Flagellin Glycosylation in Pseudomonas aeruginosa PAK Requires the O-antigen Biosynthesis Enzyme WbpO

Pseudomonas aeruginosa PAK (serotype O6) produces a single polar, glycosylated flagellum composed of a-type flagellin. To determine whether or not flagellin glycosylation in this serotype requires O-antigen genes, flagellin was isolated from the wild type, three O-antigen-deficient mutants wbpL, wbpO, and wbpP, and a wbpO mutant complemented with a plasmid containing a wild-type copy of wbpO. Flagellin from the wbpO mutant was smaller (42 kDa) than that of the wild type (45 kDa), or other mutants strains, and exhibited an altered isoelectric point (pI 4.8) when compared with PAK flagellin (pI 4.6). These differences were because of the truncation of the glycan moiety in the wbpO-flagellin. Thus, flagellin glycosylation in P. aeruginosa PAK apparently requires a functional WbpO but not WbpP. Because WbpP was previously proposed to catalyze a metabolic step in the biosynthesis of B-band O-antigen that precedes the action of WbpO, these results prompted us to reevaluate the two-step pathway catalyzed by WbpO and WbpP. Results from WbpO-WbpP-coupled enzymatic assays showed that either WbpO or WbpP is capable of initiating the two-step pathway; however, the kinetic parameters favored the WbpO reaction to occur first, converting UDP-N-acetyl-d-glucosamine to UDP-N-acetyl-d-glucuronic acid prior to the conversion to UDP-N-acetyl-d-galacturonic acid by WbpP. This is the first report to show that a C4 epimerase could utilize UDP-N-acetylhexuronic acid as a substrate. Pseudomonas aeruginosa PAK (serotype O6) produces a single polar, glycosylated flagellum composed of a-type flagellin. To determine whether or not flagellin glycosylation in this serotype requires O-antigen genes, flagellin was isolated from the wild type, three O-antigen-deficient mutants wbpL, wbpO, and wbpP, and a wbpO mutant complemented with a plasmid containing a wild-type copy of wbpO. Flagellin from the wbpO mutant was smaller (42 kDa) than that of the wild type (45 kDa), or other mutants strains, and exhibited an altered isoelectric point (pI 4.8) when compared with PAK flagellin (pI 4.6). These differences were because of the truncation of the glycan moiety in the wbpO-flagellin. Thus, flagellin glycosylation in P. aeruginosa PAK apparently requires a functional WbpO but not WbpP. Because WbpP was previously proposed to catalyze a metabolic step in the biosynthesis of B-band O-antigen that precedes the action of WbpO, these results prompted us to reevaluate the two-step pathway catalyzed by WbpO and WbpP. Results from WbpO-WbpP-coupled enzymatic assays showed that either WbpO or WbpP is capable of initiating the two-step pathway; however, the kinetic parameters favored the WbpO reaction to occur first, converting UDP-N-acetyl-d-glucosamine to UDP-N-acetyl-d-glucuronic acid prior to the conversion to UDP-N-acetyl-d-galacturonic acid by WbpP. This is the first report to show that a C4 epimerase could utilize UDP-N-acetylhexuronic acid as a substrate. Nucleotide-activated monosaccharides serve as precursors to a number of cell-surface glycoconjugates, including lipopolysaccharide (LPS), 8The abbreviations used are: LPSlipopolysaccharideCEcapillary electrophoresisCOSYcorrelation spectroscopyd-QuiNAcN-acetyl-d-quinovosamineIPTGisopropyl β-d-thiogalactopyranosideIEFisoelectric focusingMALDI-TOFmatrix-assisted laser desorption ionization-time of flightMES2-morpholinoethanesulfonic acidMSmass spectrometryNBTnitro blue tetrazoliumNOESYnuclear Overhauser effect spectroscopyPBSphosphate-buffered salineTBSTris-buffered salineTOCSYtotal correlation spectroscopyUDP-d-GalNAcAUDP-N-acetyl-d-galacturonic acidUDP-d-GalNAc-ANUDP-2-acetamido-6-amino-2-deoxy-d-galactoseUDP-d-GlcNAcUDP-N-acetyl-d-glucosamineUDP-d-GlcNAcAUDP-N-acetyl-d-glucuronic acidUDP-d-QuiNAcUDP-N-acetyl-d-quinovosamineBisTris propane1,3-bis[tris(hydroxymethyl)methylamino]propane. 8The abbreviations used are: LPSlipopolysaccharideCEcapillary electrophoresisCOSYcorrelation spectroscopyd-QuiNAcN-acetyl-d-quinovosamineIPTGisopropyl β-d-thiogalactopyranosideIEFisoelectric focusingMALDI-TOFmatrix-assisted laser desorption ionization-time of flightMES2-morpholinoethanesulfonic acidMSmass spectrometryNBTnitro blue tetrazoliumNOESYnuclear Overhauser effect spectroscopyPBSphosphate-buffered salineTBSTris-buffered salineTOCSYtotal correlation spectroscopyUDP-d-GalNAcAUDP-N-acetyl-d-galacturonic acidUDP-d-GalNAc-ANUDP-2-acetamido-6-amino-2-deoxy-d-galactoseUDP-d-GlcNAcUDP-N-acetyl-d-glucosamineUDP-d-GlcNAcAUDP-N-acetyl-d-glucuronic acidUDP-d-QuiNAcUDP-N-acetyl-d-quinovosamineBisTris propane1,3-bis[tris(hydroxymethyl)methylamino]propane. exopolysaccharide, and glycosylated proteins of pili and flagella. In some cases, these precursors may be shared between different systems. In Neisseria meningitidis, for example, more than three unlinked loci are required for synthesis and attachment of the pilin glycan (1Power P.M. Roddam L.F. Dieckelmann M. Srikhanta Y.N. Tan Y.C. Berrington A.W. Jennings M.P. Microbiology. 2000; 146: 967-979Crossref PubMed Scopus (73) Google Scholar, 2Power P.M. Roddam L.F. Rutter K. Fitzpatrick S.Z. Srikhanta Y.N. Jennings M.P. Mol. Microbiol. 2003; 49: 833-847Crossref PubMed Scopus (104) Google Scholar). One of these loci includes galE, a gene that is required for synthesis of UDP-galactose, which in turn is required both for the production of lipo-oligosaccharide and for glycosylation of the pilin subunits (3Stimson E. Virji M. Makepeace K. Dell A. Morris H.R. Payne G. Saunders J.R. Jennings M.P. Barker S. Panico M. Blench I. Moxon E.R. Mol. Microbiol. 1995; 17: 1201-1214Crossref PubMed Scopus (218) Google Scholar). In Pseudomonas aeruginosa strains, dTDP-l-rhamnose is channeled into synthesis of numerous l-rhamnose-containing cell-surface structures, including rhamnolipids (4Rahim R. Ochsner U.A. Olvera C. Graninger M. Messner P. Lam J.S. Soberon-Chavez G. Mol. Microbiol. 2001; 40: 708-718Crossref PubMed Scopus (192) Google Scholar), core oligosaccharide, and O-antigen polysaccharide (5Rahim R. Burrows L.L. Monteiro M.A. Perry M.B. Lam J.S. Microbiology. 2000; 146: 2803-2814Crossref PubMed Scopus (93) Google Scholar). This sugar-nucleotide is synthesized by proteins encoded in the rmlBDAC gene cluster, which is unlinked to the gene clusters that are required for synthesis of these glycoconjugates. lipopolysaccharide capillary electrophoresis correlation spectroscopy N-acetyl-d-quinovosamine isopropyl β-d-thiogalactopyranoside isoelectric focusing matrix-assisted laser desorption ionization-time of flight 2-morpholinoethanesulfonic acid mass spectrometry nitro blue tetrazolium nuclear Overhauser effect spectroscopy phosphate-buffered saline Tris-buffered saline total correlation spectroscopy UDP-N-acetyl-d-galacturonic acid UDP-2-acetamido-6-amino-2-deoxy-d-galactose UDP-N-acetyl-d-glucosamine UDP-N-acetyl-d-glucuronic acid UDP-N-acetyl-d-quinovosamine 1,3-bis[tris(hydroxymethyl)methylamino]propane. lipopolysaccharide capillary electrophoresis correlation spectroscopy N-acetyl-d-quinovosamine isopropyl β-d-thiogalactopyranoside isoelectric focusing matrix-assisted laser desorption ionization-time of flight 2-morpholinoethanesulfonic acid mass spectrometry nitro blue tetrazolium nuclear Overhauser effect spectroscopy phosphate-buffered saline Tris-buffered saline total correlation spectroscopy UDP-N-acetyl-d-galacturonic acid UDP-2-acetamido-6-amino-2-deoxy-d-galactose UDP-N-acetyl-d-glucosamine UDP-N-acetyl-d-glucuronic acid UDP-N-acetyl-d-quinovosamine 1,3-bis[tris(hydroxymethyl)methylamino]propane. Another example of the interrelationship between biosynthesis pathways for cell-surface glycoconjugates is pilin glycosylation in P. aeruginosa strain 1244 (serotype O7). The covalently attached pilin glycan is identical to the repeat unit from the B-band O-antigen, one of two types of O-antigen possessed by P. aeruginosa strains (6DiGiandomenico A. Matewish M.J. Bisaillon A. Stehle J.R. Lam J.S. Castric P. Mol. Microbiol. 2002; 46: PubMed Scopus Google Scholar). In a in the gene that the initiating which the first of the O-antigen unit from UDP-N-acetyl-d-quinovosamine to the pilin glycosylation as as O-antigen P. aeruginosa produces a single flagellum that to be an in and is for of the in with and with E. R. M. A. PubMed Google Scholar, M. R. S. S. A. PubMed Google Scholar). The is composed of flagellin subunits encoded by and in this flagellin is as either a-type or acid and C. C. PubMed Scopus Google Scholar, J.S. M. 49: PubMed Google Scholar, R. A. Scholar, Microbiol. 17: Google Scholar). the pilin of P. aeruginosa the a-type of strain PAK (serotype O6) are is the of the glycosylation PubMed Google Scholar, M. A. E. P. R. PubMed Scopus Google Scholar, S. PubMed Google Scholar). The flagellin of strain PAK two with to monosaccharides M. A. E. P. R. PubMed Scopus Google Scholar). the of the flagellin is that glycan with an and with and of genes, of the flagellin gene in the PAK as the flagellin glycosylation M. S. R. S. A. 2001; PubMed Scopus Google Scholar). of the in this to be required for flagellin but the of the and because of the of the of the is not to a biosynthesis pathway this The of this was to determine whether flagellin glycosylation in P. aeruginosa PAK (serotype O6) requires in B-band O-antigen biosynthesis as pilin glycosylation in strain The for biosynthesis of the serotype O-antigen and as the gene M. Burrows L.L. Lam J.S. Microbiology. PubMed Scopus Google Scholar), and the of the O-antigen of strain PAK is a repeat of A. Google Scholar). isolated and compared flagellin from wild-type PAK and three wbpL, wbpO, and in these genes, the O-antigen cluster, and of the from these mutants showed the of these for O-antigen production M. Burrows L.L. Lam J.S. Microbiology. PubMed Scopus Google Scholar). is the initiating of O-antigen biosynthesis and the first of the N-acetyl-d-quinovosamine from to the WbpP and WbpO are required for synthesis of the UDP-N-acetyl-d-galacturonic acid a to the of the WbpP as a UDP-N-acetyl-d-glucosamine C. M. Lam J.S. 2000; PubMed Scopus Google Scholar), and WbpO as a that the of and produces C. M. E. Lam J.S. 2000; PubMed Scopus Google Scholar). In this show that WbpO is required for flagellin but WbpP is This is because WbpP was proposed to catalyze an step in the metabolic pathway than WbpO, and prompted us to more of the metabolic catalyzed by these two proteins in the O-antigen biosynthesis in P. aeruginosa strain the to show that either WbpP or WbpO could the two-step the kinetic parameters of the two a for WbpO to into UDP-N-acetyl-d-glucuronic acid prior to of this by WbpP to nitro blue tetrazolium and the used in this were from β-d-thiogalactopyranoside isoelectric focusing and the were from strains used in this PubMed Scopus Google and PubMed Scopus Google were used for the of and was to the of and PubMed Google the of PubMed Scopus Google Scholar), that a was used The were with were as by PubMed Scopus Google Scholar). electrophoresis was a with as previously C. S. J.R. Lam J.S. PubMed Scopus Google Scholar). was an containing were with an of and with a of and electrophoresis was to the the was in acid for and with a of with the The was with and as by the the proteins were from the electrophoresis to a for as by the mass spectrometry flagellin was first in and a of to acid in was and was the of the flagellin were the by of the and a of flagellin proteins was a matrix-assisted laser desorption ionization-time of flight with a laser of The was including and of flagellin was in the in the with laser of were isolated from the different P. aeruginosa strains in with and were isolated and the as by and PubMed Google Scholar). The flagellin was for by of from the P. aeruginosa mutants was more than from the wild-type strain because the mutants were more to the of the in of In the the was to the was to a of in and the were by in this were but of pilin were to be in the of from to P. aeruginosa strain PAK was by A. R. 2002; PubMed Scopus Google Scholar). To the the was proteins isolated from P. aeruginosa strain a mutant that not flagella. were from strain as of a in was for and the was in and in a for This was a and to The was into The was in of phosphate-buffered saline and of were in of for a The were and were and the was as This was with and the was The of the for was by flagellin from the wild-type PAK and as a Flagellin were a and The glycosylation of PubMed Scopus Google was used with the of the proteins to the the was with of for The was in in for in the The was for with by a in The was three for in Tris-buffered saline and for in containing The was three for in and for in a of in This was by three in The was a of of and of in of The used as a was as previously C. Lam J.S. Mol. Microbiol. PubMed Scopus Google Scholar). of wbpO gene was into the with an wbpO was from aeruginosa O6) by the which an which a The of of of of and was the of of by of and step of was the and the were with and and The was by and and into the strain E. for and of the of WbpO, of containing was with of an and was to a of and was to for were by for and were WbpP and were as previously C. M. Lam J.S. 2000; PubMed Scopus Google Scholar, C. S. J.R. Lam J.S. PubMed Scopus Google Scholar). from P. aeruginosa (serotype to be a C. S. J.R. Lam J.S. PubMed Scopus Google Scholar). of WbpO or the from of was in of The were by and and were by for the was as by the with as the and containing as the the of in WbpO or WbpP was to a to the by the The of WbpO and WbpP was by and was the of to a of was as previously C. S. J.R. Lam J.S. PubMed Scopus Google Scholar). or of of WbpO, of were for were in two of the first the reaction was for by the of the and for the and of WbpO, or and of WbpO and were for of the was used as the for and and and was the for and for the of the were or in both and conversion was by The of WbpO were as previously for C. S. J.R. Lam J.S. PubMed Scopus Google Scholar), of WbpO and of the as and of and of was into in a reaction containing of and of of by from was by an a of The containing were and of was into in the as for that of WbpO was used of of the WbpO by reaction of the WbpO reaction were in of and into and single correlation correlation and from were used for and with a were used for and of J.R. G. of Scholar, J.R. in and Scholar). were with a with a for were with of the The of was used as an and of of and in a of of kinetic parameters for as the and of that from to of kinetic parameters for as a and of that from to of kinetic parameters for as an and of that from to of kinetic parameters for and of that from to were and the of the was by in a the of was from and The were the of from The of the to the and by were compared the of the and the of the could be to be a a a a of the the were to the by and the results are the of three of of E. C. PubMed Scopus Google or of E. C. R. M. 2002; PubMed Scopus Google Scholar), of with and for or for was with of an were for of of of with was with of an was by the of and was for of of with and was with of an was by the of and WbpP was for In cases, were into and by To from a were in of and by was by for by of the for of from the step was a in from the strain the were in the as of and in the of of and in were These were in the of WbpP the of and of and in the of WbpP were by the and P. J.S. M. PubMed Scopus Google Scholar). of from wild-type P. aeruginosa PAK and the mutant strains were and by and Flagellin from wild-type as as the and showed an mass of flagellin from the wbpO mutant was Flagellin isolated from the PAK wbpO mutant complemented with the plasmid which the wild-type wbpO an mass to that of the flagellin from the wild-type of the PAK with PAK flagellin showed with the flagellin proteins isolated from the and mutant strains, as as the smaller flagellin isolated from the wbpO mutant to the flagellin isolated from the wbpO mutant exhibited an altered isoelectric point when compared with isolated from either the wbpL, or acid the of the from PAK is to be but this not that may be by the glycan In flagellin from the wild type, wbpL, and mutants as from a of to with a Flagellin from the wbpO mutant to but with the focusing that is than the flagellin from the Flagellin from the complemented exhibited an to that of the wild-type flagellin. of flagellin by showed a reaction to the in and to of Flagellin glycosylation of the flagellin proteins was and the results showed a reaction with the isolated from strains for the wbpO mutant and the were in the as and of of flagellin from strain the wbpO and the wbpO mutant showed a of and Because the mass of flagellin acid is to be these that the wild-type PAK flagellin a covalently moiety to a total mass of In flagellin from the wbpO mutant an mass of that for a mass with the of two Flagellin from the wbpO mutant wbpO a plasmid showed a mass that wbpO a plasmid could the in flagellin not to the glycosylation as the and of WbpO, and was with an the with WbpO of total from an (serotype and WbpP were to the with and of and WbpP to that previously C. M. Lam J.S. 2000; PubMed Scopus Google Scholar, C. S. J.R. Lam J.S. PubMed Scopus Google Scholar). WbpO, of as from the could be from of The mass of WbpO kDa) as by with the mass of The of the was to be of WbpO and WbpP of containing and WbpO two reaction that and The that was as by with an and the as in was as by in containing and two and were the was as and the as in was as by Because is not serotype was used to this sugar-nucleotide for WbpP C. S. J.R. Lam J.S. PubMed Scopus Google Scholar). was used to into prior to the of WbpP. of these showed that WbpP was capable of converting into a that as in This was as by to is not and because is not capable of WbpO was used to for WbpP WbpO was used to into prior to the of WbpP. of that WbpP is capable of converting into a that as in This was as by with of the WbpO and by and and correlation and to in the C. S. J.R. Lam J.S. PubMed Scopus Google Scholar), was to be The for that for with from the moiety of the of and and the of and and for a the and the a single correlation spectroscopy the of was to be which the of an and a correlation spectroscopy showed the of to be which a these was to be of and of is a of and a of not WbpO is a of with between and not WbpO requires either or for The are when the is and when the is with a correlation between and the PubMed Scopus Google Scholar). that and were not because was when these were previously C. S. J.R. Lam J.S. PubMed Scopus Google Scholar), and were not the of In the the of WbpO was in a of the of WbpO with the conversion with an in to a of which conversion with of of of WbpO was by the of kinetic to the and the of from The of in the and were compared and the of the and the of between the two could be with to the three the was for kinetic because parameters than the a for than for of the for and with as the a for as the a for than for a for as the parameters of total reaction was The total reaction was in a from the of WbpP was by a was proposed for the of three of C. Lam J.S. PubMed Scopus Google Scholar). catalyze the conversion between and not show a for either or and between and Because and were not the of that from one of were for the of these to utilize and as from and WbpP strains were with and or from with from or with from reaction was was and the of was by in to from strains the both WbpP and were to either or as the of the to from previously for the acid was to catalyze the of to the but was when was used as a substrate. In not show when either of the two were used as a a of for this from assays for three of and of and for reaction is for as compared with a from a of and for reaction is for as compared with a from a in a of and in of in the of WbpP that could be in the of WbpP and could with the of WbpP in the of was the for between the moiety and the In is the for because the of and of the are as the of the moiety in P. aeruginosa PAK produces a single flagellum containing a-type which to be two with M. A. E. P. R. PubMed Scopus Google Scholar). the that O-antigen in the synthesis of these by flagellin proteins isolated from wild-type PAK and three mutant strains in O-antigen from and mutants a as the wild-type (45 kDa), and both were In flagellin from the wbpO mutant an mass of that is a in flagellin of the wbpO mutant with the plasmid containing wbpO in flagellin with of a than that of either PAK or the other mutant The mass of the glycosylated wild-type flagellin in by was to be and this mass is with that by M. A. E. P. R. PubMed Scopus Google Scholar), results that PAK flagellin was two with a of including one to showed that the wbpO mutant flagellin a mass of which to the mass of flagellin with two with a mass of M. A. E. P. R. PubMed Scopus Google proposed a for the glycan that are covalently to PAK flagellin. This that is a of two to attached to the flagellin by a the of the flagellin were the of not results that the of the WbpO is required for the synthesis of the of the flagellin The that flagellin from the wbpO mutant a with the attachment of two that the by the WbpO reaction is in this In the flagellin glycan by M. A. E. P. R. PubMed Scopus Google Scholar), the is proposed to monosaccharides of including be into the flagellin or requires by other encoded by the flagellin glycosylation into the is not because the of the flagellin is not is not to a pathway for The that in and not flagellin glycosylation that a O-antigen unit is not required flagellin glycosylation This is the that by M. A. E. P. R. PubMed Scopus Google could not the of O-antigen in the flagellin This is in to pilin glycosylation in P. aeruginosa strain an O-antigen unit is to pilin (6DiGiandomenico A. Matewish M.J. Bisaillon A. Stehle J.R. Lam J.S. Castric P. Mol. Microbiol. 2002; 46: PubMed Scopus Google Scholar). that P. aeruginosa to flagellin and is of to that in the in from with showed a of B-band polysaccharide Lam J.S. Microbiol. PubMed Google Scholar, PubMed Google Scholar), the to a of and a the is pili or E. PubMed Google Scholar). In WbpP was to C. M. Lam J.S. 2000; PubMed Scopus Google Scholar), and WbpO was to be a that possessed of C. M. E. Lam J.S. 2000; PubMed Scopus Google Scholar). this WbpP was proposed to first in the conversion of into the acid that flagellin glycosylation requires WbpO, but not prompted us to the of of WbpO and WbpP in WbpO was previously with a and to be for with WbpO from serotype as an that is the for WbpO was with an was the for and of of WbpO for kinetic either or as a substrate. To whether WbpP could or as a WbpP was the of or The of the reaction was previously by as C. S. J.R. Lam J.S. PubMed Scopus Google Scholar), and the of and were in this as and by In this the results showed that WbpP could and a C4 in to previously as a C. M. Lam J.S. 2000; PubMed Scopus Google Scholar). with the results of these results showed that in the conversion of into WbpO, and WbpP are capable of in either Because not and could be in as an this us to a kinetic and of This that are a number of between the of (serotype O6) and that of (serotype catalyze a both M. R. S. S. A. PubMed Google Scholar, C. C. PubMed Scopus Google and and both or for Because was in containing or C. S. J.R. Lam J.S. PubMed Scopus Google Scholar), this that the with or not from of or WbpO because of the of these but that and are of and was that the of the or to a in the of and The of these with the PubMed Scopus Google Scholar), the reaction is by of and by of as for C. S. J.R. Lam J.S. PubMed Scopus Google and WbpO of the effect of the of WbpO both a and a effect The effect was the because an in the of with an in of however, the conversion of to with an in an This effect is with the with The may containing than the with as the in the of the of WbpO and of with as a that WbpO a and than that WbpO is more than In WbpO is capable of as a could The from the kinetic of WbpO to WbpO a for and These results were to previously for C. M. E. Lam J.S. 2000; PubMed Scopus Google Scholar). This is to the to the WbpO in the these results showed that is more than either or a for of with to of In kinetic for WbpO not to the than to the and WbpO not This may be to the that the first step in the pathway for the biosynthesis of WbpO may catalyze either the first step or the step of the pathway for Results from the of WbpP showed that of was into the of the reaction was C. M. Lam J.S. 2000; PubMed Scopus Google Scholar). In this of the reaction acid WbpP catalyzed conversion of into the of the reaction was of the for and that the conversion of to a than the a of the of WbpP for with the of WbpO for the that WbpO is capable of of than WbpP. these that WbpO and WbpP in either is a for WbpO to first to this acid is by WbpP to is a between the which of production of and acid the of the of to the was Because the and of WbpP C. Lam J.S. PubMed Scopus Google Scholar), and that of C4 epimerase proteins acid as a were to WbpP and into the The results of this showed that WbpP these acid in a to acid The results showed that could an in the of WbpP compared with In is for between the of and C4 of the of Because the moiety of the is in the of WbpP C. Lam J.S. PubMed Scopus Google Scholar), an could to and the moiety of for This may be to a conversion of into than the the of WbpP was a was proposed for the of three of C. Lam J.S. PubMed Scopus Google Scholar). catalyze the conversion between and not show a for either or and between and In this assays were to determine which of the three of proteins are capable of C4 and as a epimerase from K. E. C. PubMed Scopus Google Scholar), was not capable of either or but this is not because could not utilize as C. M. Lam J.S. 2000; PubMed Scopus Google Scholar, C. Lam J.S. PubMed Scopus Google Scholar). a epimerase from E. C. R. M. 2002; PubMed Scopus Google Scholar), was to a conversion conversion to was could not catalyze the This is because is capable of and as as and WbpP and were to utilize either or as as for the production of the production of the when the are in the acid Thus, the of the acid to a effect the of the C4 in for and altered for In that flagellin glycosylation in P. aeruginosa PAK the O-antigen gene wbpO but not or kinetic to show WbpO and WbpP in either in the biosynthesis of is a for WbpO to into first the conversion into by WbpP In are a number of in the of serotype and WbpO serotype O6) as but is a in in that could not as a substrate. these differences and for metabolic the for is and are to determine the of and for with the and of are to and for the and and to for the of with

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