{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,20]],"date-time":"2026-02-20T03:18:40Z","timestamp":1771557520874,"version":"3.50.1"},"update-to":[{"DOI":"10.1371\/journal.ppat.1010729","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2022,8,11]],"date-time":"2022-08-11T00:00:00Z","timestamp":1660176000000}}],"reference-count":109,"publisher":"Public Library of Science (PLoS)","issue":"8","license":[{"start":{"date-parts":[[2022,8,1]],"date-time":"2022-08-01T00:00:00Z","timestamp":1659312000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/"}],"funder":[{"DOI":"10.13039\/501100001843","name":"Science and Engineering Research Board","doi-asserted-by":"publisher","award":["CRG\/2018\/000847\/HS"],"award-info":[{"award-number":["CRG\/2018\/000847\/HS"]}],"id":[{"id":"10.13039\/501100001843","id-type":"DOI","asserted-by":"publisher"}]},{"name":"JC Bose Fellowship","award":["SB\/S2\/JCB-012\/2015"],"award-info":[{"award-number":["SB\/S2\/JCB-012\/2015"]}]},{"DOI":"10.13039\/100006492","name":"Division of Intramural Research, National Institute of Allergy and Infectious Diseases","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100006492","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plospathogens.org"],"crossmark-restriction":false},"short-container-title":["PLoS Pathog"],"abstract":"<jats:p>Reversible protein phosphorylation at serine\/threonine residues is one of the most common protein modifications, widely observed in all kingdoms of life. The catalysts controlling this modification are specific serine\/threonine kinases and phosphatases that modulate various cellular pathways ranging from growth to cellular death. Genome sequencing and various omics studies have led to the identification of numerous serine\/threonine kinases and cognate phosphatases, yet the physiological relevance of many of these proteins remain enigmatic. In <jats:italic>Bacillus anthracis<\/jats:italic>, only one ser\/thr phosphatase, PrpC, has been functionally characterized; it was reported to be non-essential for bacterial growth and survival. In the present study, we characterized another ser\/thr phosphatase (PrpN) of <jats:italic>B<\/jats:italic>. <jats:italic>anthracis<\/jats:italic> by various structural and functional approaches. To examine its physiological relevance in <jats:italic>B<\/jats:italic>. <jats:italic>anthracis<\/jats:italic>, a null mutant strain of <jats:italic>prpN<\/jats:italic> was generated and shown to have defects in sporulation and reduced synthesis of toxins (PA and LF) and the toxin activator protein AtxA. We also identified CodY, a global transcriptional regulator, as a target of PrpN and ser\/thr kinase PrkC. CodY phosphorylation strongly controlled its binding to the promoter region of <jats:italic>atxA<\/jats:italic>, as shown using phosphomimetic and phosphoablative mutants. In nutshell, the present study reports phosphorylation-mediated regulation of CodY activity in the context of anthrax toxin synthesis in <jats:italic>B<\/jats:italic>. <jats:italic>anthracis<\/jats:italic> by a previously uncharacterized ser\/thr protein phosphatase\u2013PrpN.<\/jats:p>","DOI":"10.1371\/journal.ppat.1010729","type":"journal-article","created":{"date-parts":[[2022,8,1]],"date-time":"2022-08-01T19:09:12Z","timestamp":1659380952000},"page":"e1010729","update-policy":"https:\/\/doi.org\/10.1371\/journal.ppat.corrections_policy","source":"Crossref","is-referenced-by-count":12,"title":["Role of serine\/threonine protein phosphatase PrpN in the life cycle of Bacillus anthracis"],"prefix":"10.1371","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7828-4427","authenticated-orcid":true,"given":"Aakriti","family":"Gangwal","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Nitika","family":"Sangwan","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Neha","family":"Dhasmana","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Nishant","family":"Kumar","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Chetkar Chandra","family":"Keshavam","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Lalit 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H.","family":"Leppla","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3902-4355","authenticated-orcid":true,"given":"Yogendra","family":"Singh","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"340","published-online":{"date-parts":[[2022,8,1]]},"reference":[{"issue":"8","key":"ppat.1010729.ref001","doi-asserted-by":"crossref","first-page":"1337","DOI":"10.1038\/s41564-019-0435-4","article-title":"The global distribution of Bacillus anthracis and associated anthrax risk to humans, livestock and wildlife","volume":"4","author":"CJ Carlson","year":"2019","journal-title":"Nat Microbiol"},{"issue":"2","key":"ppat.1010729.ref002","doi-asserted-by":"crossref","first-page":"359","DOI":"10.20506\/rst.21.2.1336","article-title":"Anthrax and wildlife","volume":"21","author":"ME Hugh-Jones","year":"2002","journal-title":"Rev Sci 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Dis"},{"issue":"1","key":"ppat.1010729.ref012","doi-asserted-by":"crossref","first-page":"e84947","DOI":"10.1371\/journal.pone.0084947","article-title":"Toxin-independent virulence of Bacillus anthracis in rabbits","volume":"9","author":"H Levy","year":"2014","journal-title":"PLoS One"},{"issue":"12","key":"ppat.1010729.ref013","doi-asserted-by":"crossref","first-page":"1333","DOI":"10.1164\/rccm.201102-0209CI","article-title":"Anthrax infection","volume":"184","author":"DA Sweeney","year":"2011","journal-title":"Am J Respir Crit Care Med"},{"issue":"1","key":"ppat.1010729.ref014","doi-asserted-by":"crossref","first-page":"371","DOI":"10.1128\/iai.39.1.371-376.1983","article-title":"Evidence for plasmid-mediated toxin production in Bacillus anthracis","volume":"39","author":"P Mikesell","year":"1983","journal-title":"Infect Immun"},{"issue":"2","key":"ppat.1010729.ref015","first-page":"557","article-title":"Virulence and immunogenicity in experimental animals of Bacillus anthracis strains harbouring or lacking 110 MDa and 60 MDa plasmids","volume":"132","author":"I Uchida","year":"1986","journal-title":"J Gen Microbiol"},{"issue":"20","key":"ppat.1010729.ref016","doi-asserted-by":"crossref","first-page":"6509","DOI":"10.1128\/JB.181.20.6509-6515.1999","article-title":"Sequence and organization of pXO1, the large Bacillus anthracis plasmid harboring the anthrax toxin genes","volume":"181","author":"RT Okinaka","year":"1999","journal-title":"J Bacteriol"},{"key":"ppat.1010729.ref017","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1146\/annurev.cellbio.19.111301.140655","article-title":"Anthrax toxin","volume":"19","author":"RJ Collier","year":"2003","journal-title":"Annu Rev Cell Dev Biol"},{"issue":"7","key":"ppat.1010729.ref018","doi-asserted-by":"crossref","DOI":"10.3390\/toxins12070464","article-title":"Anthrax Edema and Lethal Toxins Differentially Target Human Lung and Blood Phagocytes","volume":"12","author":"VI Patel","year":"2020","journal-title":"Toxins (Basel)"},{"issue":"16","key":"ppat.1010729.ref019","doi-asserted-by":"crossref","first-page":"5188","DOI":"10.1128\/jb.176.16.5188-5192.1994","article-title":"The three Bacillus anthracis toxin genes are coordinately regulated by bicarbonate and temperature","volume":"176","author":"JC Sirard","year":"1994","journal-title":"J Bacteriol"},{"issue":"2","key":"ppat.1010729.ref020","doi-asserted-by":"crossref","first-page":"370","DOI":"10.1128\/JB.184.2.370-380.2002","article-title":"Control of anthrax toxin gene expression by the transition state regulator abrB","volume":"184","author":"E Saile","year":"2002","journal-title":"J Bacteriol"},{"issue":"3","key":"ppat.1010729.ref021","doi-asserted-by":"crossref","first-page":"761","DOI":"10.1128\/iai.45.3.761-767.1984","article-title":"Immunoelectrophoretic analysis, toxicity, and kinetics of in vitro production of the protective antigen and lethal factor components of Bacillus anthracis toxin","volume":"45","author":"JW Ezzell","year":"1984","journal-title":"Infect Immun"},{"issue":"9","key":"ppat.1010729.ref022","doi-asserted-by":"crossref","first-page":"735","DOI":"10.1016\/j.resmic.2010.09.006","article-title":"AtxA, a Bacillus anthracis global virulence regulator","volume":"161","author":"A. Fouet","year":"2010","journal-title":"Res Microbiol"},{"issue":"2","key":"ppat.1010729.ref023","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1128\/JB.186.2.307-315.2004","article-title":"atxA controls Bacillus anthracis capsule synthesis via acpA and a newly discovered regulator, acpB","volume":"186","author":"M Drysdale","year":"2004","journal-title":"J Bacteriol"},{"issue":"23","key":"ppat.1010729.ref024","doi-asserted-by":"crossref","DOI":"10.1128\/JB.00569-19","article-title":"Bacillus anthracis Virulence Regulator AtxA Binds Specifically to the pagA Promoter Region","volume":"201","author":"RM McCall","year":"2019","journal-title":"J Bacteriol"},{"issue":"17","key":"ppat.1010729.ref025","doi-asserted-by":"crossref","first-page":"5329","DOI":"10.1128\/jb.175.17.5329-5338.1993","article-title":"Cloning and characterization of a gene whose product is a trans-activator of anthrax toxin synthesis","volume":"175","author":"I Uchida","year":"1993","journal-title":"J Bacteriol"},{"issue":"6","key":"ppat.1010729.ref026","doi-asserted-by":"crossref","first-page":"1171","DOI":"10.1111\/j.1365-2958.1995.tb02340.x","article-title":"The atxA gene product activates transcription of the anthrax toxin genes and is essential for virulence","volume":"16","author":"Z Dai","year":"1995","journal-title":"Mol Microbiol"},{"key":"ppat.1010729.ref027","doi-asserted-by":"crossref","first-page":"482","DOI":"10.3389\/fmicb.2018.00482","article-title":"A Dual Role for the Bacillus anthracis Master Virulence Regulator AtxA: Control of Sporulation and Anthrax Toxin Production","volume":"9","author":"JL Dale","year":"2018","journal-title":"Front Microbiol"},{"issue":"Pt 6","key":"ppat.1010729.ref028","doi-asserted-by":"crossref","first-page":"1751","DOI":"10.1099\/mic.0.27803-0","article-title":"The DNA-binding specificity of the Bacillus anthracis AbrB protein","volume":"151","author":"MA Strauch","year":"2005","journal-title":"Microbiology (Reading)"},{"issue":"6","key":"ppat.1010729.ref029","doi-asserted-by":"crossref","first-page":"1204","DOI":"10.1128\/JB.02041-12","article-title":"Identification of CodY targets in Bacillus anthracis by genome-wide in vitro binding analysis","volume":"195","author":"A Chateau","year":"2013","journal-title":"J Bacteriol"},{"issue":"10","key":"ppat.1010729.ref030","doi-asserted-by":"crossref","first-page":"4437","DOI":"10.1128\/IAI.00716-09","article-title":"The global regulator CodY regulates toxin gene expression in Bacillus anthracis and is required for full virulence","volume":"77","author":"W van Schaik","year":"2009","journal-title":"Infect Immun"},{"issue":"14","key":"ppat.1010729.ref031","doi-asserted-by":"crossref","first-page":"9460","DOI":"10.1074\/jbc.R114.557926","article-title":"In the beginning, there was protein phosphorylation","volume":"289","author":"JM Kyriakis","year":"2014","journal-title":"J Biol Chem"},{"issue":"3\u20134","key":"ppat.1010729.ref032","first-page":"125","article-title":"Ser\/Thr\/Tyr protein phosphorylation in bacteria\u2014for long time neglected, now well established","volume":"9","author":"J Deutscher","year":"2005","journal-title":"J Mol Microbiol Biotechnol"},{"issue":"3\u20134","key":"ppat.1010729.ref033","first-page":"198","article-title":"Role of protein phosphorylation on serine\/threonine and tyrosine in the virulence of bacterial pathogens","volume":"9","author":"AJ Cozzone","year":"2005","journal-title":"J Mol Microbiol Biotechnol"},{"key":"ppat.1010729.ref034","doi-asserted-by":"crossref","first-page":"184","DOI":"10.3389\/fmicb.2016.00184","article-title":"Role of Protein Phosphorylation in the Regulation of Cell Cycle and DNA-Related Processes in Bacteria","volume":"7","author":"T Garcia-Garcia","year":"2016","journal-title":"Front Microbiol"},{"issue":"4385","key":"ppat.1010729.ref035","doi-asserted-by":"crossref","first-page":"1111","DOI":"10.1126\/science.34215","article-title":"Phosphorylation of Isocitrate dehydrogenase of Escherichia coli","volume":"203","author":"M Garnak","year":"1979","journal-title":"Science"},{"key":"ppat.1010729.ref036","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1016\/S0021-9258(18)76142-5","article-title":"Serinephosphoric acid obtained on hydrolysis of vitellinic acid","volume":"98","author":"FA Lipmann","year":"1932","journal-title":"J Biol Chem"},{"issue":"4332","key":"ppat.1010729.ref037","doi-asserted-by":"crossref","first-page":"804","DOI":"10.1038\/170804b0","article-title":"Isolation of phosphothreonine from bovine casein","volume":"170","author":"CH De Verdier","year":"1952","journal-title":"Nature"},{"key":"ppat.1010729.ref038","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1038\/s41522-017-0015-4","article-title":"Ser\/Thr protein kinase PrkC-mediated regulation of GroEL is critical for biofilm formation in Bacillus anthracis","volume":"3","author":"G Arora","year":"2017","journal-title":"NPJ Biofilms Microbiomes"},{"issue":"22","key":"ppat.1010729.ref039","doi-asserted-by":"crossref","first-page":"8930","DOI":"10.1074\/jbc.RA118.005424","article-title":"The Ser\/Thr protein kinase PrkC imprints phenotypic memory in Bacillus anthracis spores by phosphorylating the glycolytic enzyme enolase","volume":"294","author":"R Virmani","year":"2019","journal-title":"J Biol Chem"},{"issue":"4","key":"ppat.1010729.ref040","doi-asserted-by":"crossref","first-page":"520","DOI":"10.1007\/s12088-018-0744-y","article-title":"GroEL Mediates Folding of Bacillus anthracis Serine\/Threonine Protein Kinase, PrkC","volume":"58","author":"R Virmani","year":"2018","journal-title":"Indian J Microbiol"},{"key":"ppat.1010729.ref041","article-title":"Bacillus anthracis chain length, a virulence determinant, is regulated by membrane localized serine\/threonine protein kinase PrkC","author":"N Dhasmana","year":"2021","journal-title":"J Bacteriol"},{"issue":"5","key":"ppat.1010729.ref042","doi-asserted-by":"crossref","first-page":"715","DOI":"10.1007\/s10534-013-9646-y","article-title":"Zinc regulates the activity of kinase-phosphatase pair (BasPrkC\/BasPrpC) in Bacillus anthracis","volume":"26","author":"G Arora","year":"2013","journal-title":"Biometals"},{"issue":"32","key":"ppat.1010729.ref043","doi-asserted-by":"crossref","first-page":"26749","DOI":"10.1074\/jbc.M112.351304","article-title":"Unveiling the novel dual specificity protein kinases in Bacillus anthracis: identification of the first prokaryotic dual specificity tyrosine phosphorylation-regulated kinase (DYRK)-like kinase","volume":"287","author":"G Arora","year":"2012","journal-title":"J Biol Chem"},{"issue":"8","key":"ppat.1010729.ref044","doi-asserted-by":"crossref","first-page":"1833","DOI":"10.1128\/JB.01401-10","article-title":"Autoregulatory characteristics of a Bacillus anthracis serine\/threonine kinase","volume":"193","author":"KM Bryant-Hudson","year":"2011","journal-title":"J Bacteriol"},{"issue":"2","key":"ppat.1010729.ref045","doi-asserted-by":"crossref","first-page":"400","DOI":"10.1128\/JB.01221-09","article-title":"Regulatory interactions of a virulence-associated serine\/threonine phosphatase-kinase pair in Bacillus anthracis","volume":"192","author":"SM Shakir","year":"2010","journal-title":"J Bacteriol"},{"issue":"6","key":"ppat.1010729.ref046","doi-asserted-by":"crossref","first-page":"1527","DOI":"10.1111\/j.1365-2958.2010.07182.x","article-title":"ExsB, an unusually highly phosphorylated protein required for the stable attachment of the exosporium of Bacillus anthracis","volume":"76","author":"SA McPherson","year":"2010","journal-title":"Mol Microbiol"},{"issue":"16","key":"ppat.1010729.ref047","doi-asserted-by":"crossref","first-page":"5134","DOI":"10.1128\/JB.00422-09","article-title":"A new minimal 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Wang","year":"2012","journal-title":"PLoS Pathog"},{"issue":"1","key":"ppat.1010729.ref054","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1128\/MMBR.00031-15","article-title":"Staying in Shape: the Impact of Cell Shape on Bacterial Survival in Diverse Environments","volume":"80","author":"DC Yang","year":"2016","journal-title":"Microbiol Mol Biol Rev"},{"issue":"1","key":"ppat.1010729.ref055","doi-asserted-by":"crossref","first-page":"17910","DOI":"10.1038\/s41598-020-74609-5","article-title":"Cell morphology maintenance in Bacillus subtilis through balanced peptidoglycan synthesis and hydrolysis","volume":"10","author":"J Sassine","year":"2020","journal-title":"Sci Rep"},{"issue":"8","key":"ppat.1010729.ref056","doi-asserted-by":"crossref","first-page":"601","DOI":"10.1038\/nrmicro1205","article-title":"Bacterial cell shape","volume":"3","author":"MT Cabeen","year":"2005","journal-title":"Nat Rev 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Errington","year":"2003","journal-title":"Nat Rev Microbiol"},{"issue":"3","key":"ppat.1010729.ref065","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1128\/MMBR.64.3.503-514.2000","article-title":"Programmed death in bacteria","volume":"64","author":"K. Lewis","year":"2000","journal-title":"Microbiol Mol Biol Rev"},{"issue":"12","key":"ppat.1010729.ref066","doi-asserted-by":"crossref","first-page":"931","DOI":"10.1016\/j.micres.2014.05.001","article-title":"Life cycle and spore resistance of spore-forming Bacillus atrophaeus","volume":"169","author":"SR Sella","year":"2014","journal-title":"Microbiol Res"},{"issue":"4","key":"ppat.1010729.ref067","doi-asserted-by":"crossref","first-page":"306","DOI":"10.1111\/j.1472-765X.2012.03209.x","article-title":"Comparative analysis of virulence factors secreted by Bacillus anthracis Sterne at host body temperature","volume":"54","author":"SK Kim","year":"2012","journal-title":"Lett Appl Microbiol"},{"issue":"4","key":"ppat.1010729.ref068","doi-asserted-by":"crossref","first-page":"1649","DOI":"10.1128\/IAI.00530-08","article-title":"Killed but metabolically active Bacillus anthracis vaccines induce broad and protective immunity against anthrax","volume":"77","author":"J Skoble","year":"2009","journal-title":"Infect Immun"},{"key":"ppat.1010729.ref069","doi-asserted-by":"crossref","first-page":"18908","DOI":"10.1038\/srep18908","article-title":"Next-Generation Bacillus anthracis Live Attenuated Spore Vaccine Based on the htrA(-) (High Temperature Requirement A) Sterne Strain","volume":"6","author":"T Chitlaru","year":"2016","journal-title":"Sci Rep"},{"issue":"5","key":"ppat.1010729.ref070","doi-asserted-by":"crossref","first-page":"714","DOI":"10.1007\/s00284-016-1120-y","article-title":"Late-Exponential Gene Expression in codY-Deficient Bacillus anthracis in a Host-Like Environment","volume":"73","author":"SK Kim","year":"2016","journal-title":"Curr Microbiol"},{"key":"ppat.1010729.ref071","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1186\/s12864-016-3004-8","article-title":"Changes in Bacillus anthracis CodY regulation under host-specific environmental factor deprived conditions","volume":"17","author":"SK Kim","year":"2016","journal-title":"BMC Genomics"},{"issue":"3","key":"ppat.1010729.ref072","doi-asserted-by":"crossref","first-page":"672","DOI":"10.1016\/j.bbrc.2015.12.019","article-title":"Overexpression of the pleiotropic regulator CodY decreases sporulation, attachment and pellicle formation in Bacillus anthracis","volume":"469","author":"M Gopalani","year":"2016","journal-title":"Biochem Biophys Res Commun"},{"issue":"12","key":"ppat.1010729.ref073","doi-asserted-by":"crossref","first-page":"4445","DOI":"10.1096\/fj.11-188912","article-title":"CodY regulation is required for full virulence and heme iron acquisition in Bacillus anthracis","volume":"25","author":"A Chateau","year":"2011","journal-title":"FASEB J"},{"issue":"4","key":"ppat.1010729.ref074","doi-asserted-by":"crossref","first-page":"697","DOI":"10.1074\/mcp.M600464-MCP200","article-title":"The serine\/threonine\/tyrosine phosphoproteome of the model bacterium Bacillus subtilis","volume":"6","author":"B Macek","year":"2007","journal-title":"Mol Cell Proteomics"},{"issue":"12","key":"ppat.1010729.ref075","doi-asserted-by":"crossref","first-page":"4127","DOI":"10.1128\/JB.187.12.4127-4139.2005","article-title":"A region of Bacillus subtilis CodY protein required for interaction with DNA","volume":"187","author":"P Joseph","year":"2005","journal-title":"J Bacteriol"},{"issue":"7","key":"ppat.1010729.ref076","doi-asserted-by":"crossref","first-page":"2714","DOI":"10.1074\/jbc.M116.754309","article-title":"Structure of the Branched-chain Amino Acid and GTP-sensing Global Regulator, CodY, from Bacillus subtilis","volume":"292","author":"VM Levdikov","year":"2017","journal-title":"J Biol Chem"},{"issue":"3","key":"ppat.1010729.ref077","doi-asserted-by":"crossref","first-page":"798","DOI":"10.1128\/JB.01115-07","article-title":"Interaction of Bacillus subtilis CodY with GTP","volume":"190","author":"LD Handke","year":"2008","journal-title":"J Bacteriol"},{"issue":"6","key":"ppat.1010729.ref078","doi-asserted-by":"crossref","first-page":"1911","DOI":"10.1128\/JB.185.6.1911-1922.2003","article-title":"Additional targets of the Bacillus subtilis global regulator CodY identified by chromatin immunoprecipitation and genome-wide transcript analysis","volume":"185","author":"V Molle","year":"2003","journal-title":"J Bacteriol"},{"issue":"2","key":"ppat.1010729.ref079","doi-asserted-by":"crossref","first-page":"599","DOI":"10.1111\/j.1365-2958.2004.04135.x","article-title":"Activation of the Bacillus subtilis global regulator CodY by direct interaction with branched-chain amino acids","volume":"53","author":"RP Shivers","year":"2004","journal-title":"Mol Microbiol"},{"issue":"11","key":"ppat.1010729.ref080","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1038\/s41579-019-0243-0","article-title":"Protein post-translational modifications in bacteria","volume":"17","author":"B Macek","year":"2019","journal-title":"Nat Rev 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Dworkin","year":"2015","journal-title":"Curr Opin Microbiol"},{"key":"ppat.1010729.ref084","doi-asserted-by":"crossref","first-page":"107622","DOI":"10.1016\/j.pharmthera.2020.107622","article-title":"Metal-dependent Ser\/Thr protein phosphatase PPM family: Evolution, structures, diseases and inhibitors","volume":"215","author":"R Kamada","year":"2020","journal-title":"Pharmacol Ther"},{"issue":"5","key":"ppat.1010729.ref085","doi-asserted-by":"crossref","first-page":"217","DOI":"10.15698\/mic2019.05.677","article-title":"Ser\/Thr protein phosphatases in fungi: structure, regulation and function","volume":"6","author":"J Arino","year":"2019","journal-title":"Microb Cell"},{"issue":"2","key":"ppat.1010729.ref086","doi-asserted-by":"crossref","first-page":"324","DOI":"10.1111\/j.1742-4658.2012.08609.x","article-title":"Protein Ser\/Thr phosphatases\u2014the ugly ducklings of cell signalling","volume":"280","author":"DL Brautigan","year":"2013","journal-title":"FEBS J"},{"issue":"2","key":"ppat.1010729.ref087","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1016\/j.molbiopara.2008.06.008","article-title":"Protein Ser\/Thr phosphatases of parasitic protozoa","volume":"161","author":"MA Kutuzov","year":"2008","journal-title":"Mol Biochem Parasitol"},{"issue":"10","key":"ppat.1010729.ref088","doi-asserted-by":"crossref","first-page":"2175","DOI":"10.1016\/j.cellsig.2014.06.013","article-title":"Ser\/Thr-phosphoprotein phosphatases in chondrogenesis: neglected components of a two-player game","volume":"26","author":"C Matta","year":"2014","journal-title":"Cell Signal"},{"issue":"3","key":"ppat.1010729.ref089","doi-asserted-by":"crossref","first-page":"756","DOI":"10.1111\/febs.15454","article-title":"Plant protein phosphatases: What do we know about their mechanism of action?","volume":"288","author":"M Bheri","year":"2021","journal-title":"FEBS J"},{"key":"ppat.1010729.ref090","doi-asserted-by":"crossref","first-page":"527","DOI":"10.1146\/annurev-micro-020415-111342","article-title":"Protein Phosphatases of Pathogenic Bacteria: Role in Physiology and Virulence","volume":"69","author":"A Sajid","year":"2015","journal-title":"Annu Rev Microbiol"},{"issue":"4","key":"ppat.1010729.ref091","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1006\/scbi.1995.0028","article-title":"Flicking the switches: phosphorylation of serine\/threonine protein phosphatases","volume":"6","author":"DL Brautigan","year":"1995","journal-title":"Semin Cancer Biol"},{"issue":"46","key":"ppat.1010729.ref092","doi-asserted-by":"crossref","first-page":"24215","DOI":"10.1074\/jbc.M116.754531","article-title":"Serine\/Threonine Protein Phosphatase PstP of Mycobacterium tuberculosis Is Necessary for Accurate Cell Division and Survival of Pathogen","volume":"291","author":"AK Sharma","year":"2016","journal-title":"J Biol Chem"},{"issue":"22","key":"ppat.1010729.ref093","doi-asserted-by":"crossref","first-page":"6109","DOI":"10.1128\/JB.184.22.6109-6114.2002","article-title":"The PrpC serine-threonine phosphatase and PrkC kinase have opposing physiological roles in stationary-phase Bacillus subtilis cells","volume":"184","author":"TA Gaidenko","year":"2002","journal-title":"J Bacteriol"},{"issue":"9","key":"ppat.1010729.ref094","doi-asserted-by":"crossref","first-page":"1720","DOI":"10.1099\/mic.0.000148","article-title":"Serine\/threonine\/tyrosine phosphorylation regulates DNA binding of bacterial transcriptional regulators","volume":"161","author":"A Kalantari","year":"2015","journal-title":"Microbiology (Reading)"},{"issue":"3","key":"ppat.1010729.ref095","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1038\/s41564-019-0648-6","article-title":"Transcriptional regulation by sigma factor phosphorylation in bacteria","volume":"5","author":"SC Iyer","year":"2020","journal-title":"Nat Microbiol"},{"issue":"4","key":"ppat.1010729.ref096","first-page":"206","article-title":"Transcription regulators potentially controlled by HPr kinase\/phosphorylase in Gram-negative bacteria","volume":"5","author":"G Boel","year":"2003","journal-title":"J Mol Microbiol Biotechnol"},{"key":"ppat.1010729.ref097","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1146\/annurev-micro-102215-095331","article-title":"Feedback Control of Two-Component Regulatory Systems","volume":"70","author":"EA Groisman","year":"2016","journal-title":"Annu Rev Microbiol"},{"issue":"4","key":"ppat.1010729.ref098","doi-asserted-by":"crossref","DOI":"10.1093\/femspd\/ftx048","article-title":"Biochemical characterization of the GTP-sensing protein, CodY of Bacillus anthracis","volume":"75","author":"S Joon","year":"2017","journal-title":"Pathog Dis"},{"issue":"5","key":"ppat.1010729.ref099","doi-asserted-by":"crossref","first-page":"1129","DOI":"10.1111\/mmi.12617","article-title":"Phosphorylation of Bacillus subtilis gene regulator AbrB modulates its DNA-binding properties","volume":"92","author":"A Kobir","year":"2014","journal-title":"Mol Microbiol"},{"issue":"10","key":"ppat.1010729.ref100","doi-asserted-by":"crossref","first-page":"e0184975","DOI":"10.1371\/journal.pone.0184975","article-title":"The CodY-dependent clhAB2 operon is involved in cell shape, chaining and autolysis in Bacillus cereus ATCC 14579","volume":"12","author":"E Huillet","year":"2017","journal-title":"PLoS One"},{"issue":"8","key":"ppat.1010729.ref101","doi-asserted-by":"crossref","first-page":"2233","DOI":"10.1111\/j.1462-2920.2012.02766.x","article-title":"CodY, a pleiotropic regulator, influences multicellular behaviour and efficient production of virulence factors in Bacillus cereus","volume":"14","author":"T Lindback","year":"2012","journal-title":"Environ Microbiol"},{"issue":"2","key":"ppat.1010729.ref102","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1111\/j.1574-695X.2011.00812.x","article-title":"The CodY pleiotropic repressor controls virulence in gram-positive pathogens","volume":"62","author":"L Stenz","year":"2011","journal-title":"FEMS Immunol Med Microbiol"},{"issue":"32","key":"ppat.1010729.ref103","doi-asserted-by":"crossref","first-page":"19103","DOI":"10.1016\/S0021-9258(19)47273-6","article-title":"A deleted variant of Bacillus anthracis protective antigen is non-toxic and blocks anthrax toxin action in vivo","volume":"264","author":"Y Singh","year":"1989","journal-title":"J Biol Chem"},{"issue":"14","key":"ppat.1010729.ref104","doi-asserted-by":"crossref","first-page":"2646","DOI":"10.1128\/JB.01625-14","article-title":"HupB, a nucleoid-associated protein of Mycobacterium tuberculosis, is modified by serine\/threonine protein kinases in vivo","volume":"196","author":"M Gupta","year":"2014","journal-title":"J Bacteriol"},{"issue":"22","key":"ppat.1010729.ref105","doi-asserted-by":"crossref","DOI":"10.21769\/BioProtoc.970","article-title":"Separation and Detection of Phosphorylated and Nonphosphorylated BvgA, a Bordetella pertussis Response Regulator, in vivo and in vitro","volume":"3","author":"Q Chen","year":"2013","journal-title":"Bio Protoc"},{"issue":"3","key":"ppat.1010729.ref106","doi-asserted-by":"crossref","first-page":"855","DOI":"10.1111\/1462-2920.12548","article-title":"clpC operon regulates cell architecture and sporulation in Bacillus anthracis","volume":"17","author":"LK Singh","year":"2015","journal-title":"Environ Microbiol"},{"issue":"2","key":"ppat.1010729.ref107","doi-asserted-by":"crossref","first-page":"170","DOI":"10.1007\/s12088-021-00927-5","article-title":"ClpC-Mediated Sporulation Regulation at Engulfment Stage in Bacillus anthracis","volume":"61","author":"N Kumar","year":"2021","journal-title":"Indian J 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