{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T12:38:50Z","timestamp":1792067930016,"version":"4.4.0"},"reference-count":107,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2020,5,21]],"date-time":"2020-05-21T00:00:00Z","timestamp":1590019200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Microbiol."],"DOI":"10.3389\/fmicb.2020.01028","type":"journal-article","created":{"date-parts":[[2020,5,21]],"date-time":"2020-05-21T05:14:30Z","timestamp":1590038070000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":153,"title":["Persistence of Staphylococcus aureus: Multiple Metabolic Pathways Impact the Expression of Virulence Factors in Small-Colony Variants (SCVs)"],"prefix":"10.3389","volume":"11","author":[{"given":"Lorena","family":"Tuchscherr","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bettina","family":"L\u00f6ffler","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Richard A.","family":"Proctor","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2020,5,21]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.00331-18","article-title":"Genome plasticity of agr-defective Staphylococcus aureus during clinical infection.","volume":"86","author":"Altman","year":"2018","journal-title":"Infect. Immun."},{"key":"B2","first-page":"126","article-title":"Staphylococcus aureus: methicillin resistance and small colony variants from pyogenic infections of skin, soft tissue and bone.","volume":"13","author":"Ansari","year":"2015","journal-title":"J. Nepal Health Res. Counc."},{"key":"B3","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1038\/s41579-019-0196-3","article-title":"Definitions and guidelines for research on antibiotic persistence.","volume":"17","author":"Balaban","year":"2019","journal-title":"Nat. Rev. Microbiol."},{"key":"B4","doi-asserted-by":"publisher","first-page":"1033","DOI":"10.1093\/infdis\/170.4.1033","article-title":"Gentamicin-resistant menadione and hemin auxotrophic Staphylococcus aureus persist within cultured endothelial cells.","volume":"170","author":"Balwit","year":"1994","journal-title":"J. Infect. Dis."},{"key":"B5","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-018-21925-6","article-title":"Fatty acid supplementation reverses the small colony variant phenotype in Triclosan-adapted Staphylococcus aureus: genetic, proteomic and phenotypic analyses.","volume":"8","author":"Bazaid","year":"2018","journal-title":"Sci. Rep."},{"key":"B6","doi-asserted-by":"publisher","first-page":"475","DOI":"10.4103\/0976-9668.160046","article-title":"Breast abscess in a case of duct atresia caused by CO2-auxotrophic small colony variants of Staphylococcus aureus: case report and review of literature.","volume":"6","author":"Bhattacharyya","year":"2015","journal-title":"J. Nat. Sci. Biol. Med."},{"key":"B7","doi-asserted-by":"publisher","first-page":"497","DOI":"10.1016\/s0140-6736(00)74210-3","article-title":"Treatment of staphylococcal infections with penicillin by intermittent sterilisation.","volume":"244","author":"Bigger","year":"1944","journal-title":"Lancet"},{"key":"B8","doi-asserted-by":"publisher","first-page":"320","DOI":"10.1038\/nrmicro.2016.34","article-title":"Distinguishing between resistance, tolerance and persistence to antibiotic treatment.","volume":"14","author":"Brauner","year":"2016","journal-title":"Nat. Rev. Microbiol."},{"key":"B9","doi-asserted-by":"publisher","first-page":"417","DOI":"10.1007\/s00294-016-0656-5","article-title":"CodY, a master integrator of metabolism and virulence in Gram-positive bacteria.","volume":"63","author":"Brinsmade","year":"2017","journal-title":"Curr. Genet."},{"key":"B10","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1042\/EBC20160061","article-title":"Antibiotic tolerance and the alternative lifestyles of Staphylococcus aureus.","volume":"61","author":"Bui","year":"2017","journal-title":"Essays Biochem."},{"key":"B11","doi-asserted-by":"publisher","first-page":"345","DOI":"10.1016\/j.meegid.2015.10.011","article-title":"A full genomic characterization of the development of a stable small colony variant cell-type by a clinical Staphylococcus aureus strain.","volume":"36","author":"Bui","year":"2015","journal-title":"Infect. Genet. Evol."},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1128\/mBio.00358-17","article-title":"Alternative evolutionary pathways for drug-resistant small colony variant mutants in Staphylococcus aureus.","volume":"8","author":"Cao","year":"2017","journal-title":"MBio"},{"key":"B13","doi-asserted-by":"publisher","first-page":"4488","DOI":"10.1128\/jb.187.13.4488-4496.2005","article-title":"Staphylococcus aureus ClpC is required for stress resistance, aconitase activity, growth recovery, and death.","volume":"187","author":"Chatterjee","year":"2005","journal-title":"J. Bacteriol."},{"key":"B14","doi-asserted-by":"publisher","first-page":"2936","DOI":"10.1128\/jb.01444-06","article-title":"Enhanced post-stationary-phase survival of a clinical thymidine-dependent small-colony variant of Staphylococcus aureus results from lack of a functional tricarboxylic acid cycle.","volume":"189","author":"Chatterjee","year":"2007","journal-title":"J. Bacteriol."},{"key":"B15","doi-asserted-by":"publisher","first-page":"603","DOI":"10.1093\/jac\/39.5.603","article-title":"Decreased susceptibility to antibiotic killing of a stable small colony variant of Staphylococcus aureus in fluid phase and on fibronectin-coated surfaces.","volume":"39","author":"Chuard","year":"1997","journal-title":"J. Antimicrob. Chemother."},{"key":"B16","doi-asserted-by":"publisher","first-page":"405","DOI":"10.1099\/00221287-131-2-405","article-title":"A comparison of the patterns of extracellular proteins produced by the high alpha-toxin-secreting organism Staphylococcus aureus (Wood 46) during aerobic and anaerobic growth.","volume":"131","author":"Coleman","year":"1985","journal-title":"J. Gen. Microbiol."},{"key":"B17","doi-asserted-by":"publisher","first-page":"365","DOI":"10.1038\/nature12790","article-title":"Activated ClpP kills persisters and eradicates a chronic biofilm infection.","volume":"503","author":"Conlon","year":"2013","journal-title":"Nature"},{"key":"B18","doi-asserted-by":"crossref","DOI":"10.1038\/nmicrobiol.2016.51","article-title":"Persister formation in Staphylococcus aureus is associated with ATP depletion.","volume":"1","author":"Conlon","year":"2016","journal-title":"Nat. Microbiol."},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0054293","article-title":"CcpA-independent glucose regulation of lactate dehydrogenase 1 in Staphylococcus aureus.","volume":"8","author":"Crooke","year":"2013","journal-title":"PLoS One"},{"key":"B20","doi-asserted-by":"publisher","first-page":"E1647","DOI":"10.1073\/pnas.1204307109","article-title":"Coordinated phenotype switching with large-scale chromosome flip-flop inversion observed in bacteria.","volume":"109","author":"Cui","year":"2012","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B21","doi-asserted-by":"publisher","first-page":"2283","DOI":"10.1128\/AAC.01786-09","article-title":"Altered growth, pigmentation, and antimicrobial susceptibility properties of Staphylococcus aureus due to loss of the major cold shock gene cspB.","volume":"54","author":"Duval","year":"2010","journal-title":"Antimicrob. Agents Chemother."},{"key":"B22","doi-asserted-by":"publisher","first-page":"5404","DOI":"10.1128\/JB.00948-12","article-title":"Phenotype switching is a natural consequence of Staphylococcus aureus replication.","volume":"194","author":"Edwards","year":"2012","journal-title":"J. Bacteriol."},{"key":"B23","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1242\/jeb.200.2.217","article-title":"Coupling H+ transport and ATP synthesis in F1F0-ATP synthases: glimpses of interacting parts in a dynamic molecular machine.","volume":"200","author":"Fillingame","year":"1997","journal-title":"J. Exp. Biol."},{"key":"B24","doi-asserted-by":"publisher","first-page":"453","DOI":"10.1038\/nrmicro.2017.42","article-title":"Persistent bacterial infections and persister cells.","volume":"15","author":"Fisher","year":"2017","journal-title":"Nat. Rev. Microbiol."},{"key":"B25","doi-asserted-by":"publisher","first-page":"4809","DOI":"10.1128\/AAC.00352-15","article-title":"Altered competitive fitness, antimicrobial susceptibility, and cellular morphology in a Triclosan-induced small-colony variant of Staphylococcus aureus.","volume":"59","author":"Forbes","year":"2015","journal-title":"Antimicrob. Agents Chemother."},{"key":"B26","doi-asserted-by":"publisher","first-page":"142","DOI":"10.1016\/j.ijmm.2013.11.009","article-title":"Clp chaperones and proteases are central in stress survival, virulence and antibiotic resistance of Staphylococcus aureus.","volume":"304","author":"Frees","year":"2014","journal-title":"Int. J. Med. Microbiol."},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.1099\/mgen.0.000026","article-title":"Large tandem chromosome expansions facilitate niche adaptation during persistent infection with drug-resistant Staphylococcus aureus.","volume":"1","author":"Gao","year":"2015","journal-title":"Microb. Genom."},{"key":"B28","doi-asserted-by":"publisher","first-page":"1455","DOI":"10.1093\/jac\/dkt072","article-title":"Antibiotic activity against small-colony variants of Staphylococcus aureus: review of in vitro, animal and clinical data.","volume":"68","author":"Garcia","year":"2013","journal-title":"J. Antimicrob. Chemother."},{"key":"B29","doi-asserted-by":"publisher","first-page":"2878","DOI":"10.1128\/JCM.00520-10","article-title":"Clinical and molecular characteristics of infections with CO2-dependent small-colony variants of Staphylococcus aureus.","volume":"48","author":"Gomez-Gonzalez","year":"2010","journal-title":"J. Clin. Microbiol."},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2017.01917","article-title":"Generation of persister cells of Pseudomonas aeruginosa and Staphylococcus aureus by chemical treatment and evaluation of their susceptibility to membrane-targeting agents.","volume":"8","author":"Grassi","year":"2017","journal-title":"Front. Microbiol."},{"key":"B31","doi-asserted-by":"publisher","first-page":"20135","DOI":"10.1073\/pnas.1904861116","article-title":"Unstable chromosome rearrangements in Staphylococcus aureus cause phenotype switching associated with persistent infections.","volume":"116","author":"Guerillot","year":"2019","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B32","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-52731-3","article-title":"ClpC affects the intracellular survival capacity of Staphylococcus aureus in non-professional phagocytic cells.","volume":"9","author":"Gunaratnam","year":"2019","journal-title":"Sci. Rep."},{"key":"B33","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1016\/s0140-6736(49)90596-6","article-title":"Acute osteomyelitis due to a penicillinase-producing Staphylococcus aureus.","volume":"1","author":"Jeffery","year":"1949","journal-title":"Lancet"},{"key":"B34","doi-asserted-by":"publisher","first-page":"401","DOI":"10.1128\/CMR.00069-15","article-title":"Clinical significance and pathogenesis of staphylococcal small colony variants in persistent infections.","volume":"29","author":"Kahl","year":"2016","journal-title":"Clin. Microbiol. Rev."},{"key":"B35","doi-asserted-by":"publisher","first-page":"4119","DOI":"10.1128\/iai.73.7.4119-4126.2005","article-title":"Thymidine-dependent Staphylococcus aureus small-colony variants are associated with extensive alterations in regulator and virulence gene expression profiles.","volume":"73","author":"Kahl","year":"2005","journal-title":"Infect. Immun."},{"key":"B36","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1128\/jcm.4.4.343-348.1976","article-title":"Growth requirements of some small-colony-forming variants of Staphylococcus aureus.","volume":"4","author":"Kaplan","year":"1976","journal-title":"J. Clin. Microbiol."},{"key":"B37","doi-asserted-by":"crossref","first-page":"538","DOI":"10.1089\/mdr.2015.0262","article-title":"Small-colony variants in persistent and recurrent Staphylococcus aureus bacteremia.","volume":"22","author":"Kim","year":"2016","journal-title":"Microb. Drug Resist."},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.1128\/mBio.00696-13","article-title":"The Staphylococcus aureus SrrAB two-component system promotes resistance to nitrosative stress and hypoxia.","volume":"4","author":"Kinkel","year":"2013","journal-title":"MBio"},{"key":"B39","doi-asserted-by":"publisher","first-page":"1000","DOI":"10.1136\/jnnp.74.7.1000","article-title":"Detection of Staphylococcus aureus by 16S rRNA directed in situ hybridisation in a patient with a brain abscess caused by small colony variants.","volume":"74","author":"Kipp","year":"2003","journal-title":"J. Neurol. Neurosurg. Psychiatry"},{"key":"B40","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1002\/emmm.200900002","article-title":"The antibiotic ADEP reprogrammes ClpP, switching it from a regulated to an uncontrolled protease.","volume":"1","author":"Kirstein","year":"2009","journal-title":"EMBO Mol. Med."},{"key":"B41","doi-asserted-by":"publisher","first-page":"6351","DOI":"10.1128\/JB.00505-08","article-title":"A defect in menadione biosynthesis induces global changes in gene expression in Staphylococcus aureus.","volume":"190","author":"Kohler","year":"2008","journal-title":"J. Bacteriol."},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.1128\/mBio.01447-14","article-title":"Inactivation of thyA in Staphylococcus aureus attenuates virulence and has a strong impact on metabolism and virulence gene expression.","volume":"5","author":"Kriegeskorte","year":"2014","journal-title":"MBio"},{"key":"B43","doi-asserted-by":"publisher","first-page":"1883","DOI":"10.1099\/jmm.0.062752-0","article-title":"First isolation of oleate-dependent Enterococcus faecalis small-colony variants from the umbilical exudate of a paediatric patient with omphalitis.","volume":"62","author":"Kubota","year":"2013","journal-title":"J. Med. Microbiol."},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.1038\/s41426-018-0205-z","article-title":"Emergence of a dalbavancin induced glycopeptide\/lipoglycopeptide non-susceptible Staphylococcus aureus during treatment of a cardiac device-related endocarditis.","volume":"7","author":"Kussmann","year":"2018","journal-title":"Emerg. Microbes Infect."},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.1128\/AAC.02197-17","article-title":"Tomatidine is a lead antibiotic molecule that targets Staphylococcus aureus ATP synthase subunit C.","volume":"62","author":"Lamontagne Boulet","year":"2018","journal-title":"Antimicrob. Agents Chemother."},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0028366","article-title":"Genetic complexity of fusidic acid-resistant small colony variants (SCV) in Staphylococcus aureus.","volume":"6","author":"Lannergard","year":"2011","journal-title":"PLoS One"},{"key":"B47","doi-asserted-by":"publisher","first-page":"2059","DOI":"10.1128\/AAC.00871-08","article-title":"Genetic determinants of resistance to fusidic acid among clinical bacteremia isolates of Staphylococcus aureus.","volume":"53","author":"Lannergard","year":"2009","journal-title":"Antimicrob. Agents Chemother."},{"key":"B48","doi-asserted-by":"publisher","first-page":"1968","DOI":"10.1128\/IAI.01214-08","article-title":"Capsule expression and genotypic differences among Staphylococcus aureus isolates from patients with chronic or acute osteomyelitis.","volume":"77","author":"Lattar","year":"2009","journal-title":"Infect. Immun."},{"key":"B49","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1159\/000342449","article-title":"Staphylococcus aureus persisters tolerant to bactericidal antibiotics.","volume":"22","author":"Lechner","year":"2012","journal-title":"J. Mol. Microbiol. Biotechnol."},{"key":"B50","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1038\/nrmicro1557","article-title":"Persister cells, dormancy and infectious disease.","volume":"5","author":"Lewis","year":"2007","journal-title":"Nat. Rev. Microbiol."},{"key":"B51","doi-asserted-by":"publisher","first-page":"1807","DOI":"10.1093\/jac\/dkw060","article-title":"Emergence of a small colony variant of vancomycin-intermediate Staphylococcus aureus in a patient with septic arthritis during long-term treatment with daptomycin.","volume":"71","author":"Lin","year":"2016","journal-title":"J. Antimicrob. Chemother."},{"key":"B52","doi-asserted-by":"publisher","first-page":"170","DOI":"10.1016\/j.ijmm.2013.11.011","article-title":"Staphylococcus aureus persistence in non-professional phagocytes.","volume":"304","author":"Loffler","year":"2014","journal-title":"Int. J. Med. Microbiol."},{"key":"B53","doi-asserted-by":"publisher","first-page":"2109","DOI":"10.1038\/s41564-019-0536-0","article-title":"Bacterial metabolic state more accurately predicts antibiotic lethality than growth rate.","volume":"4","author":"Lopatkin","year":"2019","journal-title":"Nat. Microbiol."},{"key":"B54","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2019.00363","article-title":"Staphylococcus aureus small colony variants (SCVs): news from a chronic prosthetic joint infection.","volume":"9","author":"Loss","year":"2019","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"B55","doi-asserted-by":"publisher","first-page":"18","DOI":"10.1016\/j.micpath.2009.10.003","article-title":"A role for sigma factor B in the emergence of Staphylococcus aureus small-colony variants and elevated biofilm production resulting from an exposure to aminoglycosides.","volume":"48","author":"Mitchell","year":"","journal-title":"Microb. Pathog."},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0065018","article-title":"SigB is a dominant regulator of virulence in Staphylococcus aureus small-colony variants.","volume":"8","author":"Mitchell","year":"2013","journal-title":"PLoS One"},{"key":"B57","doi-asserted-by":"publisher","first-page":"1937","DOI":"10.1128\/AAC.01468-10","article-title":"Tomatidine inhibits replication of Staphylococcus aureus small-colony variants in cystic fibrosis airway epithelial cells.","volume":"55","author":"Mitchell","year":"2011","journal-title":"Antimicrob. Agents Chemother."},{"key":"B58","doi-asserted-by":"publisher","first-page":"559","DOI":"10.1093\/jac\/dkr510","article-title":"Tomatidine acts in synergy with aminoglycoside antibiotics against multiresistant Staphylococcus aureus and prevents virulence gene expression.","volume":"67","author":"Mitchell","year":"2012","journal-title":"J. Antimicrob. Chemother."},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2180-10-33","article-title":"Staphylococcus aureus sigma B-dependent emergence of small-colony variants and biofilm production following exposure to Pseudomonas aeruginosa 4-hydroxy-2-heptylquinoline-N-oxide.","volume":"10","author":"Mitchell","year":"","journal-title":"BMC Microbiol."},{"key":"B60","doi-asserted-by":"publisher","first-page":"4438","DOI":"10.1128\/aac.00328-07","article-title":"Genetic and phenotypic identification of fusidic acid-resistant mutants with the small-colony-variant phenotype in Staphylococcus aureus.","volume":"51","author":"Norstrom","year":"2007","journal-title":"Antimicrob. Agents Chemother."},{"key":"B61","doi-asserted-by":"publisher","first-page":"541","DOI":"10.1146\/annurev.genet.42.110807.091640","article-title":"Quorum sensing in Staphylococci.","volume":"42","author":"Novick","year":"2008","journal-title":"Annu. Rev. Genet."},{"key":"B62","doi-asserted-by":"publisher","first-page":"3967","DOI":"10.1002\/j.1460-2075.1993.tb06074.x","article-title":"Synthesis of staphylococcal virulence factors is controlled by a regulatory RNA molecule.","volume":"12","author":"Novick","year":"1993","journal-title":"EMBO J."},{"key":"B63","doi-asserted-by":"publisher","first-page":"1142","DOI":"10.1111\/j.1365-2958.2010.07105.x","article-title":"Redox sensing by a Rex-family repressor is involved in the regulation of anaerobic gene expression in Staphylococcus aureus.","volume":"76","author":"Pagels","year":"2010","journal-title":"Mol. Microbiol."},{"key":"B64","doi-asserted-by":"publisher","first-page":"10532","DOI":"10.1073\/pnas.1408797111","article-title":"Identification of a two-component fatty acid kinase responsible for host fatty acid incorporation by Staphylococcus aureus.","volume":"111","author":"Parsons","year":"2014","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B65","doi-asserted-by":"publisher","first-page":"5729","DOI":"10.1128\/aac.01038-13","article-title":"Staphylococcus aureus fatty acid auxotrophs do not proliferate in mice.","volume":"57","author":"Parsons","year":"2013","journal-title":"Antimicrob. Agents Chemother."},{"key":"B66","doi-asserted-by":"publisher","first-page":"544","DOI":"10.1016\/j.mib.2011.07.029","article-title":"Is bacterial fatty acid synthesis a valid target for antibacterial drug discovery?","volume":"14","author":"Parsons","year":"2011","journal-title":"Curr. Opin. Microbiol."},{"key":"B67","doi-asserted-by":"publisher","first-page":"314","DOI":"10.1021\/bi0603266","article-title":"Repression of Staphylococcus aureus SrrAB using inducible antisense srrA alters growth and virulence factor transcript levels.","volume":"46","author":"Pragman","year":"2007","journal-title":"Biochemistry"},{"key":"B68","doi-asserted-by":"publisher","first-page":"434","DOI":"10.1128\/9781555816513.ch35","article-title":"Respiration and small-colony variants of Staphylococcus aureus","author":"Proctor","year":"2006","journal-title":"Gram-Positive Pathogens"},{"key":"B69","doi-asserted-by":"publisher","first-page":"549","DOI":"10.1128\/9781555816513.ch35","article-title":"Respiration and small colony variants of Staphylococcus aureus.","volume":"7","author":"Proctor","year":"2019","journal-title":"Microbiol. Spectr."},{"key":"B70","doi-asserted-by":"publisher","first-page":"1","DOI":"10.21775\/9781910190494.01","author":"Proctor","year":"2016","journal-title":"Staphylococcus: Genetics and Physiology"},{"key":"B71","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2014.00099","article-title":"Staphylococcus aureus small colony variants (SCVs): a road map for the metabolic pathways involved in persistent infections.","volume":"4","author":"Proctor","year":"2014","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"B72","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1093\/clinids\/20.1.95","article-title":"Persistent and relapsing infections associated with small-colony variants of Staphylococcus aureus.","volume":"20","author":"Proctor","year":"1995","journal-title":"Clin. Infect. Dis."},{"key":"B73","doi-asserted-by":"publisher","first-page":"295","DOI":"10.1038\/nrmicro1384","article-title":"Small colony variants: a pathogenic form of bacteria that facilitates persistent and recurrent infections.","volume":"4","author":"Proctor","year":"2006","journal-title":"Nat. Rev. Microbiol."},{"key":"B74","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1016\/j.molcel.2018.10.022","article-title":"ATP-dependent dynamic protein aggregation regulates bacterial dormancy depth critical for antibiotic tolerance.","volume":"73","author":"Pu","year":"2019","journal-title":"Mol Cell"},{"key":"B75","doi-asserted-by":"publisher","first-page":"1126","DOI":"10.1093\/jac\/dki385","article-title":"Staphylococcus aureus small colony variants are resistant to the antimicrobial peptide lactoferricin B.","volume":"56","author":"Samuelsen","year":"2005","journal-title":"J. Antimicrob. Chemother."},{"key":"B76","doi-asserted-by":"publisher","first-page":"75","DOI":"10.1099\/00222615-24-1-75","article-title":"Environmental factors affecting toxic shock syndrome toxin-1 (TSST-1) synthesis.","volume":"24","author":"Sarafian","year":"1987","journal-title":"J. Med. Microbiol."},{"key":"B77","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2018.01863","article-title":"The energy-coupling factor transporter module EcfAA\u2019T, a novel candidate for the genetic basis of fatty acid-auxotrophic small-colony variants of Staphylococcus aureus.","volume":"9","author":"Schleimer","year":"2018","journal-title":"Front. Microbiol."},{"key":"B78","doi-asserted-by":"publisher","first-page":"631","DOI":"10.1016\/j.ijmm.2018.02.003","article-title":"Staphylococcus aureus in the airways of cystic fibrosis patients \u2013 a retrospective long-term study.","volume":"308","author":"Schwerdt","year":"2018","journal-title":"Int. J. Med. Microbiol."},{"key":"B79","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1093\/jac\/dkl450","article-title":"Small-colony variants: a novel mechanism for triclosan resistance in methicillin-resistant Staphylococcus aureus.","volume":"59","author":"Seaman","year":"2007","journal-title":"J. Antimicrob. Chemother."},{"key":"B80","doi-asserted-by":"publisher","first-page":"7765","DOI":"10.1128\/jb.00774-06","article-title":"Reporter metabolite analysis of transcriptional profiles of a Staphylococcus aureus strain with normal phenotype and its isogenic hemB mutant displaying the small-colony-variant phenotype.","volume":"188","author":"Seggewiss","year":"2006","journal-title":"J. Bacteriol."},{"key":"B81","doi-asserted-by":"publisher","first-page":"7397","DOI":"10.1128\/jb.187.21.7397-7406.2005","article-title":"sigmaB activity in a Staphylococcus aureus hemB mutant.","volume":"187","author":"Senn","year":"2005","journal-title":"J. Bacteriol."},{"key":"B82","doi-asserted-by":"publisher","DOI":"10.1128\/JB.00409-19","article-title":"Timing is everything: impact of naturally occurring Staphylococcus aureus AgrC cytoplasmic domain adaptive mutations on autoinduction.","volume":"201","author":"Sloan","year":"2019","journal-title":"J. Bacteriol."},{"key":"B83","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-017-14110-8","article-title":"Cytolytic toxin production by Staphylococcus aureus is dependent upon the activity of the protoheme IX farnesyltransferase.","volume":"7","author":"Stevens","year":"2017","journal-title":"Sci. Rep."},{"key":"B84","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2018.00018","article-title":"Mutation of Agr is associated with the adaptation of Staphylococcus aureus to the host during chronic osteomyelitis.","volume":"8","author":"Suligoy","year":"2018","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"B85","doi-asserted-by":"publisher","first-page":"774","DOI":"10.1038\/nri.2017.103","article-title":"Regulation of immunity and inflammation by hypoxia in immunological niches.","volume":"17","author":"Taylor","year":"2017","journal-title":"Nat. Rev. Immunol."},{"key":"B86","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1004870","article-title":"Sigma factor SigB is crucial to mediate Staphylococcus aureus adaptation during chronic infections.","volume":"11","author":"Tuchscherr","year":"2015","journal-title":"PLoS Pathog."},{"key":"B87","doi-asserted-by":"publisher","DOI":"10.3390\/pathogens6030031","article-title":"Staphylococcus aureus regulator sigma B is important to develop chronic infections in hematogenous murine osteomyelitis model.","volume":"6","author":"Tuchscherr","year":"2017","journal-title":"Pathogens"},{"key":"B88","doi-asserted-by":"publisher","first-page":"1031","DOI":"10.1086\/656047","article-title":"Staphylococcus aureus small-colony variants are adapted phenotypes for intracellular persistence.","volume":"202","author":"Tuchscherr","year":"","journal-title":"J. Infect. Dis."},{"key":"B89","doi-asserted-by":"publisher","first-page":"438","DOI":"10.1093\/jac\/dkv371","article-title":"Staphylococcus aureus develops increased resistance to antibiotics by forming dynamic small colony variants during chronic osteomyelitis.","volume":"71","author":"Tuchscherr","year":"2016","journal-title":"J. Antimicrob. Chemother."},{"key":"B90","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1007\/s00294-015-0503-0","article-title":"Staphylococcus aureus dynamically adapts global regulators and virulence factor expression in the course from acute to chronic infection.","volume":"62","author":"Tuchscherr","year":"2015","journal-title":"Curr. Genet."},{"key":"B91","doi-asserted-by":"publisher","first-page":"1823","DOI":"10.2217\/fmb.10.147","article-title":"Staphylococcus aureus adaptation to the host and persistence: role of loss of capsular polysaccharide expression.","volume":"5","author":"Tuchscherr","year":"","journal-title":"Future Microbiol."},{"key":"B92","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1002\/emmm.201000115","article-title":"Staphylococcus aureus phenotype switching: an effective bacterial strategy to escape host immune response and establish a chronic infection.","volume":"3","author":"Tuchscherr","year":"2011","journal-title":"EMBO Mol. Med."},{"key":"B93","doi-asserted-by":"publisher","DOI":"10.3390\/toxins11030135","article-title":"Clinical S. aureus isolates vary in their virulence to promote adaptation to the host.","volume":"11","author":"Tuchscherr","year":"2019","journal-title":"Toxins"},{"key":"B94","doi-asserted-by":"publisher","first-page":"5738","DOI":"10.1128\/IAI.00874-08","article-title":"Antibodies to capsular polysaccharide and clumping factor a prevent mastitis and the emergence of unencapsulated and small-colony variants of Staphylococcus aureus in mice.","volume":"76","author":"Tuchscherr","year":"2008","journal-title":"Infect. Immun."},{"key":"B95","doi-asserted-by":"publisher","first-page":"739","DOI":"10.1093\/infdis\/173.3.739","article-title":"Staphylococcus aureus small colony variants are induced by the endothelial cell intracellular milieu.","volume":"173","author":"Vesga","year":"1996","journal-title":"J. Infect. Dis."},{"key":"B96","doi-asserted-by":"publisher","first-page":"687","DOI":"10.1128\/jb.188.2.687-693.2006","article-title":"Phenotype microarray profiling of Staphylococcus aureus menD and hemB mutants with the small-colony-variant phenotype.","volume":"188","author":"von Eiff","year":"2006","journal-title":"J. Bacteriol."},{"key":"B97","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-018-06527-0","article-title":"Prolonged bacterial lag time results in small colony variants that represent a sub-population of persisters.","volume":"9","author":"Vulin","year":"2018","journal-title":"Nat. Commun."},{"key":"B98","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2015.01437","article-title":"Transposon mutagenesis identifies novel genes associated with Staphylococcus aureus persister formation.","volume":"6","author":"Wang","year":"2015","journal-title":"Front. Microbiol."},{"key":"B99","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-018-29123-0","article-title":"Inactivation of TCA cycle enhances Staphylococcus aureus persister cell formation in stationary phase.","volume":"8","author":"Wang","year":"2018","journal-title":"Sci. Rep."},{"key":"B100","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0011576","article-title":"Activation of hypoxia inducible factor 1 is a general phenomenon in infections with human pathogens.","volume":"5","author":"Werth","year":"2010","journal-title":"PLoS One"},{"key":"B101","doi-asserted-by":"publisher","first-page":"1027","DOI":"10.1016\/S2213-2600(19)30365-0","article-title":"Prevalence and clinical associations of Staphylococcus aureus small-colony variant respiratory infection in children with cystic fibrosis (SCVSA): a multicentre, observational study.","volume":"7","author":"Wolter","year":"2019","journal-title":"Lancet Respir. Med."},{"key":"B102","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1038\/s41564-019-0597-0","article-title":"Staphylococcus aureus small colony variants impair host immunity by activating host cell glycolysis and inducing necroptosis.","volume":"5","author":"Wong Fok Lung","year":"2020","journal-title":"Nat. Microbiol."},{"key":"B103","doi-asserted-by":"publisher","first-page":"7116","DOI":"10.1128\/AEM.02636-13","article-title":"Bacterial persister cell formation and dormancy.","volume":"79","author":"Wood","year":"2013","journal-title":"Appl. Environ. Microbiol."},{"key":"B104","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2180-3-16","article-title":"Effects of disruption of heat shock genes on susceptibility of Escherichia coli to fluoroquinolones.","volume":"3","author":"Yamaguchi","year":"2003","journal-title":"BMC Microbiol."},{"key":"B105","doi-asserted-by":"publisher","DOI":"10.1128\/mBio.00415-18","article-title":"Novel insights into Staphylococcus aureus deep bone infections: the involvement of osteocytes.","volume":"9","author":"Yang","year":"2018","journal-title":"MBio"},{"key":"B106","doi-asserted-by":"publisher","first-page":"1113","DOI":"10.1128\/jb.183.4.1113-1123.2001","article-title":"Identification of a novel two-component regulatory system that acts in global regulation of virulence factors of Staphylococcus aureus.","volume":"183","author":"Yarwood","year":"2001","journal-title":"J. Bacteriol."},{"key":"B107","doi-asserted-by":"publisher","first-page":"615","DOI":"10.1093\/jac\/dkx433","article-title":"Bicarbonate induces high-level resistance to the human antimicrobial peptide LL-37 in Staphylococcus aureus small colony variants.","volume":"73","author":"Zhang","year":"2018","journal-title":"J. Antimicrob. Chemother."}],"container-title":["Frontiers in Microbiology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/article\/10.3389\/fmicb.2020.01028\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,10,24]],"date-time":"2022-10-24T12:24:22Z","timestamp":1666614262000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/article\/10.3389\/fmicb.2020.01028\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,5,21]]},"references-count":107,"alternative-id":["10.3389\/fmicb.2020.01028"],"URL":"https:\/\/doi.org\/10.3389\/fmicb.2020.01028","relation":{},"ISSN":["1664-302X"],"issn-type":[{"value":"1664-302X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,5,21]]},"article-number":"1028"}}