol. Biotechnol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/j.cell.2010.06.028","article-title":"Two to Tango: Regulation of Mammalian Iron Metabolism","volume":"142","author":"Hentze","year":"2010","journal-title":"Cell"},{"key":"ref_5","first-page":"241","article-title":"The Iron Metallome in Eukaryotic Organisms","volume":"12","author":"Dlouhy","year":"2013","journal-title":"Met. Ions Life Sci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"2980","DOI":"10.1091\/mbc.e06-11-1054","article-title":"Mechanism Underlying the Iron-dependent Nuclear Export of the Iron-responsive Transcription Factor Aft1p in Saccharomyces cerevisiae","volume":"18","author":"Ueta","year":"2007","journal-title":"Mol. Biol. Cell"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"18914","DOI":"10.1074\/jbc.M200949200","article-title":"Subcellular Localization of Aft1 Transcription Factor Responds to Iron Status in Saccharomyces cerevisiae","volume":"277","author":"Ueta","year":"2002","journal-title":"J. Biol. Chem."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1016\/j.cmet.2008.04.010","article-title":"Cooperation of Two mRNA-Binding Proteins Drives Metabolic Adaptation to Iron Deficiency","volume":"7","author":"Puig","year":"2008","journal-title":"Cell Metab."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"14823","DOI":"10.1074\/jbc.M109.091710","article-title":"Metabolic Response to Iron Deficiency in Saccharomyces cerevisiae","volume":"285","author":"Protchenko","year":"2010","journal-title":"J. Biol. Chem."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1233","DOI":"10.1091\/mbc.e03-09-0642","article-title":"Transcriptional Remodeling in Response to Iron Deprivation inSaccharomyces cerevisiae","volume":"15","author":"Tiedeman","year":"2004","journal-title":"Mol. Biol. Cell"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"22181","DOI":"10.1074\/jbc.M414663200","article-title":"Post-transcriptional Regulation of the Yeast High Affinity Iron Transport System","volume":"280","author":"Felice","year":"2005","journal-title":"J. Biol. Chem."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1021\/bi3015339","article-title":"Iron Content ofSaccharomyces cerevisiaeCells Grown under Iron-Deficient and Iron-Overload Conditions","volume":"52","author":"Jhurry","year":"2013","journal-title":"Biochemistry"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/j.molcel.2005.05.021","article-title":"Multiple RNA Surveillance Pathways Limit Aberrant Expression of Iron Uptake mRNAs and Prevent Iron Toxicity in S. cerevisiae","volume":"19","author":"Lee","year":"2005","journal-title":"Mol. Cell"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"10709","DOI":"10.1074\/jbc.275.14.10709","article-title":"Desferrioxamine-mediated Iron Uptake in Saccharomyces cerevisiae","volume":"275","author":"Yun","year":"2000","journal-title":"J. Biol. Chem."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1111\/j.1432-1033.1995.00337.x","article-title":"Iron sequestration by the yeast vacuole. A study with vacuolar mutants of Saccharomyces cerevisiae","volume":"228","author":"Bode","year":"1995","journal-title":"Eur. J. Biochem."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1016\/0014-5793(88)80742-7","article-title":"Iron storage in Saccharomyces cerevisiae","volume":"231","author":"Raguzzi","year":"1988","journal-title":"Febs Lett."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"29515","DOI":"10.1074\/jbc.M103944200","article-title":"CCC1 Is a Transporter That Mediates Vacuolar Iron Storage in Yeast","volume":"276","author":"Li","year":"2001","journal-title":"J. Biol. Chem."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1262","DOI":"10.1039\/c2mt20179h","article-title":"The interplay between vacuolar and siderophore-mediated iron storage in Aspergillus fumigatus","volume":"4","author":"Gsaller","year":"2012","journal-title":"Metallom. Integr. Biomet. Sci."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"28129","DOI":"10.1074\/jbc.R114.592618","article-title":"Lysosome-related Organelles as Mediators of Metal Homeostasis","volume":"289","author":"Merchant","year":"2014","journal-title":"J. Biol. Chem."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"2434","DOI":"10.1104\/pp.17.00672","article-title":"Wheat Vacuolar Iron Transporter TaVIT2 Transports Fe and Mn and Is Effective for Biofortification","volume":"174","author":"Connorton","year":"2017","journal-title":"Plant. Physiol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1295","DOI":"10.1126\/science.1132563","article-title":"Localization of Iron in Arabidopsis Seed Requires the Vacuolar Membrane Transporter VIT1","volume":"314","author":"Kim","year":"2006","journal-title":"Science"},{"key":"ref_22","first-page":"308","article-title":"Crystal structure of plant vacuolar iron transporter Vitnat","volume":"5","author":"Kato","year":"2019","journal-title":"Plants"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"16498","DOI":"10.1074\/jbc.M114.571562","article-title":"A Bacterial Iron Exporter for Maintenance of Iron Homeostasis","volume":"289","author":"Sankari","year":"2014","journal-title":"J. Biol. Chem."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"863","DOI":"10.1099\/mic.0.076802-0","article-title":"Roles of Agrobacterium tumefaciens membrane-bound ferritin (MbfA) in iron transport and resistance to iron under acidic conditions","volume":"160","author":"Bhubhanil","year":"2014","journal-title":"Microbiology"},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Huang, G., Ulrich, P.N., Storey, M., Johnson, D., Tischer, J., Tovar, J.A., Moreno, S.N.J., Orlando, R., and Docampo, R. (2014). Proteomic Analysis of the Acidocalcisome, an Organelle Conserved from Bacteria to Human Cells. Plos Pathog., 10.","DOI":"10.1371\/journal.ppat.1004555"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"10403","DOI":"10.1038\/ncomms10403","article-title":"A vacuolar iron-transporter homologue acts as a detoxifier in Plasmodium","volume":"7","author":"Slavic","year":"2016","journal-title":"Nat. Commun."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"400","DOI":"10.1111\/j.1365-313X.2012.05088.x","article-title":"Vacuolar membrane transporters OsVIT1 and OsVIT2 modulate iron translocation between flag leaves and seeds in rice","volume":"72","author":"Zhang","year":"2012","journal-title":"Plant J."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"370","DOI":"10.1128\/MCB.01033-14","article-title":"The Basic Leucine Zipper Stress Response Regulator Yap5 Senses High-Iron Conditions by Coordination of [2Fe-2S] Clusters","volume":"35","author":"Rietzschel","year":"2015","journal-title":"Mol. Cell. Biol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1326","DOI":"10.1128\/MCB.01219-07","article-title":"Yap5 Is an Iron-Responsive Transcriptional Activator That Regulates Vacuolar Iron Storage in Yeast","volume":"28","author":"Li","year":"2008","journal-title":"Mol. Cell. Biol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"35709","DOI":"10.1074\/jbc.M112.395533","article-title":"A Role for Iron-Sulfur Clusters in the Regulation of Transcription Factor Yap5-dependent High Iron Transcriptional Responses in Yeast","volume":"287","author":"Li","year":"2012","journal-title":"J. Biol. Chem."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Pimentel, C., Vicente, C., Menezes, R., Caetano, S., Carreto, L., and Rodrigues-Pousada, C. (2012). The Role of the Yap5 Transcription Factor in Remodeling Gene Expression in Response to Fe Bioavailability. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0037434"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"15577","DOI":"10.1074\/jbc.M117.802504","article-title":"The glucose sensor Snf1 and the transcription factors Msn2 and Msn4 regulate transcription of the vacuolar iron importer gene CCC1 and iron resistance in yeast","volume":"292","author":"Li","year":"2017","journal-title":"J. Biol. Chem."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"2519","DOI":"10.1093\/nar\/24.13.2519","article-title":"A new efficient gene disruption cassette for repeated use in budding yeast","volume":"24","author":"Guldener","year":"1996","journal-title":"Nucl. Acids Res."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"472","DOI":"10.1016\/j.bbagrm.2017.02.001","article-title":"Mediator, SWI\/SNF and SAGA complexes regulate Yap8-dependent transcriptional activation of ACR2 in response to arsenate","volume":"1860","author":"Menezes","year":"2017","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Morawe, C.B.R., Cloetens, P., Lantelme, B., Peffen, J.-C., and Vivo, A. (2015, January 26). Graded multilayers for figured Kirkpatrick-Baez mirrors on the new ESRF end station ID16A. Proceedings of the Advances in X-ray\/Euv Optics and Components, San Diego, CA, USA.","DOI":"10.1117\/12.2189279"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/j.sab.2006.12.002","article-title":"A multiplatform code for the analysis of energy-dispersive X-ray fluorescence spectra","volume":"62","author":"Papillon","year":"2007","journal-title":"Spectrochim. Acta Part B Spectrosc."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"D348","DOI":"10.1093\/nar\/gkx842","article-title":"YEASTRACT: An upgraded database for the analysis of transcription regulatory networks in Saccharomyces cerevisiae","volume":"46","author":"Teixeira","year":"2018","journal-title":"Nucleic Acids Res."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1671","DOI":"10.1016\/j.bbapap.2013.05.022","article-title":"RAPID: Fast and accurate sequence-based prediction of intrinsic disorder content on proteomic scale","volume":"1834","author":"Yan","year":"2013","journal-title":"Biochim. Biophys. Acta Proteins Proteom."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"6979","DOI":"10.1007\/s00216-014-8004-4","article-title":"Correlative organelle fluorescence microscopy and synchrotron X-ray chemical element imaging in single cells","volume":"406","author":"Roudeau","year":"2014","journal-title":"Anal. Bioanal. Chem."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1166","DOI":"10.1128\/EC.00077-07","article-title":"Saccharomyces cerevisiae Vacuole in Zinc Storage and Intracellular Zinc Distribution","volume":"6","author":"Simm","year":"2007","journal-title":"Eukaryot. Cell"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"7810","DOI":"10.1038\/ncomms8810","article-title":"The development and characterization of synthetic minimal yeast promoters","volume":"6","author":"Redden","year":"2015","journal-title":"Nat. Commun."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"5712","DOI":"10.1021\/pr300538f","article-title":"Amino Termini of Many Yeast Proteins Map to Downstream Start Codons","volume":"11","author":"Fournier","year":"2012","journal-title":"J. Proteome Res."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"2891","DOI":"10.1021\/acs.jproteome.5b00996","article-title":"N-Terminal Peptide Detection with Optimized Peptide-Spectrum Matching and Streamlined Sequence Libraries","volume":"15","author":"Lycette","year":"2016","journal-title":"J. Proteome Res."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1974","DOI":"10.1101\/gr.243378.118","article-title":"Chromatin-sensitive cryptic promoters putatively drive expression of alternative protein isoforms in yeast","volume":"29","author":"Wei","year":"2019","journal-title":"Genome Res."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"977","DOI":"10.1101\/gr.150342.112","article-title":"Roles for transcript leaders in translation and mRNA decay revealed by transcript leader sequencing","volume":"23","author":"Arribere","year":"2013","journal-title":"Genome Res."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"5160","DOI":"10.1016\/S0021-9258(19)89228-1","article-title":"Structure of yeast LEUThe 5\u2019 flanking region contains features that predict two modes of control and two productive translation starts","volume":"261","author":"Beltzer","year":"1986","journal-title":"J. Biol. Chem."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/0092-8674(82)90384-1","article-title":"Two differentially regulated mRNAs with different 5\u2032 ends encode secreted and intracellular forms of yeast invertase","volume":"28","author":"Carlson","year":"1982","journal-title":"Cell"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1016\/S0021-9258(19)57354-9","article-title":"The yeast VAS1 gene encodes both mitochondrial and cytoplasmic valyl-tRNA synthetases","volume":"263","author":"Chatton","year":"1988","journal-title":"J. Biol. Chem."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"987","DOI":"10.1093\/genetics\/132.4.987","article-title":"Hts1 Encodes Both the Cytoplasmic and Mitochondrial Histidyl-Trna Synthetase of Saccharomyces Cerevisiae: Mutations Alter the Specificity of Compartmentation","volume":"132","author":"Chiu","year":"1992","journal-title":"Genetics"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"5172","DOI":"10.1073\/pnas.84.15.5172","article-title":"Amino-terminal extension generated from an upstream AUG codon is not required for mitochondrial import of yeast N2,N2-dimethylguanosine-specific tRNA methyltransferase","volume":"84","author":"Ellis","year":"1987","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"190083","DOI":"10.1098\/rsob.190083","article-title":"Tales of tails in transporters","volume":"9","author":"Mikros","year":"2019","journal-title":"Open Biol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1128\/MCB.19.1.817","article-title":"The Two Forms of Karyogamy Transcription Factor Kar4p Are Regulated by Differential Initiation of Transcription, Translation, and Protein Turnover","volume":"19","author":"Gammie","year":"1999","journal-title":"Mol. Cell. Biol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"858","DOI":"10.15252\/msb.20156662","article-title":"Positional proteomics reveals differences in N-terminal proteoform stability","volume":"12","author":"Gawron","year":"2016","journal-title":"Mol. Syst. Biol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"3712","DOI":"10.1016\/j.csbj.2020.10.044","article-title":"Structure and function of the vacuolar Ccc1\/VIT1 family of iron transporters and its regulation in fungi","volume":"18","author":"Peris","year":"2020","journal-title":"Comput. Struct. Biotechnol. J."}],"container-title":["Microorganisms"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2607\/9\/6\/1337\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T06:19:46Z","timestamp":1760163586000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2607\/9\/6\/1337"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,6,20]]},"references-count":54,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2021,6]]}},"alternative-id":["microorganisms9061337"],"URL":"https:\/\/doi.org\/10.3390\/microorganisms9061337","relation":{},"ISSN":["2076-2607"],"issn-type":[{"value":"2076-2607","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,6,20]]}}}