Shield as Signal Lipopolysaccharides and the Evolution of Immunity to Gram-Negative Bacteria 英文参考文献.docVIP

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Shield as Signal Lipopolysaccharides and the Evolution of Immunity to Gram-Negative Bacteria 英文参考文献.doc

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Shield as Signal Lipopolysaccharides and the Evolution of Immunity to Gram-Negative Bacteria 英文参考文献

Opinions ShieldasSignal:Lipopolysaccharides andtheEvolutionofImmunity toGram-NegativeBacteria Robert S.Munford*, Alan W. Varley A ccordingtotheinnateimmunityconcept[1],animals defendthemselvesfrommicrobesbyrecognizing pathogen-associatedmolecularpatterns.Todetect manyGram-negativebacteria,animalsusetheCD14–MD-2– TLR4receptormechanismtorecognizethelipidAmoietyof thecellwalllipopolysaccharide(LPS).LipidAisa antimicrobialpeptides(CAMPs)[3].AsnotedbyMiller[4] andothers,increasedresistancetoCAMPsandotherhost moleculesmayexplainwhyGram-negativebacteriathat colonizemucosaeusuallymakeLPSwithsixormoreacyl chains(Figure1).Althoughwefoundnodemonstrationthat thislipidAstructureenablescommensalbacteriatothriveon mucosalsurfaces,theevidencethatitdoessoforcolonizers andpathogensisextensive.Havinghexaacyl(ratherthan pentaacyl)lipidAenablesBordetellaandHaemophilusspeciesto persistintherespiratorytract[5–7]andNeisseriagonorrhoeae tosurvivewithinepithelialcells[8].Pseudomonasaeruginosa livesinwater,producesapredominantlypentaacylatedLPS, anddoesnotcolonizethemucosaeofnormalhumans.When P.aeruginosacolonizetheairwaysofchildrenwithcystic ?brosis,however,thebacteriaoftenadaptbyproducing hexaacylated(andevenheptaacylated)lipidA[9].Mucosal lipidAisalsofoundinintestinalpathogens:Shigellaand Salmonella,pathogenicEscherichiacoli[10],Aeromonasspecies, Plesiomonasshigelloides,andVibriocholeraeO1.InSalmonellaand someothers,aPhoP/PhoQ-regulatedtranscriptionalprogram promoteslipidApalmitoylation(heptaacylation)alongwith otherchangesthatincreaseresistancetoCAMPs[4,11–13]. OthermucosalbacteriamayalsoproducelipidAthatismore hydrophobicthanmucosallipidA,withlongersecondary chains(Campylobacterjejuni)ormoreofthem:heptaacyl (Moraxella)oroctaacyl(V.choleraeO139).Thosethatproduce lesshydrophobiclipidAseemtobespecialcases:the pentaacylLPSofChlamydiaspeciesisfoundinspore-like elementarybodies,andHelicobacterpylori,withtetraacylLPS,is adaptedtoliveinthestomach. glucosaminedisaccharidethatcarriesphosphatesat