{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,30]],"date-time":"2026-10-30T02:02:58Z","timestamp":1793325778667,"version":"4.5.0"},"reference-count":73,"publisher":"Oxford University Press (OUP)","issue":"12","license":[{"start":{"date-parts":[[2023,10,26]],"date-time":"2023-10-26T00:00:00Z","timestamp":1698278400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01AI148844"],"award-info":[{"award-number":["R01AI148844"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["AI078958"],"award-info":[{"award-number":["AI078958"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["DE023080"],"award-info":[{"award-number":["DE023080"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R21AI163663"],"award-info":[{"award-number":["R21AI163663"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001807","name":"Funda\u00e7\u00e3o de Amparo \u00e0 Pesquisa do Estado de S\u00e3o Paulo","doi-asserted-by":"publisher","award":["2020\/02678-8"],"award-info":[{"award-number":["2020\/02678-8"]}],"id":[{"id":"10.13039\/501100001807","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Chemical Biology Interface NIH training","award":["T32GM138826"],"award-info":[{"award-number":["T32GM138826"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2023,12,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Spirochetes cause Lyme disease, leptospirosis, syphilis, and several other human illnesses. Unlike other bacteria, spirochete flagella are enclosed within the periplasmic space where the filaments distort and push the cell body by the action of the flagellar motors. We previously demonstrated that the oral pathogen Treponema denticola (Td) and Lyme disease pathogen Borreliella burgdorferi (Bb) form covalent lysinoalanine (Lal) cross-links between conserved cysteine and lysine residues of the FlgE protein that composes the flagellar hook. In Td, Lal is unnecessary for hook assembly but is required for motility, presumably due to the stabilizing effect of the cross-link. Herein, we extend these findings to other, representative spirochete species across the phylum. We confirm the presence of Lal cross-linked peptides in recombinant and in vivo-derived samples from Treponema spp., Borreliella spp., Brachyspira spp., and Leptospira spp. As was observed with Td, a mutant strain of Bb unable to form the cross-link has greatly impaired motility. FlgE from Leptospira spp. does not conserve the Lal-forming cysteine residue which is instead substituted by serine. Nevertheless, Leptospira interrogans FlgE also forms Lal, with several different Lal isoforms being detected between Ser-179 and Lys-145, Lys-148, and Lys-166, thereby highlighting species or order-specific differences within the phylum. Our data reveal that the Lal cross-link is a conserved and necessary posttranslational modification across the spirochete phylum and may thus represent an effective target for the development of spirochete-specific antimicrobials.<\/jats:p>","DOI":"10.1093\/pnasnexus\/pgad349","type":"journal-article","created":{"date-parts":[[2023,10,26]],"date-time":"2023-10-26T13:42:20Z","timestamp":1698327740000},"source":"Crossref","is-referenced-by-count":8,"title":["Lysinoalanine cross-linking is a conserved post-translational modification in the spirochete flagellar hook"],"prefix":"10.1093","volume":"2","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3569-4749","authenticated-orcid":false,"given":"Michael J","family":"Lynch","sequence":"first","affiliation":[{"name":"Department of Chemistry and Chemical Biology, Cornell University , Ithaca, NY 14853 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maithili","family":"Deshpande","sequence":"additional","affiliation":[{"name":"Department of Chemistry and Chemical Biology, Cornell University , Ithaca, NY 14853 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kurni","family":"Kurniyati","sequence":"additional","affiliation":[{"name":"Philips Institute for Oral Health Research, Virginia Commonwealth University School of Dentistry , Richmond, VA 23298 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kai","family":"Zhang","sequence":"additional","affiliation":[{"name":"Philips Institute for Oral Health Research, Virginia Commonwealth University School of Dentistry , Richmond, VA 23298 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Milinda","family":"James","sequence":"additional","affiliation":[{"name":"Department of Microbiology, Immunology, and Cell Biology, Robert C. Byrd Health Sciences Center, West Virginia University , Morgantown, WV 26505 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael","family":"Miller","sequence":"additional","affiliation":[{"name":"Department of Biochemistry, Robert C. Byrd Health Sciences Center, West Virginia University , Morgantown, WV 26505 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8206-1007","authenticated-orcid":false,"given":"Sheng","family":"Zhang","sequence":"additional","affiliation":[{"name":"Proteomics and Metabolomics Facility, Institute of Biotechnology, Cornell University , Ithaca, NY 14853 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0305-7160","authenticated-orcid":false,"given":"Felipe J","family":"Passalia","sequence":"additional","affiliation":[{"name":"Department of Epidemiology of Microbial Diseases, Yale School of Public Health , New Haven, CT 06510 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5239-8511","authenticated-orcid":false,"suffix":"Jr.","given":"Elsio A","family":"Wunder","sequence":"additional","affiliation":[{"name":"Department of Epidemiology of Microbial Diseases, Yale School of Public Health , New Haven, CT 06510 , USA"},{"name":"Department of Pathobiology and Veterinary Science, University of Connecticut , Storrs, CT 06269 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-2734-7467","authenticated-orcid":false,"given":"Nyles W","family":"Charon","sequence":"additional","affiliation":[{"name":"Department of Microbiology, Immunology, and Cell Biology, Robert C. Byrd Health Sciences Center, West Virginia University , Morgantown, WV 26505 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1872-4870","authenticated-orcid":false,"given":"Chunhao","family":"Li","sequence":"additional","affiliation":[{"name":"Philips Institute for Oral Health Research, Virginia Commonwealth University School of Dentistry , Richmond, VA 23298 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8234-9991","authenticated-orcid":false,"given":"Brian R","family":"Crane","sequence":"additional","affiliation":[{"name":"Department of Chemistry and Chemical Biology, Cornell University , Ithaca, NY 14853 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2023,10,26]]},"reference":[{"key":"2023122115210803900_pgad349-B1","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1146\/annurev-micro-092611-150145","article-title":"The unique paradigm of spirochete motility and chemotaxis","volume":"66","author":"Charon","year":"2012","journal-title":"Annu Rev Microbiol"},{"key":"2023122115210803900_pgad349-B2","doi-asserted-by":"crossref","first-page":"1413","DOI":"10.2217\/fmb.10.102","article-title":"Leptospira as an emerging pathogen: a review of its biology, pathogenesis and host immune responses","volume":"5","author":"Evangelista","year":"2010","journal-title":"Future Microbiol"},{"key":"2023122115210803900_pgad349-B3","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1002\/jobm.201100335","article-title":"Flagella and bacterial pathogenicity","volume":"53","author":"Duan","year":"2013","journal-title":"J Basic Microbiol"},{"key":"2023122115210803900_pgad349-B4","doi-asserted-by":"crossref","first-page":"322","DOI":"10.3389\/fcimb.2018.00322","article-title":"Editorial: pathogenesis of Leptospira","volume":"8","author":"Wunder","year":"2018","journal-title":"Fronti Cell Infect Microb"},{"key":"2023122115210803900_pgad349-B5","doi-asserted-by":"crossref","first-page":"744","DOI":"10.1038\/nrmicro.2016.141","article-title":"Treponema pallidum, the syphilis spirochete: making a living as a stealth pathogen","volume":"14","author":"Radolf","year":"2016","journal-title":"Nat Rev Microbiol"},{"key":"2023122115210803900_pgad349-B6","doi-asserted-by":"crossref","first-page":"104193","DOI":"10.1016\/j.micpath.2020.104193","article-title":"Oral spirochetes: pathogenic mechanisms in periodontal disease","volume":"144","author":"Yousefi","year":"2020","journal-title":"Microb Pathog"},{"key":"2023122115210803900_pgad349-B7","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1016\/j.semcdb.2015.10.015","article-title":"Flagellar motility of the pathogenic spirochetes","volume":"46","author":"Wolgemuth","year":"2015","journal-title":"Semin Cell Dev Biol"},{"key":"2023122115210803900_pgad349-B8","doi-asserted-by":"crossref","first-page":"2115","DOI":"10.1099\/ijsem.0.004027","article-title":"Treponema phagedenis (ex Noguchi 1912) Brumpt 1922 sp. nov., nom. rev., isolated from bovine digital dermatitis","volume":"70","author":"Kuhnert","year":"2020","journal-title":"Int J Syst Evol Microbiol"},{"key":"2023122115210803900_pgad349-B9","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1177\/0300985816653795","article-title":"Swine dysentery","volume":"54","author":"Burrough","year":"2017","journal-title":"Vet Pathol"},{"key":"2023122115210803900_pgad349-B10","doi-asserted-by":"crossref","first-page":"4154","DOI":"10.1128\/iai.64.10.4154-4162.1996","article-title":"Dual flaA1 flaB1 mutant of Serpulina hyodysenteriae expressing periplasmic flagella is severely attenuated in a murine model of swine dysentery","volume":"64","author":"Rosey","year":"1996","journal-title":"Infect Immun"},{"key":"2023122115210803900_pgad349-B11","doi-asserted-by":"crossref","first-page":"baaa062","DOI":"10.1093\/database\/baaa062","article-title":"NCBI Taxonomy: a comprehensive update on curation, resources and tools","volume":"2020","author":"Schoch","year":"2020","journal-title":"Database (Oxford)"},{"key":"2023122115210803900_pgad349-B12","first-page":"217","article-title":"A phylogenomic and molecular signature based approach for characterization of the phylum spirochaetes and its major clades: proposal for a taxonomic revision of the phylum","volume":"4","author":"Gupta","year":"2013","journal-title":"Front Microbiol"},{"key":"2023122115210803900_pgad349-B13","doi-asserted-by":"crossref","first-page":"4442","DOI":"10.1093\/nar\/gkz246","article-title":"AnnoTree: visualization and exploration of a functionally annotated microbial tree of life","volume":"47","author":"Mendler","year":"2019","journal-title":"Nucleic Acids Res"},{"key":"2023122115210803900_pgad349-B14","doi-asserted-by":"crossref","first-page":"1188","DOI":"10.1101\/gr.849004","article-title":"WebLogo: a sequence logo generator","volume":"14","author":"Crooks","year":"2004","journal-title":"Genome Res"},{"key":"2023122115210803900_pgad349-B15","doi-asserted-by":"crossref","first-page":"1049","DOI":"10.1007\/s10482-014-0164-x","article-title":"A phylogenomic and molecular marker based proposal for the division of the genus Borrelia into two genera: the emended genus Borrelia containing only the members of the relapsing fever Borrelia, and the genus Borreliella gen. nov. containing the members of the Lyme disease Borrelia (Borrelia burgdorferi sensu lato complex)","volume":"105","author":"Adeolu","year":"2014","journal-title":"Antonie Van Leeuwenhoek"},{"key":"2023122115210803900_pgad349-B16","doi-asserted-by":"crossref","first-page":"e0221397","DOI":"10.1371\/journal.pone.0221397","article-title":"Distinction between Borrelia and Borreliella is more robustly supported by molecular and phenotypic characteristics than all other neighbouring prokaryotic genera: response to Margos\u2019 et al. \u201cThe genus Borrelia reloaded\u201d (PLoS ONE 13(12): e0208432)","volume":"14","author":"Gupta","year":"2019","journal-title":"PLoS One"},{"key":"2023122115210803900_pgad349-B17","doi-asserted-by":"crossref","first-page":"1180","DOI":"10.1128\/AEM.01913-06","article-title":"Identification of specific chemoattractants and genetic complementation of a Borrelia burgdorferi chemotaxis mutant: flow cytometry-based capillary tube chemotaxis assay","volume":"73","author":"Bakker","year":"2007","journal-title":"Appl Environ Microbiol"},{"key":"2023122115210803900_pgad349-B18","author":"Bakker","year":"2006"},{"key":"2023122115210803900_pgad349-B19","first-page":"355","article-title":"Chemotaxis in pathogenic spirochetes: directed movement toward targeting tissues?","volume":"2","author":"Lux","year":"2000","journal-title":"J Mol Microbiol Biotechnol"},{"key":"2023122115210803900_pgad349-B20","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1146\/annurev.genet.36.041602.134359","article-title":"Genetics of motility and chemotaxis of a fascinating group of Bacteria: the spirochetes","volume":"36","author":"Charon","year":"2002","journal-title":"Annu Rev Genet"},{"key":"2023122115210803900_pgad349-B21","doi-asserted-by":"crossref","first-page":"6276","DOI":"10.1128\/IAI.69.10.6276-6283.2001","article-title":"Motility and chemotaxis in tissue penetration of oral epithelial cell layers by Treponema denticola","volume":"69","author":"Lux","year":"2001","journal-title":"Infect Immun"},{"key":"2023122115210803900_pgad349-B22","doi-asserted-by":"crossref","first-page":"2485","DOI":"10.1128\/IAI.00145-12","article-title":"Borrelia burgdorferi needs chemotaxis to establish infection in mammals and to accomplish its enzootic cycle","volume":"80","author":"Sze","year":"2012","journal-title":"Infect Immun"},{"key":"2023122115210803900_pgad349-B23","first-page":"345","article-title":"Spirochete periplasmic flagella and motility","volume":"2","author":"Li","year":"2000","journal-title":"J Mol Microbiol Biotechnol"},{"key":"2023122115210803900_pgad349-B24","doi-asserted-by":"crossref","first-page":"634","DOI":"10.1038\/nrmicro2405","article-title":"A field guide to bacterial swarming motility","volume":"8","author":"Kearns","year":"2010","journal-title":"Nat Rev Microbiol"},{"key":"2023122115210803900_pgad349-B25","doi-asserted-by":"crossref","first-page":"550","DOI":"10.3390\/biom10040550","article-title":"Spirochete flagella and motility","volume":"10","author":"Nakamura","year":"2020","journal-title":"Biomolecules"},{"key":"2023122115210803900_pgad349-B26","doi-asserted-by":"crossref","first-page":"294","DOI":"10.1016\/j.tim.2022.09.010","article-title":"Diving into the complexity of the spirochetal endoflagellum","volume":"31","author":"San Martin","year":"2023","journal-title":"Trends Microbiol"},{"key":"2023122115210803900_pgad349-B27","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1016\/j.tim.2014.12.011","article-title":"The bacterial flagellar motor and its structural diversity","volume":"23","author":"Minamino","year":"2015","journal-title":"Trends Microbiol"},{"key":"2023122115210803900_pgad349-B28","doi-asserted-by":"crossref","first-page":"e2006989","DOI":"10.1371\/journal.pbio.2006989","article-title":"Hook length of the bacterial flagellum is optimized for maximal stability of the flagellar bundle","volume":"16","author":"Sp\u00f6ring","year":"2018","journal-title":"PLoS Biol"},{"key":"2023122115210803900_pgad349-B29","doi-asserted-by":"crossref","first-page":"13425","DOI":"10.1038\/ncomms13425","article-title":"Complete structure of the bacterial flagellar hook reveals extensive set of stabilizing interactions","volume":"7","author":"Matsunami","year":"2016","journal-title":"Nat Commun"},{"key":"2023122115210803900_pgad349-B30","doi-asserted-by":"crossref","first-page":"2972","DOI":"10.1038\/emboj.2011.186","article-title":"Structural diversity of bacterial flagellar motors","volume":"30","author":"Chen","year":"2011","journal-title":"EMBO J"},{"key":"2023122115210803900_pgad349-B31","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1111\/mmi.13544","article-title":"A novel glycan modifies the flagellar filament proteins of the oral bacterium Treponema denticola","volume":"103","author":"Kurniyati","year":"2017","journal-title":"Mol Microbiol"},{"key":"2023122115210803900_pgad349-B32","doi-asserted-by":"crossref","first-page":"2596","DOI":"10.1128\/JB.01502-09","article-title":"Differential regulation of the multiple flagellins in spirochetes","volume":"192","author":"Li","year":"2010","journal-title":"J Bacteriol"},{"key":"2023122115210803900_pgad349-B33","doi-asserted-by":"crossref","first-page":"5607","DOI":"10.1128\/JB.00319-08","article-title":"Genetic analysis of spirochete flagellin proteins and their involvement in motility, filament assembly, and flagellar morphology","volume":"190","author":"Li","year":"2008","journal-title":"J Bacteriol"},{"key":"2023122115210803900_pgad349-B34","doi-asserted-by":"crossref","first-page":"e53672","DOI":"10.7554\/eLife.53672","article-title":"An asymmetric sheath controls flagellar supercoiling and motility in the Leptospira spirochete","volume":"9","author":"Gibson","year":"2020","journal-title":"Elife"},{"key":"2023122115210803900_pgad349-B35","author":"Brady","year":"2022"},{"key":"2023122115210803900_pgad349-B36","doi-asserted-by":"crossref","first-page":"959","DOI":"10.1038\/s41589-019-0341-3","article-title":"Structure and chemistry of lysinoalanine crosslinking in the spirochaete flagella hook","volume":"15","author":"Lynch","year":"2019","journal-title":"Nat Chem Biol"},{"key":"2023122115210803900_pgad349-B37","doi-asserted-by":"crossref","first-page":"16134","DOI":"10.1038\/nmicrobiol.2016.134","article-title":"Spirochaete flagella hook proteins self-catalyse a lysinoalanine covalent crosslink for motility","volume":"1","author":"Miller","year":"2016","journal-title":"Nat Microbiol"},{"key":"2023122115210803900_pgad349-B38","doi-asserted-by":"crossref","first-page":"e98338","DOI":"10.1371\/journal.pone.0098338","article-title":"Initial characterization of the FlgE hook high molecular weight complex of Borrelia burgdorferi","volume":"9","author":"Miller","year":"2014","journal-title":"PLoS One"},{"key":"2023122115210803900_pgad349-B39","doi-asserted-by":"crossref","first-page":"928","DOI":"10.1038\/s41589-018-0122-4","article-title":"Substrate-assisted enzymatic formation of lysinoalanine in duramycin","volume":"14","author":"An","year":"2018","journal-title":"Nat Chem Biol"},{"key":"2023122115210803900_pgad349-B40","doi-asserted-by":"crossref","first-page":"474","DOI":"10.1021\/acs.biochem.8b01083","article-title":"Design, validation, and application of an enzyme-coupled hydrogen sulfide detection assay","volume":"59","author":"Lynch","year":"2019","journal-title":"Biochemistry"},{"key":"2023122115210803900_pgad349-B41","doi-asserted-by":"crossref","first-page":"5439","DOI":"10.1128\/jb.176.17.5439-5449.1994","article-title":"Roles of FliK and FlhB in determination of flagellar hook length in Salmonella typhimurium","volume":"176","author":"Hirano","year":"1994","journal-title":"J Bacteriol"},{"key":"2023122115210803900_pgad349-B42","doi-asserted-by":"publisher","author":"Gelberg","year":"2017","DOI":"10.1016\/B978-0-323-35775-3.00007-2"},{"key":"2023122115210803900_pgad349-B43","doi-asserted-by":"publisher","author":"Pillay","year":"1996","DOI":"10.1007\/978-1-62703-185-1_19"},{"key":"2023122115210803900_pgad349-B44","doi-asserted-by":"crossref","first-page":"390","DOI":"10.1016\/0076-6879(94)36029-4","article-title":"Culture of Treponema pallidum","volume":"236","author":"Cox","year":"1994","journal-title":"Methods Enzymol"},{"key":"2023122115210803900_pgad349-B45","doi-asserted-by":"crossref","first-page":"e01153","DOI":"10.1128\/mBio.01153-18","article-title":"Long-term in vitro culture of the syphilis spirochete Treponema pallidum subsp. pallidum","volume":"9","author":"Edmondson","year":"2018","journal-title":"mBio"},{"key":"2023122115210803900_pgad349-B46","doi-asserted-by":"crossref","first-page":"e0281187","DOI":"10.1371\/journal.pone.0281187","article-title":"Clonal isolates of Treponema pallidum subsp. pallidum Nichols provide evidence for the occurrence of microevolution during experimental rabbit infection and in vitro culture","volume":"18","author":"Edmondson","year":"2023","journal-title":"PLoS One"},{"key":"2023122115210803900_pgad349-B47","doi-asserted-by":"crossref","first-page":"e44","DOI":"10.1002\/cpz1.44","article-title":"In vitro cultivation of the syphilis spirochete Treponema pallidum","volume":"1","author":"Edmondson","year":"2021","journal-title":"Curr Protoc"},{"key":"2023122115210803900_pgad349-B48","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1038\/nrmicro.2017.5","article-title":"Virulence of the zoonotic agent of leptospirosis: still terra incognita?","volume":"15","author":"Picardeau","year":"2017","journal-title":"Nat Rev Microbiol"},{"key":"2023122115210803900_pgad349-B49","doi-asserted-by":"crossref","first-page":"816","DOI":"10.3389\/fmicb.2018.00816","article-title":"Biodiversity of environmental Leptospira: improving identification and revisiting the diagnosis","volume":"9","author":"Thibeaux","year":"2018","journal-title":"Front Microbiol"},{"key":"2023122115210803900_pgad349-B50","doi-asserted-by":"crossref","first-page":"005560","DOI":"10.1099\/ijsem.0.005560","article-title":"Leptospira sanjuanensis sp. nov., a pathogenic species of the genus Leptospira isolated from soil in Puerto Rico","volume":"72","author":"Fernandes","year":"2022","journal-title":"Int J Syst Evol Microbiol"},{"key":"2023122115210803900_pgad349-B51","doi-asserted-by":"crossref","first-page":"005148","DOI":"10.1099\/ijsem.0.005148","article-title":"Leptospira ainlahdjerensis sp. nov., Leptospira ainazelensis sp. nov. Leptospira abararensis sp. nov. and Leptospira chreensis sp. nov., four new species isolated from water sources in Algeria","volume":"71","author":"Korba","year":"2021","journal-title":"Int J Syst Evol Microbiol"},{"key":"2023122115210803900_pgad349-B52","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1007\/978-1-4615-4853-9_10","article-title":"Lysinoalanine in food and in antimicrobial proteins","volume":"459","author":"Friedman","year":"1999","journal-title":"Adv Exp Med Biol"},{"key":"2023122115210803900_pgad349-B53","doi-asserted-by":"crossref","first-page":"e0184648","DOI":"10.1371\/journal.pone.0184648","article-title":"Periplasmic flagella in Borrelia burgdorferi function to maintain cellular integrity upon external stress","volume":"12","author":"Kumar","year":"2017","journal-title":"PLoS One"},{"key":"2023122115210803900_pgad349-B54","first-page":"221","article-title":"The flagellar cytoskeleton of the spirochetes","volume":"11","author":"Wolgemuth","year":"2006","journal-title":"J Mol Microbiol Biotechnol"},{"key":"2023122115210803900_pgad349-B55","doi-asserted-by":"crossref","first-page":"1563","DOI":"10.1111\/j.1365-2958.2010.07078.x","article-title":"Inactivation of a putative flagellar motor switch protein FliG1 prevents Borrelia burgdorferi from swimming in highly viscous media and blocks its infectivity","volume":"75","author":"Li","year":"2010","journal-title":"Mol Microbiol"},{"key":"2023122115210803900_pgad349-B56","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1111\/mmi.13403","article-title":"A novel flagellar sheath protein, FcpA, determines filament coiling, translational motility and virulence for the Leptospira spirochete","volume":"101","author":"Wunder","year":"2016","journal-title":"Mol Microbiol"},{"key":"2023122115210803900_pgad349-B57","doi-asserted-by":"crossref","first-page":"2019","DOI":"10.1128\/IAI.00131-12","article-title":"Flaa proteins in Leptospira interrogans are essential for motility and virulence but are not required for formation of the flagellum sheath","volume":"80","author":"Lambert","year":"2012","journal-title":"Infect Immun"},{"key":"2023122115210803900_pgad349-B58","doi-asserted-by":"crossref","first-page":"5457","DOI":"10.1021\/acs.chemrev.6b00591","article-title":"Mechanistic understanding of lanthipeptide biosynthetic enzymes","volume":"117","author":"Repka","year":"2017","journal-title":"Chem Rev"},{"key":"2023122115210803900_pgad349-B59","doi-asserted-by":"crossref","first-page":"2273","DOI":"10.1016\/j.bpj.2013.10.004","article-title":"Viscous dynamics of Lyme disease and syphilis spirochetes reveal flagellar torque and drag","volume":"105","author":"Harman","year":"2013","journal-title":"Biophys J"},{"key":"2023122115210803900_pgad349-B60","doi-asserted-by":"crossref","first-page":"24433","DOI":"10.1073\/pnas.94.26.14433","article-title":"Absence of a barrier to backwards rotation of the bacterial flagellar motor demonstrated with optical tweezers","volume":"94","author":"Berry","year":"1997","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2023122115210803900_pgad349-B61","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/j.bpj.2013.11.1118","article-title":"Direct measurement of helical cell motion of the spirochete Leptospira","volume":"106","author":"Nakamura","year":"2014","journal-title":"Biophys J"},{"key":"2023122115210803900_pgad349-B62","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1038\/msb.2011.75","article-title":"Fast, scalable generation of high-quality protein multiple sequence alignments using Clustal Omega","volume":"7","author":"Sievers","year":"2011","journal-title":"Mol Syst Biol"},{"key":"2023122115210803900_pgad349-B63","doi-asserted-by":"crossref","first-page":"6097","DOI":"10.1093\/nar\/18.20.6097","article-title":"Sequence logos: a new way to display consensus sequences","volume":"18","author":"Schneider","year":"1990","journal-title":"Nucleic Acids Res"},{"key":"2023122115210803900_pgad349-B64","doi-asserted-by":"crossref","first-page":"1270","DOI":"10.1128\/IAI.00871-10","article-title":"Inactivation of bb0184, which encodes carbon storage regulator A, represses the infectivity of Borrelia burgdorferi","volume":"79","author":"Sze","year":"2011","journal-title":"Infect Immun"},{"key":"2023122115210803900_pgad349-B65","doi-asserted-by":"crossref","first-page":"6698","DOI":"10.1128\/JB.182.23.6698-6706.2000","article-title":"The spirochete FlaA periplasmic flagellar sheath protein impacts flagellar helicity","volume":"182","author":"Li","year":"2000","journal-title":"J Bacteriol"},{"key":"2023122115210803900_pgad349-B66","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1128\/jb.166.1.105-112.1986","article-title":"Treponema phagedenis has at least two proteins residing together on its periplasmic flagella","volume":"166","author":"Limberger","year":"1986","journal-title":"J Bacteriol"},{"key":"2023122115210803900_pgad349-B67","doi-asserted-by":"crossref","first-page":"2105","DOI":"10.1128\/IAI.00094-16","article-title":"Real-time PCR reveals rapid dissemination of Leptospira interrogans after intraperitoneal and conjunctival inoculation of hamsters","volume":"84","author":"Wunder","year":"2016","journal-title":"Infect Immun"},{"key":"2023122115210803900_pgad349-B68","doi-asserted-by":"crossref","first-page":"384","DOI":"10.1128\/jb.105.1.384-395.1971","article-title":"Fine structure and isolation of the hook-basal body complex of flagella from Escherichia coli and Bacillus subtilis","volume":"105","author":"DePamphilis","year":"1971","journal-title":"J Bacteriol"},{"key":"2023122115210803900_pgad349-B69","doi-asserted-by":"crossref","first-page":"1122","DOI":"10.1111\/mmi.14482","article-title":"Flhf regulates the number and configuration of periplasmic flagella in Borrelia burgdorferi","volume":"113","author":"Zhang","year":"2020","journal-title":"Mol Microbiol"},{"key":"2023122115210803900_pgad349-B70","doi-asserted-by":"crossref","first-page":"6169","DOI":"10.1073\/pnas.092010499","article-title":"Asymmetrical flagellar rotation in Borrelia burgdorferi nonchemotactic mutants","volume":"99","author":"Li","year":"2002","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2023122115210803900_pgad349-B71","doi-asserted-by":"crossref","first-page":"D439","DOI":"10.1093\/nar\/gkab1061","article-title":"AlphaFold Protein Structure Database: massively expanding the structural coverage of protein-sequence space with high-accuracy models","volume":"50","author":"Varadi","year":"2022","journal-title":"Nucleic Acids Res"},{"key":"2023122115210803900_pgad349-B72","doi-asserted-by":"crossref","first-page":"260","DOI":"10.3390\/biom9070260","article-title":"Architecture of the bacterial flagellar distal rod and hook of Salmonella","volume":"9","author":"Saijo-Hamano","year":"2019","journal-title":"Biomolecules"},{"key":"2023122115210803900_pgad349-B73"}],"container-title":["PNAS Nexus"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/pnasnexus\/advance-article-pdf\/doi\/10.1093\/pnasnexus\/pgad349\/52600767\/pgad349.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/pnasnexus\/article-pdf\/2\/12\/pgad349\/54579502\/pgad349.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/pnasnexus\/article-pdf\/2\/12\/pgad349\/54579502\/pgad349.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,12,21]],"date-time":"2023-12-21T10:36:32Z","timestamp":1703154992000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/pnasnexus\/article\/doi\/10.1093\/pnasnexus\/pgad349\/7330553"}},"subtitle":[],"editor":[{"given":"Bruce","family":"Levine","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2023,10,26]]},"references-count":73,"journal-issue":{"issue":"12","published-print":{"date-parts":[[2023,12,1]]}},"URL":"https:\/\/doi.org\/10.1093\/pnasnexus\/pgad349","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2023.06.13.544825","asserted-by":"object"}]},"ISSN":["2752-6542"],"issn-type":[{"value":"2752-6542","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2023,12]]},"published":{"date-parts":[[2023,10,26]]},"article-number":"pgad349"}}