{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,19]],"date-time":"2026-02-19T08:26:49Z","timestamp":1771489609143,"version":"3.50.1"},"reference-count":39,"publisher":"American Society for Microbiology","issue":"14","license":[{"start":{"date-parts":[[2007,7,15]],"date-time":"2007-07-15T00:00:00Z","timestamp":1184457600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/journals.asm.org\/non-commercial-tdm-license"}],"content-domain":{"domain":["journals.asm.org"],"crossmark-restriction":true},"short-container-title":["J Bacteriol"],"published-print":{"date-parts":[[2007,7,15]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n          <jats:p>\n            Mutations in\n            <jats:italic>pmrA<\/jats:italic>\n            were recombined into\n            <jats:italic>Salmonella<\/jats:italic>\n            strain ATCC 14028\n            <jats:italic>msbB<\/jats:italic>\n            to determine if\n            <jats:italic>pmrA<\/jats:italic>\n            -regulated modifications of lipopolysaccharide could suppress\n            <jats:italic>msbB<\/jats:italic>\n            growth defects. A mutation that functions to constitutively activate\n            <jats:italic>pmrA<\/jats:italic>\n            [\n            <jats:italic>pmrA<\/jats:italic>\n            (Con)] suppresses\n            <jats:italic>msbB<\/jats:italic>\n            growth defects on EGTA-containing media. Lipid A structural analysis showed that\n            <jats:italic>Salmonella msbB pmrA<\/jats:italic>\n            (Con) strains, compared to\n            <jats:italic>Salmonella msbB<\/jats:italic>\n            strains, have increased amounts of palmitate and phosphoethanolamine but no aminoarabinose addition, suggesting that aminoarabinose is not incorporated into\n            <jats:italic>msbB<\/jats:italic>\n            lipid A. Surprisingly, loss-of-function mutations in the aminoarabinose biosynthetic genes restored EGTA and polymyxin sensitivity to\n            <jats:italic>Salmonella msbB pmrA<\/jats:italic>\n            (Con) strains. These blocks in aminoarabinose biosynthesis also prevented lipid A phosphoethanolamine incorporation and reduced the levels of palmitate addition, indicating previously unknown roles for the aminoarabinose biosynthetic enzymes. Lipid A structural analysis of the EGTA- and polymyxin-resistant triple mutant\n            <jats:italic>msbB pmrA<\/jats:italic>\n            (Con)\n            <jats:italic>pagP<\/jats:italic>\n            ::Tn\n            <jats:italic>10<\/jats:italic>\n            , which contains phosphoethanolamine but no palmitoylated lipid A, suggests that phosphoethanolamine addition is sufficient to confer EGTA and polymyxin resistance on\n            <jats:italic>Salmonella msbB<\/jats:italic>\n            strains. Additionally, palmitoylated lipid A was observed only in wild-type\n            <jats:italic>Salmonella<\/jats:italic>\n            grown in the presence of salt in rich media. Thus, we correlate EGTA resistance and polymyxin resistance with phosphoethanolamine-decorated lipid A and demonstrate that the aminoarabinose biosynthetic proteins play an essential role in lipid A phosphoethanolamine addition and affect lipid A palmitate addition in\n            <jats:italic>Salmonella msbB<\/jats:italic>\n            strains.\n          <\/jats:p>","DOI":"10.1128\/jb.01969-06","type":"journal-article","created":{"date-parts":[[2007,4,21]],"date-time":"2007-04-21T00:41:19Z","timestamp":1177116079000},"page":"5161-5169","update-policy":"https:\/\/doi.org\/10.1128\/asmj-crossmark-policy-page","source":"Crossref","is-referenced-by-count":28,"title":["PmrA(Con) Confers\n            <i>pmrHFIJKL<\/i>\n            -Dependent EGTA and Polymyxin Resistance on\n            <i>msbB Salmonella<\/i>\n            by Decorating Lipid A with Phosphoethanolamine"],"prefix":"10.1128","volume":"189","author":[{"given":"Sean R.","family":"Murray","sequence":"first","affiliation":[{"name":"Department of Biology, Yale University, New Haven, Connecticut"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Robert K.","family":"Ernst","sequence":"additional","affiliation":[{"name":"Department of Microbiology, University of Washington, Seattle, Washington"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"David","family":"Bermudes","sequence":"additional","affiliation":[{"name":"Vion Pharmaceuticals Inc., New Haven, Connecticut"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Samuel I.","family":"Miller","sequence":"additional","affiliation":[{"name":"Department of Microbiology, University of Washington, Seattle, Washington"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"K. Brooks","family":"Low","sequence":"additional","affiliation":[{"name":"Department of Therapeutic Radiology, Yale University School of Medicine, New Haven, Connecticut"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"235","reference":[{"key":"e_1_3_2_2_2","doi-asserted-by":"crossref","first-page":"5071","DOI":"10.1093\/emboj\/19.19.5071","volume":"19","year":"2000","unstructured":"Bishop, R. E., H. S. Gibbons, T. Guina, M. S. Trent, S. I. Miller, and C. R. Raetz. 2000. Transfer of palmitate from phospholipids to lipid A in outer membranes of gram-negative bacteria. EMBO J. 19 : 5071-5080.","journal-title":"EMBO J."},{"key":"e_1_3_2_3_2","doi-asserted-by":"crossref","first-page":"14154","DOI":"10.1074\/jbc.M414265200","volume":"280","year":"2005","unstructured":"Breazeale, S. D., A. A. Ribeiro, A. L. McClerren, and C. R. Raetz. 2005. A formyltransferase required for polymyxin resistance in Escherichia coli and the modification of lipid A with 4-amino-4-deoxy-l-arabinose. Identification and function oF UDP-4-deoxy-4-formamido-l-arabinose. J. Biol. Chem. 280 : 14154-14167.","journal-title":"J. Biol. Chem."},{"key":"e_1_3_2_4_2","doi-asserted-by":"crossref","first-page":"24731","DOI":"10.1074\/jbc.M304043200","volume":"278","year":"2003","unstructured":"Breazeale, S. D., A. A. Ribeiro, and C. R. Raetz. 2003. Origin of lipid A species modified with 4-amino-4-deoxy-l-arabinose in polymyxin-resistant mutants of Escherichia coli. An aminotransferase (ArnB) that generates UDP-4-deoxyl-l-arabinose. J. Biol. Chem. 278 : 24731-24739.","journal-title":"J. Biol. Chem."},{"key":"e_1_3_2_5_2","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1016\/0008-6215(88)84149-1","volume":"175","year":"1988","unstructured":"Caroff, M., A. Tacken, and L. Szabo. 1988. Detergent-accelerated hydrolysis of bacterial endotoxins and determination of the anomeric configuration of the glycosyl phosphate present in the \u201cisolated lipid A\u201d fragment of the Bordetella pertussis endotoxin. Carbohydr. Res. 175 : 273-282.","journal-title":"Carbohydr. Res."},{"key":"e_1_3_2_6_2","doi-asserted-by":"publisher","DOI":"10.1128\/jb.155.2.831-838.1983"},{"key":"e_1_3_2_7_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.120163297"},{"key":"e_1_3_2_8_2","unstructured":"Advanced bacterial genetics. 1980"},{"key":"e_1_3_2_9_2","doi-asserted-by":"crossref","first-page":"1561","DOI":"10.1126\/science.286.5444.1561","volume":"286","year":"1999","unstructured":"Ernst, R. K., E. C. Yi, L. Guo, K. B. Lim, J. L. Burns, M. Hackett, and S. I. Miller. 1999. Specific lipopolysaccharide found in cystic fibrosis airway Pseudomonas aeruginosa. Science 286 : 1561-1565.","journal-title":"Science"},{"key":"e_1_3_2_10_2","doi-asserted-by":"crossref","first-page":"1171","DOI":"10.1046\/j.1365-2958.1998.00757.x","volume":"27","year":"1998","unstructured":"Gunn, J. S., K. B. Lim, J. Krueger, K. Kim, L. Guo, M. Hackett, and S. I. Miller. 1998. PmrA-PmrB-regulated genes necessary for 4-aminoarabinose lipid A modification and polymyxin resistance. Mol. Microbiol. 27 : 1171-1182.","journal-title":"Mol. Microbiol."},{"key":"e_1_3_2_11_2","doi-asserted-by":"publisher","DOI":"10.1128\/jb.178.23.6857-6864.1996"},{"key":"e_1_3_2_12_2","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.68.11.6139-6146.2000"},{"key":"e_1_3_2_13_2","doi-asserted-by":"crossref","first-page":"250","DOI":"10.1126\/science.276.5310.250","volume":"276","year":"1997","unstructured":"Guo, L., K. B. Lim, J. S. Gunn, B. Bainbridge, R. P. Darveau, M. Hackett, and S. I. Miller. 1997. Regulation of lipid A modifications by Salmonella typhimurium virulence genes phoP-phoQ. Science 276 : 250-253.","journal-title":"Science"},{"key":"e_1_3_2_14_2","doi-asserted-by":"crossref","first-page":"481","DOI":"10.1111\/j.1365-2958.1994.tb00329.x","volume":"11","year":"1994","unstructured":"Helander, I. M., I. Kilpelainen, and M. Vaara. 1994. Increased substitution of phosphate groups in lipopolysaccharides and lipid A of the polymyxin-resistant pmrA mutants of Salmonella typhimurium: a 31P-NMR study. Mol. Microbiol. 11 : 481-487.","journal-title":"Mol. Microbiol."},{"key":"e_1_3_2_15_2","doi-asserted-by":"crossref","first-page":"44966","DOI":"10.1074\/jbc.M404963200","volume":"279","year":"2004","unstructured":"Jia, W., A. E. Zoeiby, T. N. Petruzziello, B. Jayabalasingham, S. Seyedirashti, and R. E. Bishop. 2004. Lipid trafficking controls endotoxin acylation in outer membranes of Escherichia coli. J. Biol. Chem. 279 : 44966-44975.","journal-title":"J. Biol. Chem."},{"key":"e_1_3_2_16_2","doi-asserted-by":"crossref","first-page":"2302","DOI":"10.1101\/gad.1230804","volume":"18","year":"2004","unstructured":"Kato, A., and E. A. Groisman. 2004. Connecting two-component regulatory systems by a protein that protects a response regulator from dephosphorylation by its cognate sensor. Genes Dev. 18 : 2302-2313.","journal-title":"Genes Dev."},{"key":"e_1_3_2_17_2","doi-asserted-by":"crossref","first-page":"571","DOI":"10.1046\/j.1365-2958.1998.00952.x","volume":"29","year":"1998","unstructured":"Khan, S. A., P. Everest, S. Servos, N. Foxwell, U. Zahringer, H. Brade, E. T. Rietschel, G. Dougan, I. G. Charles, and D. J. Maskell. 1998. A lethal role for lipid A in Salmonella infections. Mol. Microbiol. 29 : 571-579.","journal-title":"Mol. Microbiol."},{"key":"e_1_3_2_18_2","doi-asserted-by":"crossref","first-page":"4422","DOI":"10.1073\/pnas.80.14.4422","volume":"80","year":"1983","unstructured":"Kolodkin, A. L., M. A. Capage, E. I. Golub, and K. B. Low. 1983. F sex factor of Escherichia coli K-12 codes for a single-stranded DNA binding protein. Proc. Natl. Acad. Sci. USA 80 : 4422-4426.","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"e_1_3_2_19_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.186.13.4124-4133.2004"},{"key":"e_1_3_2_20_2","first-page":"37","volume":"17","year":"1999","unstructured":"Low, K. B., M. Ittensohn, T. Le, J. Platt, S. Sodi, M. Amoss, O. Ash, E. Carmichael, A. Chakraborty, J. Fischer, S. L. Lin, X. Luo, S. I. Miller, L. Zheng, I. King, J. M. Pawelek, and D. Bermudes. 1999. Lipid A mutant Salmonella with suppressed virulence and TNF\u03b1 induction retain tumor-targeting in vivo. Nat. Biotechnol. 17 : 37-41.","journal-title":"Nat. Biotechnol."},{"key":"e_1_3_2_21_2","first-page":"47","volume":"90","year":"2004","unstructured":"Low, K. B., M. Ittensohn, X. Luo, L. M. Zheng, I. King, J. M. Pawelek, and D. Bermudes. 2004. Construction of VNP20009: a novel, genetically stable antibiotic-sensitive strain of tumor-targeting Salmonella for parenteral administration in humans. Methods Mol. Med. 90 : 47-60.","journal-title":"Methods Mol. Med."},{"key":"e_1_3_2_22_2","unstructured":"Experiments in molecular genetics. 1972"},{"key":"e_1_3_2_23_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.183.19.5554-5561.2001"},{"key":"e_1_3_2_24_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.186.24.8516-8523.2004"},{"key":"e_1_3_2_25_2","first-page":"29","year":"1996","unstructured":"Nikaido, H. 1996. Outer membrane, p. 29-47. In F. Neidhardt, R. Curtis, J. Ingraham, E. C. C. Lin, K. B. Low, B. Magasanik, W. Reznikoff, M. Riley, M. Schaechter, and H. E. Umbarger (ed.), Escherichia coli and Salmonella: cellular and molecular biology, 2nd ed. ASM Press, Washington, DC.","journal-title":"Escherichia coli and Salmonella: cellular and molecular biology"},{"key":"e_1_3_2_26_2","doi-asserted-by":"crossref","first-page":"156","DOI":"10.1007\/BF00315809","volume":"223","year":"1990","unstructured":"O'Callaghan, D., and A. Charbit. 1990. High efficiency transformation of Salmonella typhimurium and Salmonella typhi by electroporation. Mol. Gen. Genet. 223 : 156-158.","journal-title":"Mol. Gen. Genet."},{"key":"e_1_3_2_27_2","doi-asserted-by":"publisher","DOI":"10.1128\/jb.175.13.4154-4164.1993"},{"key":"e_1_3_2_28_2","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1172\/JCI118423","volume":"97","year":"1996","unstructured":"Somerville, J. E., Jr., L. Cassiano, B. Bainbridge, M. D. Cunningham, and R. P. Darveau. 1996. A novel Escherichia coli lipid A mutant that produces an antiinflammatory lipopolysaccharide. J. Clin. Investig. 97 : 359-365.","journal-title":"J. Clin. Investig."},{"key":"e_1_3_2_29_2","doi-asserted-by":"publisher","DOI":"10.1128\/jb.178.23.6796-6801.1996"},{"key":"e_1_3_2_30_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.187.10.3391-3399.2005"},{"key":"e_1_3_2_31_2","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.70.12.6770-6778.2002"},{"key":"e_1_3_2_32_2","doi-asserted-by":"crossref","first-page":"28186","DOI":"10.1074\/jbc.M505020200","volume":"280","year":"2005","unstructured":"Tran, A. X., M. E. Lester, C. M. Stead, C. R. Raetz, D. J. Maskell, S. C. McGrath, R. J. Cotter, and M. S. Trent. 2005. Resistance to the antimicrobial peptide polymyxin requires myristoylation of Escherichia coli and Salmonella typhimurium lipid A. J. Biol. Chem. 280 : 28186-28194.","journal-title":"J. Biol. Chem."},{"key":"e_1_3_2_33_2","doi-asserted-by":"crossref","first-page":"43132","DOI":"10.1074\/jbc.M106962200","volume":"276","year":"2001","unstructured":"Trent, M. S., A. A. Ribeiro, W. T. Doerrler, S. Lin, R. J. Cotter, and C. R. Raetz. 2001. Accumulation of a polyisoprene-linked amino sugar in polymyxin-resistant Salmonella typhimurium and Escherichia coli: structural characterization and transfer to lipid A in the periplasm. J. Biol. Chem. 276 : 43132-43144.","journal-title":"J. Biol. Chem."},{"key":"e_1_3_2_34_2","doi-asserted-by":"crossref","first-page":"43122","DOI":"10.1074\/jbc.M106961200","volume":"276","year":"2001","unstructured":"Trent, M. S., A. A. Ribeiro, S. Lin, R. J. Cotter, and C. R. Raetz. 2001. An inner membrane enzyme in Salmonella and Escherichia coli that transfers 4-amino-4-deoxy-l-arabinose to lipid A: induction on polymyxin-resistant mutants and role of a novel lipid-linked donor. J. Biol. Chem. 276 : 43122-43131.","journal-title":"J. Biol. Chem."},{"key":"e_1_3_2_35_2","doi-asserted-by":"publisher","DOI":"10.1128\/jb.148.2.426-434.1981"},{"key":"e_1_3_2_36_2","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/0014-5793(81)80777-6","volume":"129","year":"1981","unstructured":"Vaara, M., T. Vaara, M. Jensen, I. Helander, M. Nurminen, E. T. Rietschel, and P. H. Makela. 1981. Characterization of the lipopolysaccharide from the polymyxin-resistant pmrA mutants of Salmonella typhimurium. FEBS Lett. 129 : 145-149.","journal-title":"FEBS Lett."},{"key":"e_1_3_2_37_2","doi-asserted-by":"crossref","first-page":"23000","DOI":"10.1074\/jbc.M501534200","volume":"280","year":"2005","unstructured":"Williams, G. J., S. D. Breazeale, C. R. Raetz, and J. H. Naismith. 2005. Structure and function of both domains of ArnA, a dual function decarboxylase and a formyltransferase, involved in 4-amino-4-deoxy-l-arabinose biosynthesis. J. Biol. Chem. 280 : 23000-23008.","journal-title":"J. Biol. Chem."},{"key":"e_1_3_2_38_2","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1016\/S0092-8674(00)00092-1","volume":"103","year":"2000","unstructured":"Wosten, M. M., L. F. Kox, S. Chamnongpol, F. C. Soncini, and E. A. Groisman. 2000. A signal transduction system that responds to extracellular iron. Cell 103 : 113-125.","journal-title":"Cell"},{"key":"e_1_3_2_39_2","doi-asserted-by":"crossref","first-page":"18503","DOI":"10.1074\/jbc.274.26.18503","volume":"274","year":"1999","unstructured":"Zhou, Z., S. Lin, R. J. Cotter, and C. R. Raetz. 1999. Lipid A modifications characteristic of Salmonella typhimurium are induced by NH4VO3 in Escherichia coli K12. Detection of 4-amino-4-deoxy-l-arabinose, phosphoethanolamine and palmitate. J. Biol. Chem. 274 : 18503-18514.","journal-title":"J. Biol. Chem."},{"key":"e_1_3_2_40_2","doi-asserted-by":"crossref","first-page":"43111","DOI":"10.1074\/jbc.M106960200","volume":"276","year":"2001","unstructured":"Zhou, Z., A. A. Ribeiro, S. Lin, R. J. Cotter, S. I. Miller, and C. R. Raetz. 2001. Lipid A modifications in polymyxin-resistant Salmonella typhimurium: PMRA-dependent 4-amino-4-deoxy-l-arabinose, and phosphoethanolamine incorporation. J. Biol. Chem. 276 : 43111-43121.","journal-title":"J. Biol. Chem."}],"container-title":["Journal of Bacteriology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.asm.org\/doi\/pdf\/10.1128\/JB.01969-06","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/journals.asm.org\/doi\/pdf\/10.1128\/JB.01969-06","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,7,29]],"date-time":"2021-07-29T17:34:51Z","timestamp":1627580091000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.asm.org\/doi\/10.1128\/JB.01969-06"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2007,7,15]]},"references-count":39,"journal-issue":{"issue":"14","published-print":{"date-parts":[[2007,7,15]]}},"alternative-id":["10.1128\/JB.01969-06"],"URL":"https:\/\/doi.org\/10.1128\/jb.01969-06","relation":{},"ISSN":["0021-9193","1098-5530"],"issn-type":[{"value":"0021-9193","type":"print"},{"value":"1098-5530","type":"electronic"}],"subject":[],"published":{"date-parts":[[2007,7,15]]},"assertion":[{"value":"2006-12-29","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2007-04-06","order":1,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2020-12-20","order":2,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}]}}