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Lipopolysaccharides of bacterial pathogens from the genus Yersinia: a mini-review
Biochimie 85 (2003) 145–152 www.elsevier.com/locate/biochi
Review
Lipopolysaccharides of bacterial pathogens from the genus Yersinia: a mini-review Maud Bruneteau a,*, Samuel Minka b a
Laboratoire d’écologie microbienne du sol, UMR-CNRS 5557, université Claude-Bernard, Lyon 1, bâtiment 741, 43, boulevard du 11 novembre 1918, 69622 Villeurbanne cedex, France b Faculté des sciences, département de biochimie, BP 812, Yaoundé, Cameroun Received 22 October 2002; accepted 6 January 2003
Abstract This review summarizes the state of knowledge on the composition and structure of the lipopolysaccharides (LPS) from three species of Yersinia known to produce disease in humans: Y. pseudotuberculosis, Y. enterocolitica and Y. pestis. We also mention recent data on the genome sequence of Yersinia pestis and the role of LPS in relation to the virulence of this bacteria. © 2003 Éditions scientifiques et médicales Elsevier SAS and Société française de biochimie et biologie moléculaire. All rights reserved. Keywords: Lipopolysaccharides; Chemical structure; Virulence; Yersinia
1. Introduction The genus Yersinia belongs to the family Enterobacteriaceae. It consists so far of eleven species, three of which are human pathogens: Y. pestis, Y. pseudotuberculosis and Y. enterocolitica [1]. Yersinia pestis is the causative agent of bubonic plague, whereas the latter two species cause mild gastroenteritis [2,3]. Being gram-negative bacteria, these organisms have lipopolysaccharides (LPS) as major components of their outer membrane. The full LPS molecule (Sform) includes three structurally distinct regions: the lipid A anchored in the outer membrane, an oligosaccharide core and the O-antigenic polysaccharide [4–7]. Most studied so far are the antigens of Y. pseudotuberculosis and Y. enterocolitica in which the chemical composition and the structure of the
Abbreviations: Abe, abequose (3,6-dideoxy-D-xylo-hexose); Alt, altrose; 6dAlt, 6-deoxyaltrose; Asc, ascarylose (3,6-dideoxy-L-arabinohexose); Col, colitose (3,6-dideoxy-L-xylo-hexose); Fo, formyl; 6dGul, 6-deoxygulose; D,D-Hep, D-glycero-D-manno-heptose; L,D-Hep, L-glycero-D-manno-heptose; Kdo, 3-deoxy-a-D-manno-oct-2-ulosonic acid; D-man-6dHep, 6-deoxy-D-manno-heptose; P, phosphate; Par, paratose (3,6-dideoxy-D-ribo-hexose); Rha4N, 4-amino-4-deoxyrhamnose; Tyv, tyvelose (3,6-dideoxy-D-arabino-hexose); D-Xlu, xylulose (D-threopentulose). * Corresponding author. E-mail address: [email protected] (M. Bruneteau).
oligosaccharide repeating units are very diverse [8]. The majority of them are characterized by a linear sugar backbone with single sugar residues as branches. Yersinia pestis, the bubonic plague bacterium, is an R-form which has a genetic defect in the biosynthesis of complete LPS and it lacks O-specific chains [9,10]. Today the chemical composition of the inner and outer core regions of the LPS of several strains from the three species of Yersinia mentioned above is known [6] but so far only little information has been available of their detailed structure. Actually, detailed knowledge on the structure of the core-lipid A region of Yersinia LPS has been essentially deduced from more recent studies on serotypes O: 3, O: 8 and O: 9 of Y. enterocolitica and of Y. pestis, strain EV. In earlier work, the data concerning the structural features of the core of the LPS from various serovars of Y. pseudotuberculosis have been reported by Ovodov et al. [11]. More detailed structural work has been carried out on the inner core region of Y. enterocolitica O: 3 LPS [12]. More recently, the chemical structure of the core region for serotypes O: 8, O: 9 of Y. enterocolitica were elucidated [13,14]. The core structure of the lipopolysaccharide from Y. pestis has also been recently reported [15]. Recent structural analyses have been reported on the lipid A [16]. The structure of the Yersinia lipid A has been shown to be identical to those of other enterobacteria [13].
© 2003 Éditions scientifiques et médicales Elsevier SAS and Société française de biochimie et biologie moléculaire. All rights reserved. DOI: 1 0 . 1 0 1 6 / S 0 3 0 0 - 9 0 8 4 ( 0 3 ) 0 0 0 0 5 - 1
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Table 1 Structures of O-antigens of Y. pseudotuberculosis (Knirel and Kochetkov [8])
This mini-review describes the state of knowledge on the composition and chemical structure of the LPS from three species of Yersinia known to produce human disease. 2. Chemical composition and structural analysis of O-antigens For many serotypes of Y. pseudotuberculosis and Y. enterocolitica, the chemical structure of the O-antigens is known and has been described in earlier reviews [7,8].The O-specific chains determine the O-antigenic specificity of the bacteria. Strains of Y. pseudotuberculosis are divided into seven serogroups. Several studies were performed on the O-specific polysaccharides of LPS from groups I to VII. Their chemical composition was shown to be highly diverse. All polysaccharides of Y. pseudotuberculosis are branched
hexosaminoglycans characterized by the presence of hexoses, 6-deoxy-hexoses, N-acetyl-2-amino-2-deoxy-hexoses and 3,6-dideoxy-hexoses [8]. Their structures, as described by Knirel and Kochetkov [8], are shown in tables 1 and 2 . These authors observed that the main difference between the polysaccharides of the seven serogroups is the nature of the terminal group identified as 3,6-dideoxy-hexoses (in serotypes IB, IIC, III, IVA, VA, VII [18–22,26]), the 6-deoxy-Laltrofuranose (in serotype VB [23]), the 6-deoxy-D-mannoheptose (in serotype IA [17]) and yersiniose A (4C-[(R)-1hydroxyethl]-3,6-dideoxy-D-xylo-hexose) (in serotype VI [24,25]). Yersinia enterocolitica has also been divided into different serotypes depending on their O-somatic antigens [27]. The O-specific Y. enterocolitica polysaccharides are either homopolysaccharides or branched heteropolysaccharides. The
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Table 2 Structures of O-antigens of Y. pseudotuberculosis and Y. enterocolitica (Knirel and Kochetkov [8])
O-antigens of serogroups: O: 1,2a,3 and O: 2,3 contain 6-deoxy-L-altrofuranose as the sole sugar constituent [28,29]. The O-antigen of serogroup O: 9 as described by Caroff et al. [30], is also a homopolymer of 1,2 linked 4,6-dideoxy-4-formamido-D-mannopyranosyl residues. O-Acyl groups are present in the antigens of serogroup O: 2a,2b,3 [28] and O: 4,32 [31]. Most of the O-specific Y. enterocolitica heteroglycans are branched; only the O-antigen of serogroup O: 6,31 is a linear polysaccharide composed of galactosyl and 6-deoxy-D-gulosyl residues linked through O-3 [32]. The O-antigen of serogroup O: 8 is a heteropolymer of branched pentasaccharide repeating units of 2-amino-2-deoxy-D-glucose, galactose, fucose, mannose
and 6d-gulose [33]. Recently Perry and MacLean [34] have reported the structure of the O-antigen polysaccharide component of the lipopolysaccharide produced by Y. enterocolitica serotype O: 28. This O-antigen was found to be a polymer of repeating branched hexasaccharide units composed of L-rhamnose, 2-amino-2-deoxy-D-glucose and 2-amino-2-deoxy-D-galacturonic acid. The antigens O: 5,5,27 [35] and O: 5,27 [36] are neutral branched polysaccharides characterized respectively by the presence of yersiniose B (4C-[(S)-1-hydroxyethyl]-3,6-dideoxy-D-xylohexose) and of D-threo-pentulose (xylulose). The structure of the respective units described by Knirel and Kochetkov [8], are shown in Table 2 and Table 3.
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Table 3 Structures of O-antigens of Y. enterocolitica
3. Chemical composition and structural analysis of the core The same chemical composition was observed for the core regions of Y. pseudotuberculosis, Y. enterocolitica and Y. pestis. They contain glucose, galactose, 2-amino-2-deoxy-Dglucose, 3-deoxy-D-manno-octulonic acid, L-glycero-Dmanno- and D-glycero-D-manno-heptose [10,12,24,37,38]. Until recently, not much information was available on the structure of the core region of Yersinia LPS. Only a partial structure of the core oligosaccharide isolated from LPS of an R-form of Y. pseudotuberculosis has been reported by Tomshich et al. [39]. However, extensive investigations carried out in recent years on the chemical structure of LPS isolated from Y. enterocolitica have now led to a detailed knowledge of the yersinial core. In previous work the structure of the complete inner core region of serotype O: 3 (Y75) has been reported. Radziejewska-Lebrecht et al. [12] isolated a heptasaccharide composed of D-glucose, L,D-heptose, D,D-heptose and 3-deoxy-D-manno-octulosonic acid which structure is shown in Table 4 . Structural determination of the core region of LPS of serotypes O: 8 and O: 9 have also been elucidated. The carbohydrate backbone of the major portion of the core of these LPS shown in Table 4 corresponds to a branched octosaccharide. Apparently the Y. enterocolitica core structures are basically conserved among the different serotypes so far ana-
lyzed. The sole difference mentioned by Oertelt et al. [14], between the core region of Y. enterocolitica O: 3 and O: 9 LPS is the substitution of the terminal D,D Hep residue by a phosphate at position O7 in the LPS of the rough O: 3. In the core region of Y. enterocolitica O: 8 the substitution of the L,D-heptose (II) unit by a 2-amino-2-deoxy-D-glucose unit is also the sole difference between the core regions of Y. enterocolitica O: 8 and O: 9 LPS. The partial structure of the LPS core of Y. pestis analyzed recently by Vinogradov et al. [15], shows a backbone partially differing from that of the LPS core of Y. enterocolitica as shown in Table 4. Like in the Enterobacteriaceae family, the internal Hep-> Hep-> Hep-> Kdo region is also characteristic of the core type of Yersiniae. 4. Chemical composition and structural analysis of lipid A In the past lipids A of Yersinia LPS have been described essentially in terms of fatty acid composition [10,40–42]. Several investigations on the partial structure of lipid A of LPS from Y. pestis have been published [42]. More recently, Aussel et al. [16], determined and compared the lipid A structure of the endotoxins of Y. enterocolitica and Y. pestis. These authors reported that the structure of the Yersinia lipid A skeleton is identical to that of other Enterobacteria. The
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Table 4 Structures of the core of Y. enterocolitica and Y. pestis
lipid A is a tetraacylated and bisphosphorylated glucosamine disaccharide [43]. The position of fatty acid residues as shown in Fig. 1 is in accordance with recent reports of Oertelt et al. [14]. The hydroxyl groups of the disaccharide are acylated with 3-hydroxytetradecanoic and 3-dodecanoyloxytetradecanoic acids. The reducing glucosamine residue is amidified with 3-hydroxytetradecanoic acid. The nonreducing glucosamine residue is amidified with 3-dodecanoyloxytetradecanoic acid in the lipid A of Y. enterocolitica and with 3-hexadecenoyltetradecanoic acid in the lipid A of Y. pestis. It appears that the lipid A structures differed essentially in the secondary acylation of the amide-linked 3-hydroxytetradecanoic acid in the non reducing glucosaminyl residue. Comparison of the Yersinia lipid A structures shows that they do not differ significantly in their general architecture from those of Enterobacteria [13,16].
5. Discussion Chemical composition and structural analysis of Yersinia LPS reported in this review show common structural features mainly in the core-lipid A regions. LPS is an important factor in the virulence of Yersinia but until now the role of LPS in the virulence of Y. pestis was not clear. Achtman et al. [44] showed that Y. pestis and Y. pseudotuberculosis are genetically very closely related. Recently Skurnik et al. [45] observed that Y. pestis has evolved from Y. pseudotuberculosis. These authors provide evidence that one step in the evolution of Y. pestis from Y. pseudotuberculosis O: 1b was the inactivation of the O-antigen gene cluster. It seems that the expression of this antigen is not beneficial for the virulence of the agent of bubonic plague. However the O-side chain of LPS is an important factor in the virulence of a range of pathogens including Y. enterocolitica
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Fig. 1. Schematic representation of the major lipid A component of (a) Y. enterocolitica O: 11,23 and O: 11,24; Y. ruckeri O: 1 and O: 2 (b) Y. enterocolitica O: 3 and O: 9. (c) Y. pestis. (d) E. coli. The variable part of the structures is presented in bold. The configuration of the double bond of the C16:1 was not determined (Aussel et al. [16]).
[46]. The genome sequence of Y. pestis reported recently by Parkhill et al. [47], revealed genes which encode unknown surface antigens having a role in the virulence of this bacterium.
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V. Zähringer, B. Lindner, E. Th. Rietschel, Chemical structure of lipid A: Recent advances in structural analysis of biological active molecules, in: H. Brade, S.M. Opal, S.N. Vogel, D.C. Morrison (Eds.), Endotoxin in Health and Disease, Marcel Dekker, New-York USA, 1999, pp. 93–113.
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P.E. Jansson, The Chemistry of O-polysaccharide chains in bacterial lipopolysaccharides: Recent advances in structural analysis of biological active molecules, in: H. Brade, S.M. Opal, S.N. Vogel, D.C. Morrison (Eds.), Endotoxin in Health and Disease, Marcel Dekker, New York USA, 1999, pp. 155–178.
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A.Y. Knirel, N.K. Kochetkov, The structure of lipopolysaccharides of Gram-Negative bacteria. III. The structure of O-antigens: A Review, Biochemistry (Moscow) 59 (1994) 1325–1383.
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Acknowledgements We thank Dr H. Mayer for carefully reading this review. We also thank Drs J. Radziejewska-Lebrecht, M.TH. Pommier, Y. Knirel and G. Pellon for help in searching the literature.
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Review
Lipopolysaccharides of bacterial pathogens from the genus Yersinia: a mini-review Maud Bruneteau a,*, Samuel Minka b a
Laboratoire d’écologie microbienne du sol, UMR-CNRS 5557, université Claude-Bernard, Lyon 1, bâtiment 741, 43, boulevard du 11 novembre 1918, 69622 Villeurbanne cedex, France b Faculté des sciences, département de biochimie, BP 812, Yaoundé, Cameroun Received 22 October 2002; accepted 6 January 2003
Abstract This review summarizes the state of knowledge on the composition and structure of the lipopolysaccharides (LPS) from three species of Yersinia known to produce disease in humans: Y. pseudotuberculosis, Y. enterocolitica and Y. pestis. We also mention recent data on the genome sequence of Yersinia pestis and the role of LPS in relation to the virulence of this bacteria. © 2003 Éditions scientifiques et médicales Elsevier SAS and Société française de biochimie et biologie moléculaire. All rights reserved. Keywords: Lipopolysaccharides; Chemical structure; Virulence; Yersinia
1. Introduction The genus Yersinia belongs to the family Enterobacteriaceae. It consists so far of eleven species, three of which are human pathogens: Y. pestis, Y. pseudotuberculosis and Y. enterocolitica [1]. Yersinia pestis is the causative agent of bubonic plague, whereas the latter two species cause mild gastroenteritis [2,3]. Being gram-negative bacteria, these organisms have lipopolysaccharides (LPS) as major components of their outer membrane. The full LPS molecule (Sform) includes three structurally distinct regions: the lipid A anchored in the outer membrane, an oligosaccharide core and the O-antigenic polysaccharide [4–7]. Most studied so far are the antigens of Y. pseudotuberculosis and Y. enterocolitica in which the chemical composition and the structure of the
Abbreviations: Abe, abequose (3,6-dideoxy-D-xylo-hexose); Alt, altrose; 6dAlt, 6-deoxyaltrose; Asc, ascarylose (3,6-dideoxy-L-arabinohexose); Col, colitose (3,6-dideoxy-L-xylo-hexose); Fo, formyl; 6dGul, 6-deoxygulose; D,D-Hep, D-glycero-D-manno-heptose; L,D-Hep, L-glycero-D-manno-heptose; Kdo, 3-deoxy-a-D-manno-oct-2-ulosonic acid; D-man-6dHep, 6-deoxy-D-manno-heptose; P, phosphate; Par, paratose (3,6-dideoxy-D-ribo-hexose); Rha4N, 4-amino-4-deoxyrhamnose; Tyv, tyvelose (3,6-dideoxy-D-arabino-hexose); D-Xlu, xylulose (D-threopentulose). * Corresponding author. E-mail address: [email protected] (M. Bruneteau).
oligosaccharide repeating units are very diverse [8]. The majority of them are characterized by a linear sugar backbone with single sugar residues as branches. Yersinia pestis, the bubonic plague bacterium, is an R-form which has a genetic defect in the biosynthesis of complete LPS and it lacks O-specific chains [9,10]. Today the chemical composition of the inner and outer core regions of the LPS of several strains from the three species of Yersinia mentioned above is known [6] but so far only little information has been available of their detailed structure. Actually, detailed knowledge on the structure of the core-lipid A region of Yersinia LPS has been essentially deduced from more recent studies on serotypes O: 3, O: 8 and O: 9 of Y. enterocolitica and of Y. pestis, strain EV. In earlier work, the data concerning the structural features of the core of the LPS from various serovars of Y. pseudotuberculosis have been reported by Ovodov et al. [11]. More detailed structural work has been carried out on the inner core region of Y. enterocolitica O: 3 LPS [12]. More recently, the chemical structure of the core region for serotypes O: 8, O: 9 of Y. enterocolitica were elucidated [13,14]. The core structure of the lipopolysaccharide from Y. pestis has also been recently reported [15]. Recent structural analyses have been reported on the lipid A [16]. The structure of the Yersinia lipid A has been shown to be identical to those of other enterobacteria [13].
© 2003 Éditions scientifiques et médicales Elsevier SAS and Société française de biochimie et biologie moléculaire. All rights reserved. DOI: 1 0 . 1 0 1 6 / S 0 3 0 0 - 9 0 8 4 ( 0 3 ) 0 0 0 0 5 - 1
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Table 1 Structures of O-antigens of Y. pseudotuberculosis (Knirel and Kochetkov [8])
This mini-review describes the state of knowledge on the composition and chemical structure of the LPS from three species of Yersinia known to produce human disease. 2. Chemical composition and structural analysis of O-antigens For many serotypes of Y. pseudotuberculosis and Y. enterocolitica, the chemical structure of the O-antigens is known and has been described in earlier reviews [7,8].The O-specific chains determine the O-antigenic specificity of the bacteria. Strains of Y. pseudotuberculosis are divided into seven serogroups. Several studies were performed on the O-specific polysaccharides of LPS from groups I to VII. Their chemical composition was shown to be highly diverse. All polysaccharides of Y. pseudotuberculosis are branched
hexosaminoglycans characterized by the presence of hexoses, 6-deoxy-hexoses, N-acetyl-2-amino-2-deoxy-hexoses and 3,6-dideoxy-hexoses [8]. Their structures, as described by Knirel and Kochetkov [8], are shown in tables 1 and 2 . These authors observed that the main difference between the polysaccharides of the seven serogroups is the nature of the terminal group identified as 3,6-dideoxy-hexoses (in serotypes IB, IIC, III, IVA, VA, VII [18–22,26]), the 6-deoxy-Laltrofuranose (in serotype VB [23]), the 6-deoxy-D-mannoheptose (in serotype IA [17]) and yersiniose A (4C-[(R)-1hydroxyethl]-3,6-dideoxy-D-xylo-hexose) (in serotype VI [24,25]). Yersinia enterocolitica has also been divided into different serotypes depending on their O-somatic antigens [27]. The O-specific Y. enterocolitica polysaccharides are either homopolysaccharides or branched heteropolysaccharides. The
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Table 2 Structures of O-antigens of Y. pseudotuberculosis and Y. enterocolitica (Knirel and Kochetkov [8])
O-antigens of serogroups: O: 1,2a,3 and O: 2,3 contain 6-deoxy-L-altrofuranose as the sole sugar constituent [28,29]. The O-antigen of serogroup O: 9 as described by Caroff et al. [30], is also a homopolymer of 1,2 linked 4,6-dideoxy-4-formamido-D-mannopyranosyl residues. O-Acyl groups are present in the antigens of serogroup O: 2a,2b,3 [28] and O: 4,32 [31]. Most of the O-specific Y. enterocolitica heteroglycans are branched; only the O-antigen of serogroup O: 6,31 is a linear polysaccharide composed of galactosyl and 6-deoxy-D-gulosyl residues linked through O-3 [32]. The O-antigen of serogroup O: 8 is a heteropolymer of branched pentasaccharide repeating units of 2-amino-2-deoxy-D-glucose, galactose, fucose, mannose
and 6d-gulose [33]. Recently Perry and MacLean [34] have reported the structure of the O-antigen polysaccharide component of the lipopolysaccharide produced by Y. enterocolitica serotype O: 28. This O-antigen was found to be a polymer of repeating branched hexasaccharide units composed of L-rhamnose, 2-amino-2-deoxy-D-glucose and 2-amino-2-deoxy-D-galacturonic acid. The antigens O: 5,5,27 [35] and O: 5,27 [36] are neutral branched polysaccharides characterized respectively by the presence of yersiniose B (4C-[(S)-1-hydroxyethyl]-3,6-dideoxy-D-xylohexose) and of D-threo-pentulose (xylulose). The structure of the respective units described by Knirel and Kochetkov [8], are shown in Table 2 and Table 3.
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Table 3 Structures of O-antigens of Y. enterocolitica
3. Chemical composition and structural analysis of the core The same chemical composition was observed for the core regions of Y. pseudotuberculosis, Y. enterocolitica and Y. pestis. They contain glucose, galactose, 2-amino-2-deoxy-Dglucose, 3-deoxy-D-manno-octulonic acid, L-glycero-Dmanno- and D-glycero-D-manno-heptose [10,12,24,37,38]. Until recently, not much information was available on the structure of the core region of Yersinia LPS. Only a partial structure of the core oligosaccharide isolated from LPS of an R-form of Y. pseudotuberculosis has been reported by Tomshich et al. [39]. However, extensive investigations carried out in recent years on the chemical structure of LPS isolated from Y. enterocolitica have now led to a detailed knowledge of the yersinial core. In previous work the structure of the complete inner core region of serotype O: 3 (Y75) has been reported. Radziejewska-Lebrecht et al. [12] isolated a heptasaccharide composed of D-glucose, L,D-heptose, D,D-heptose and 3-deoxy-D-manno-octulosonic acid which structure is shown in Table 4 . Structural determination of the core region of LPS of serotypes O: 8 and O: 9 have also been elucidated. The carbohydrate backbone of the major portion of the core of these LPS shown in Table 4 corresponds to a branched octosaccharide. Apparently the Y. enterocolitica core structures are basically conserved among the different serotypes so far ana-
lyzed. The sole difference mentioned by Oertelt et al. [14], between the core region of Y. enterocolitica O: 3 and O: 9 LPS is the substitution of the terminal D,D Hep residue by a phosphate at position O7 in the LPS of the rough O: 3. In the core region of Y. enterocolitica O: 8 the substitution of the L,D-heptose (II) unit by a 2-amino-2-deoxy-D-glucose unit is also the sole difference between the core regions of Y. enterocolitica O: 8 and O: 9 LPS. The partial structure of the LPS core of Y. pestis analyzed recently by Vinogradov et al. [15], shows a backbone partially differing from that of the LPS core of Y. enterocolitica as shown in Table 4. Like in the Enterobacteriaceae family, the internal Hep-> Hep-> Hep-> Kdo region is also characteristic of the core type of Yersiniae. 4. Chemical composition and structural analysis of lipid A In the past lipids A of Yersinia LPS have been described essentially in terms of fatty acid composition [10,40–42]. Several investigations on the partial structure of lipid A of LPS from Y. pestis have been published [42]. More recently, Aussel et al. [16], determined and compared the lipid A structure of the endotoxins of Y. enterocolitica and Y. pestis. These authors reported that the structure of the Yersinia lipid A skeleton is identical to that of other Enterobacteria. The
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Table 4 Structures of the core of Y. enterocolitica and Y. pestis
lipid A is a tetraacylated and bisphosphorylated glucosamine disaccharide [43]. The position of fatty acid residues as shown in Fig. 1 is in accordance with recent reports of Oertelt et al. [14]. The hydroxyl groups of the disaccharide are acylated with 3-hydroxytetradecanoic and 3-dodecanoyloxytetradecanoic acids. The reducing glucosamine residue is amidified with 3-hydroxytetradecanoic acid. The nonreducing glucosamine residue is amidified with 3-dodecanoyloxytetradecanoic acid in the lipid A of Y. enterocolitica and with 3-hexadecenoyltetradecanoic acid in the lipid A of Y. pestis. It appears that the lipid A structures differed essentially in the secondary acylation of the amide-linked 3-hydroxytetradecanoic acid in the non reducing glucosaminyl residue. Comparison of the Yersinia lipid A structures shows that they do not differ significantly in their general architecture from those of Enterobacteria [13,16].
5. Discussion Chemical composition and structural analysis of Yersinia LPS reported in this review show common structural features mainly in the core-lipid A regions. LPS is an important factor in the virulence of Yersinia but until now the role of LPS in the virulence of Y. pestis was not clear. Achtman et al. [44] showed that Y. pestis and Y. pseudotuberculosis are genetically very closely related. Recently Skurnik et al. [45] observed that Y. pestis has evolved from Y. pseudotuberculosis. These authors provide evidence that one step in the evolution of Y. pestis from Y. pseudotuberculosis O: 1b was the inactivation of the O-antigen gene cluster. It seems that the expression of this antigen is not beneficial for the virulence of the agent of bubonic plague. However the O-side chain of LPS is an important factor in the virulence of a range of pathogens including Y. enterocolitica
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Fig. 1. Schematic representation of the major lipid A component of (a) Y. enterocolitica O: 11,23 and O: 11,24; Y. ruckeri O: 1 and O: 2 (b) Y. enterocolitica O: 3 and O: 9. (c) Y. pestis. (d) E. coli. The variable part of the structures is presented in bold. The configuration of the double bond of the C16:1 was not determined (Aussel et al. [16]).
[46]. The genome sequence of Y. pestis reported recently by Parkhill et al. [47], revealed genes which encode unknown surface antigens having a role in the virulence of this bacterium.
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Acknowledgements We thank Dr H. Mayer for carefully reading this review. We also thank Drs J. Radziejewska-Lebrecht, M.TH. Pommier, Y. Knirel and G. Pellon for help in searching the literature.
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