{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,2]],"date-time":"2026-06-02T07:09:16Z","timestamp":1780384156000,"version":"3.54.1"},"reference-count":66,"publisher":"Public Library of Science (PLoS)","issue":"6","license":[{"start":{"date-parts":[[2021,6,1]],"date-time":"2021-06-01T00:00:00Z","timestamp":1622505600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000060","name":"National Institute of Allergy and Infectious Diseases","doi-asserted-by":"publisher","award":["R01 AI141656"],"award-info":[{"award-number":["R01 AI141656"]}],"id":[{"id":"10.13039\/100000060","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS ONE"],"abstract":"<jats:p><jats:italic>Borrelia burgdorferi (Bb)<\/jats:italic>, the etiological agent of Lyme disease, produces a series of simple glycolipids where diacylglycerol and cholesterol serve as the precursor. The cholesterol-based glycolipids, cholesteryl 6-<jats:italic>O<\/jats:italic>-acyl-\u03b2-D-galactopyranoside (ACGal) and cholesteryl-\u03b2-D-galactopyranoside (CGal) are immunogenic and proposed to contribute to the pathogenesis of Lyme disease. Detailed studies of CGal and ACGal in <jats:italic>Bb<\/jats:italic> have been hampered by a lack of knowledge of their underlying biosynthetic processes. The genome of <jats:italic>Bb<\/jats:italic> encodes four putative glycosyltransferases, and only one of these, BB0572, was predicted to be an inverting family 2 glycosyltransferase (GT2 enzyme) capable of using UDP-galactose as a substrate and forming a \u03b2-glycosidic bond. Comparison of the 42 kDa BB0572 amino acid sequence from <jats:italic>Bb<\/jats:italic> with other <jats:italic>Borrelia<\/jats:italic> spp demonstrates that this protein is highly conserved. To establish BB0572 as the galactosyltransferase capable of cholesterol glycolipid formation in <jats:italic>Bb<\/jats:italic>, the protein was produced as a recombinant product in <jats:italic>Escherichia coli<\/jats:italic> and tested in a cell-free assay with <jats:sup>14<\/jats:sup>C-cholesterol and UDP-galactose as the substrates. This experiment resulted in a radiolabeled lipid that migrated with the cholesterol glycolipid standard of CGal when evaluated by thin layer chromatography. Additionally, mutation in the predicted active site of BB0572 resulted in a recombinant protein that was unable to catalyze the formation of the cholesterol glycolipid. These data characterize BB0572 as a putative cholesterol galactosyltransferase. This provides the first step in understanding how <jats:italic>Bb<\/jats:italic> cholesterol glycolipids are formed and will allow investigations into their involvement in pathogen transmission and disease development.<\/jats:p>","DOI":"10.1371\/journal.pone.0252214","type":"journal-article","created":{"date-parts":[[2021,6,1]],"date-time":"2021-06-01T20:04:18Z","timestamp":1622577858000},"page":"e0252214","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":8,"title":["Identification and functional analysis of a galactosyltransferase capable of cholesterol glycolipid formation in the Lyme disease spirochete Borrelia burgdorferi"],"prefix":"10.1371","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4779-5264","authenticated-orcid":true,"given":"Petronella R.","family":"Hove","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Forgivemore","family":"Magunda","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria Angela","family":"de Mello Marques","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"M. 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Acta"},{"issue":"38","key":"pone.0252214.ref016","doi-asserted-by":"crossref","first-page":"27765","DOI":"10.1074\/jbc.M603345200","article-title":"Cloning of a Cholesterol-\u03b1-glucosyltransferase from Helicobacter pylori","volume":"281","author":"A-H Lebrun","year":"2006","journal-title":"Journal of Biological Chemistry"},{"key":"pone.0252214.ref017","doi-asserted-by":"crossref","first-page":"D490","DOI":"10.1093\/nar\/gkt1178","article-title":"The carbohydrate-active enzymes database (CAZy) in 2013","volume":"42","author":"V Lombard","year":"2014","journal-title":"Nucleic Acids Research"},{"issue":"1","key":"pone.0252214.ref018","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1128\/IAI.01061-13","article-title":"Presence of Arp Specifically Contributes to Joint Tissue Edema Associated with Early-Onset Lyme Arthritis","volume":"82","author":"PR Hove","year":"2014","journal-title":"Infection and Immunity"},{"issue":"1","key":"pone.0252214.ref019","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1186\/s12866-016-0806-4","article-title":"Investigating the potential role of non-vls genes on linear plasmid 28\u20131 in virulence and persistence by Borrelia burgdorferi","volume":"16","author":"PRH Magunda","year":"2016","journal-title":"BMC Microbiology"},{"issue":"4","key":"pone.0252214.ref020","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1006\/meth.2001.1262","article-title":"Analysis of relative gene expression data using real-time quantitative PCR and the 2(-Delta Delta C(T)) Method","volume":"25","author":"KJ Livak","year":"2001","journal-title":"Methods (San Diego, Calif)."},{"issue":"24","key":"pone.0252214.ref021","doi-asserted-by":"crossref","first-page":"22572","DOI":"10.1074\/jbc.M500549200","article-title":"Dissecting the Structural Determinants of the Stability of Cholesterol Oxidase Containing Covalently Bound Flavin","volume":"280","author":"L Caldinelli","year":"2005","journal-title":"Journal of Biological Chemistry"},{"key":"pone.0252214.ref022","doi-asserted-by":"crossref","first-page":"W13","DOI":"10.1093\/nar\/gkr245","article-title":"T-Coffee: a web server for the multiple sequence alignment of protein and RNA sequences using structural information and homology extension","volume":"39","author":"P Di Tommaso","year":"2011","journal-title":"Nucleic Acids Res"},{"issue":"W1","key":"pone.0252214.ref023","doi-asserted-by":"crossref","first-page":"W3","DOI":"10.1093\/nar\/gkv310","article-title":"TCS: a web server for multiple sequence alignment evaluation and phylogenetic reconstruction","volume":"43","author":"JM Chang","year":"2015","journal-title":"Nucleic Acids Res"},{"issue":"6","key":"pone.0252214.ref024","doi-asserted-by":"crossref","first-page":"1625","DOI":"10.1093\/molbev\/msu117","article-title":"TCS: a new multiple sequence alignment reliability measure to estimate alignment accuracy and improve phylogenetic tree reconstruction","volume":"31","author":"JM Chang","year":"2014","journal-title":"Molecular biology and evolution"},{"issue":"6","key":"pone.0252214.ref025","doi-asserted-by":"crossref","first-page":"1547","DOI":"10.1093\/molbev\/msy096","article-title":"MEGA X: Molecular Evolutionary Genetics Analysis across Computing Platforms","volume":"35","author":"S Kumar","year":"2018","journal-title":"Molecular biology and evolution"},{"key":"pone.0252214.ref026","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1016\/B978-1-4832-2734-4.50017-6","volume-title":"Evolving Genes and Proteins","author":"E Zuckerkandl","year":"1965"},{"issue":"13","key":"pone.0252214.ref027","doi-asserted-by":"crossref","first-page":"1608","DOI":"10.1093\/bioinformatics\/btq249","article-title":"PSORTb 3.0: improved protein subcellular localization prediction with refined localization subcategories and predictive capabilities for all prokaryotes","volume":"26","author":"NY Yu","year":"2010","journal-title":"Bioinformatics (Oxford, England)"},{"issue":"3","key":"pone.0252214.ref028","doi-asserted-by":"crossref","first-page":"200","DOI":"10.1007\/s10930-019-09838-3","article-title":"A Brief History of Protein Sorting Prediction","volume":"38","author":"H Nielsen","year":"2019","journal-title":"The Protein Journal"},{"issue":"2","key":"pone.0252214.ref029","doi-asserted-by":"crossref","first-page":"29R","DOI":"10.1093\/glycob\/cwj016","article-title":"Structures and mechanisms of glycosyltransferases","volume":"16","author":"C Breton","year":"2005","journal-title":"Glycobiology"},{"key":"pone.0252214.ref030","doi-asserted-by":"crossref","first-page":"D233","DOI":"10.1093\/nar\/gkn663","article-title":"The Carbohydrate-Active EnZymes database (CAZy): an expert resource for Glycogenomics","volume":"37","author":"BL Cantarel","year":"2009","journal-title":"Nucleic Acids Research"},{"issue":"D1","key":"pone.0252214.ref031","doi-asserted-by":"crossref","first-page":"D200","DOI":"10.1093\/nar\/gkw1129","article-title":"CDD\/SPARCLE: functional classification of proteins via subfamily domain architectures","volume":"45","author":"A Marchler-Bauer","year":"2017","journal-title":"Nucleic acids research"},{"key":"pone.0252214.ref032","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1016\/j.mcp.2016.08.003","article-title":"Emerging borreliae\u2013Expanding beyond Lyme borreliosis","volume":"31","author":"SJ Cutler","year":"2017","journal-title":"Molecular and Cellular Probes"},{"issue":"2","key":"pone.0252214.ref033","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1016\/j.idc.2015.02.005","article-title":"Laboratory Diagnosis of Lyme Disease: Advances and Challenges","volume":"29","author":"AR Marques","year":"2015","journal-title":"Infectious Disease Clinics of North America"},{"issue":"5","key":"pone.0252214.ref034","doi-asserted-by":"crossref","first-page":"556","DOI":"10.1016\/S1473-3099(15)00464-8","article-title":"Identification of a novel pathogenic Borrelia species causing Lyme borreliosis with unusually high spirochaetaemia: a descriptive study","volume":"16","author":"BS Pritt","year":"2016","journal-title":"The Lancet Infectious Diseases"},{"issue":"3","key":"pone.0252214.ref035","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1016\/0168-1702(85)90051-6","article-title":"On the predictive recognition of signal peptide sequences","volume":"3","author":"DJ McGeoch","year":"1985","journal-title":"Virus Research"},{"issue":"2","key":"pone.0252214.ref036","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1002\/prot.340110203","article-title":"Expert system for predicting protein localization sites in gram-negative bacteria","volume":"11","author":"K Nakai","year":"1991","journal-title":"Proteins: Structure, Function, and Bioinformatics"},{"issue":"23","key":"pone.0252214.ref037","doi-asserted-by":"crossref","first-page":"3698","DOI":"10.1128\/JB.00625-15","article-title":"The Essential Role of Cholesterol Metabolism in the Intracellular Survival of Mycobacterium leprae Is Not Coupled to Central Carbon Metabolism and Energy Production","volume":"197","author":"MA Marques","year":"2015","journal-title":"J Bacteriol"},{"issue":"1","key":"pone.0252214.ref038","doi-asserted-by":"crossref","first-page":"521","DOI":"10.1146\/annurev.biochem.76.061005.092322","article-title":"Glycosyltransferases: Structures, Functions, and Mechanisms","volume":"77","author":"LL Lairson","year":"2008","journal-title":"Annual Review of Biochemistry"},{"issue":"10","key":"pone.0252214.ref039","doi-asserted-by":"crossref","first-page":"e47532","DOI":"10.1371\/journal.pone.0047532","article-title":"Analysis of an ordered, comprehensive STM mutant library in infectious Borrelia burgdorferi: insights into the genes required for mouse infectivity","volume":"7","author":"T Lin","year":"2012","journal-title":"PloS one"},{"key":"pone.0252214.ref040","first-page":"65","volume-title":"Essentials of Glycobiology","author":"JM Rini","year":"2015"},{"issue":"2","key":"pone.0252214.ref041","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1016\/S0022-2836(03)00307-3","article-title":"An evolving hierarchical family classification for glycosyltransferases","volume":"328","author":"PM Coutinho","year":"2003","journal-title":"J Mol Biol"},{"issue":"1","key":"pone.0252214.ref042","doi-asserted-by":"crossref","first-page":"570","DOI":"10.1128\/JB.124.1.570-572.1975","article-title":"Cholesterol incorporation into bacterial membranes","volume":"124","author":"S Razin","year":"1975","journal-title":"Journal of Bacteriology"},{"issue":"5","key":"pone.0252214.ref043","doi-asserted-by":"crossref","first-page":"1289","DOI":"10.1006\/bbrc.2001.2023","article-title":"Sterols and Acylated Proteins in Mycoplasmas","volume":"292","author":"S Rottem","year":"2002","journal-title":"Biochemical and Biophysical Research Communications"},{"issue":"18","key":"pone.0252214.ref044","doi-asserted-by":"crossref","first-page":"5327","DOI":"10.1128\/JB.177.18.5327-5333.1995","article-title":"Unique cholesteryl glucosides in Helicobacter pylori: composition and structural analysis","volume":"177","author":"Y Hirai","year":"1995","journal-title":"Journal of Bacteriology"},{"issue":"3","key":"pone.0252214.ref045","doi-asserted-by":"crossref","first-page":"986","DOI":"10.1128\/jb.108.3.986-991.1971","article-title":"Biosynthesis of cholesteryl glucoside by Mycoplasma gallinarum","volume":"108","author":"PF Smith","year":"1971","journal-title":"Journal of bacteriology"},{"issue":"8","key":"pone.0252214.ref046","doi-asserted-by":"crossref","first-page":"713","DOI":"10.1016\/0300-9084(93)90102-X","article-title":"UDP-glucose sterol \u00df-d-glucosyltransferase, a plasma membrane-bound enzyme of plants: Enzymatic properties and lipid dependence","volume":"75","author":"P Ullmann","year":"1993","journal-title":"Biochimie"},{"issue":"9","key":"pone.0252214.ref047","doi-asserted-by":"crossref","first-page":"989","DOI":"10.1007\/BF02522694","article-title":"Sterol composition ofPhaeodactylum tricornutum as influenced by growth temperature and light spectral quality","volume":"31","author":"B V\u00e9ron","year":"1996","journal-title":"Lipids"},{"issue":"2","key":"pone.0252214.ref048","doi-asserted-by":"crossref","first-page":"322","DOI":"10.1016\/0005-2760(80)90213-1","article-title":"Cholesteryl glucoside in Candida bogoriensis","volume":"620","author":"T Kastelic-Suhadolc","year":"1980","journal-title":"Biochimica et Biophysica Acta (BBA)\u2014Lipids and Lipid Metabolism"},{"issue":"2","key":"pone.0252214.ref049","doi-asserted-by":"crossref","first-page":"308","DOI":"10.1016\/0005-2760(77)90026-1","article-title":"Cholesterol \u03b2-d-glucoside-6\u2019-O-palmitate, a metabolite of Pythium sylvaticum","volume":"486","author":"TC McMorris","year":"1977","journal-title":"Biochimica et Biophysica Acta (BBA)\u2014Lipids and Lipid Metabolism"},{"issue":"1","key":"pone.0252214.ref050","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1042\/bj1410133","article-title":"The cell-envelope glycolipids of baker\u2019s yeast","volume":"141","author":"K Ty\u00f6rinoja","year":"1974","journal-title":"Biochemical Journal"},{"issue":"1","key":"pone.0252214.ref051","doi-asserted-by":"crossref","first-page":"486","DOI":"10.1074\/jbc.272.1.486","article-title":"Heat Stress Induces a Glycosylation of Membrane Sterol in Myxoamoebae of a True Slime Mold, Physarum polycephalum","volume":"272","author":"K Murakami-Murofushi","year":"1997","journal-title":"Journal of Biological Chemistry"},{"issue":"2","key":"pone.0252214.ref052","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1016\/0005-2744(79)90156-6","article-title":"Specificity of sterol-glucosylating enzymes from Sinapis alba and Physarum polycephalum","volume":"570","author":"ZA Wojciechowski","year":"1979","journal-title":"Biochimica et Biophysica Acta (BBA)\u2014Enzymology"},{"issue":"4","key":"pone.0252214.ref053","doi-asserted-by":"crossref","first-page":"427","DOI":"10.1016\/S0022-2275(20)38823-4","article-title":"Lipids of chicken epidermis","volume":"27","author":"PW Wertz","year":"1986","journal-title":"Journal of Lipid Research"},{"issue":"4","key":"pone.0252214.ref054","doi-asserted-by":"crossref","first-page":"446","DOI":"10.1016\/S0022-2275(20)38695-8","article-title":"Glucosylsterol and acylglucosylsterol of snake epidermis: structure determination","volume":"28","author":"W Abraham","year":"1987","journal-title":"Journal of Lipid Research"},{"issue":"2","key":"pone.0252214.ref055","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1016\/0009-3084(94)90038-8","article-title":"In vivo incorporation of [2-14C]mevalonic acid into rat brain complex glycolipids: characterization of glycosylsterols in mammalian brain","volume":"69","author":"BL Hungund","year":"1994","journal-title":"Chemistry and Physics of Lipids"},{"issue":"10","key":"pone.0252214.ref056","doi-asserted-by":"crossref","first-page":"1117","DOI":"10.1016\/S0022-2275(20)37721-X","article-title":"Characterization of human liver 3-O-beta-D-glucopyranuronosyl-cholesterol by mass spectrometry and nuclear magnetic resonance spectroscopy","volume":"25","author":"IA Muhiudeen","year":"1984","journal-title":"Journal of Lipid Research"},{"issue":"1","key":"pone.0252214.ref057","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1128\/iai.20.1.215-220.1978","article-title":"Lipid metabolism of Borrelia hermsi","volume":"20","author":"BP Livermore","year":"1978","journal-title":"Infection and Immunity"},{"issue":"20","key":"pone.0252214.ref058","doi-asserted-by":"crossref","first-page":"13326","DOI":"10.1074\/jbc.M809575200","article-title":"Acylated Cholesteryl Galactosides Are Specific Antigens of Borrelia Causing Lyme Disease and Frequently Induce Antibodies in Late Stages of Disease","volume":"284","author":"G St\u00fcbs","year":"2009","journal-title":"Journal of Biological Chemistry"},{"issue":"11","key":"pone.0252214.ref059","doi-asserted-by":"crossref","first-page":"3536","DOI":"10.1002\/chem.200902205","article-title":"Chemoenzymatic Synthesis of a Glycolipid Library and Elucidation of the Antigenic Epitope for Construction of a Vaccine Against Lyme Disease","volume":"16","author":"G St\u00fcbs","year":"2010","journal-title":"Chemistry\u2013A European Journal"},{"key":"pone.0252214.ref060","doi-asserted-by":"crossref","first-page":"1030","DOI":"10.1038\/nm1480","article-title":"Cholesterol glucosylation promotes immune evasion by Helicobacter pylori","volume":"12","author":"C Wunder","year":"2006","journal-title":"Nature Medicine"},{"issue":"6","key":"pone.0252214.ref061","doi-asserted-by":"crossref","first-page":"2897","DOI":"10.1128\/AAC.00016-11","article-title":"Cholesterol enhances Helicobacter pylori resistance to antibiotics and LL-37","volume":"55","author":"DJ McGee","year":"2011","journal-title":"Antimicrobial agents and chemotherapy"},{"issue":"4","key":"pone.0252214.ref062","doi-asserted-by":"crossref","first-page":"331","DOI":"10.1016\/j.chom.2010.09.001","article-title":"Cholesterol lipids of Borrelia burgdorferi form lipid rafts and are required for the bactericidal mechanism of a complement-independent antibody","volume":"8","author":"TJ LaRocca","year":"2010","journal-title":"Cell host & microbe"},{"issue":"2","key":"pone.0252214.ref063","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.00899-14","article-title":"Selective Association of Outer Surface Lipoproteins with the Lipid Rafts of Borrelia burgdorferi","volume":"5","author":"A 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