**11. References**


[8] Steeghs L, Jennings MP, Poolman JT, van der Ley P. (1997) Isolation and characterization of the *Neisseria meningitidis lpx*D-*fab*Z-*lpx*A gene cluster involved in lipid A biosynthesis. Gene 190: 263–70.

94 The Complex World of Polysaccharides

interchange produced by lateral transference of genes which leads to strain selection with

One of the most important aspects of LPS function is its participation as immunogenic molecule and its role in bacterial classification based on O antigen and its variability. In general, it is seen that the modifications of O antigen play an important role in the process of infection including the adherence, the colonization, and the ability to evade defensive

The study of the events of variation of LPS and its effects on pathogenecity and virulence represents a field of study of great interest to understand bacterial physiology and its

The immunogenicity and variability of O antigen confer to gram-negative bacteria an important characteristic for its serological typification. The 0-antigen is subject to an intense selection on the part of immune system, which could be the principal factor for the different forms in which it is presented. For this reason, the variability of O antigen has been an area of

*Departamento de Medicina Experimental, Instituto, Nacional de Pediatría, México D.F., México* 

*Departamento de Sistemas Biológicos, División CBS, Universidad Autónoma Metropolitana Unidad,* 

[1] Erridge C, Bennett-Guerrero E, Poxton IR. (2002) Structure and function of

[2] Silhavy TJ, Kahne D, Walker S. (2010) The bacterial cell envelope. Cold Spring Harb

[3] Flemming H-C, Neu TR, Wozniak DJ. (2007) The EPS matrix: the «house of biofilm cells».

[4] Pier GB. (2007) *Pseudomonas aeruginosa* lipopolysaccharide: a major virulence factor, initiator of inflammation and target for effective immunity. Int. J. Med. Microbiol

[5] Franklin MJ, Nivens DE, Weadge JT, Howell PL. (2011) Biosynthesis of the *Pseudomonas aeruginosa* extracellular polysaccharides, Alginate, Pel, and Psl. Front Microbiol 2:167. [6] Erridge C, Stewart J, Bennett-Guerrero E, McIntosh TJ, Poxton IR. (2002) The biological activity of a liposomal complete core lipopolysaccharide vaccine. J. Endotoxin Res 8:39–46. [7] Lerouge I, Vanderleyden J. (2002) O-antigen structural variation: mechanisms and possible roles in animal/plant-microbe interactions. FEMS Microbiol. Rev 26:17–47.

Rosa Eugenia Reyes, Alejandra Aquino Andrade, Rafael Coria Jiménez,

Carolina Romo González and Maribel Ortiz Herrera

lipopolysaccharides. Microbes Infect 4:837–51.

*Xochimilco, Coyoacán, México D.F., México* 

Perspect Biol 2:a000414.

J. Bacteriol 189:7945–7.

new characteristics and the evolution of the bacteria by modification of this structure.

mechanisms of the host especially the innate resistance.

mechanisms of adaptation and evolution.

intense research.

**Author details** 

Rosa Eugenia Reyes

**11. References** 

297:277–95.

	- [26] Gunn JS. (2008) The *Salmonella* PmrAB regulon: lipopolysaccharide modifications, antimicrobial peptide resistance and more. Trends Microbiol. 16:284–90.
	- [27] Marchal K, De Keersmaecker S, Monsieurs P, van Boxel N, Lemmens K, Thijs G, Vanderleyden J, De Moor B. (2004) In silico identification and experimental validation of PmrAB targets in *Salmonella typhimurium* by regulatory motif detection. Genome Biol. 5:R9.
	- [28] Heinrichs DE, Yethon JA, Whitfield C. (1998) Molecular basis for structural diversity in the core regions of the lipopolysaccharides of *Escherichia coli* and *Salmonella enterica*. Mol. Microbiol. 30:221–32.
	- [29] Rietschel ET, Kirikae T, Schade FU, Mamat U, Schmidt G, Loppnow H, Ulmer AJ, Zähringer U, Seydel U, Di Padova F. (1994) Bacterial endotoxin: molecular relationships of structure to activity and function. FASEB J. 8:217–25.
	- [30] Bailey MJ, Hughes C, Koronakis V. (1997) RfaH and the ops element, components of a novel system controlling bacterial transcription elongation. Mol. Microbiol. 26:845–51.
	- [31] Geerlof A, Lewendon A, Shaw WV. (1999) Purification and characterization of phosphopantetheine adenylyltransferase from *Escherichia coli*. J. Biol. Chem. 274:27105–11.
	- [32] Feng L, Han W, Wang Q, Bastin DA, Wang L. (2005) Characterization of *Escherichia coli*  O86 O-antigen gene cluster and identification of O86-specific genes. Vet. Microbiol. 106:241–8.
	- [33] Clark CG, Kropinski AM, Parolis H, Grant CCR, Trout-Yakel KM, Franklin K, Ng L-K, Paramonov NA, Parolis LAS, Rahn K, Tabor H. (2009) *Escherichia coli* O123 O antigen genes and polysaccharide structure are conserved in some *Salmonella enterica* serogroups. J. Med. Microbiol. 58:884–94.
	- [34] Sozhamannan S, Deng YK, Li M, Sulakvelidze A, Kaper JB, Johnson JA, Nair GB, Morris JG Jr. (1999) Cloning and sequencing of the genes downstream of the *wbf* gene cluster of *Vibrio cholerae* serogroup O139 and analysis of the junction genes in other serogroups. Infect. Immun. 67:5033–40.
	- [35] Keenleyside WJ, Whitfield C. (1996) A Novel Pathway for O-Polysaccharide Biosynthesis in *Salmonella enterica* Serovar Borreze. J. Biol. Chem. 271:28581–92.
	- [36] Wang L, Jensen S, Hallman R, Reeves PR. (1998) Expression of the O antigen gene cluster is regulated by RfaH through the JUMPstart sequence. FEMS Microbiol. Lett. 165:201–6.
	- [37] Liu B, Knirel YA, Feng L, Perepelov AV, Senchenkova SN, Wang Q, Reeves PR, Wang L. (2008) Structure and genetics of *Shigella* O antigens. FEMS Microbiol. Rev. 32:627–53.
	- [38] Samuel G, Reeves P. (2003) Biosynthesis of O-antigens: genes and pathways involved in nucleotide sugar precursor synthesis and O-antigen assembly. Carbohydr. Res. 338:2503–19.
	- [39] Olson MV, Kas A, Bubb K, Qui R, Smith EE, Raymond CK, Kaul R. (2004) Hypervariability, suppressed recombination and the genetics of individuality. Philos. Trans. R. Soc. Lond., B, Biol. Sci .359:129–40.
	- [40] Dixon DR, Darveau RP. (2005) Lipopolysaccharide heterogeneity: innate host responses to bacterial modification of lipid a structure. J. Dent. Res. 84:584–95.
	- [41] Lawson AJ, Chart H, Dassama MU, Threlfall EJ. (2002) Heterogeneity in expression of lipopolysaccharide by strains of *Salmonella enterica* serotype Typhimurium definitive phage type 104 and related phage types. Lett. Appl. Microbiol. 34:428–32.
	- [42] Tso MD, Dooley JS. (1995) Temperature-dependent protein and lipopolysaccharide expression in clinical *Aeromonas* isolates. J. Med. Microbiol. 42:32–8.

[43] Reyes RE, Ramírez Saad H, Pérez J, Meléndez E, Navarro A, Murata Ch, Cravioto A. (2010) Intra-strain variation of lipopolysaccharides from *Citrobacter freundii* E9750 by *in vitro* passages. Afr. J. Microbiol. Res. 4:1324–31.

96 The Complex World of Polysaccharides

Mol. Microbiol. 30:221–32.

106:241–8.

[26] Gunn JS. (2008) The *Salmonella* PmrAB regulon: lipopolysaccharide modifications,

[27] Marchal K, De Keersmaecker S, Monsieurs P, van Boxel N, Lemmens K, Thijs G, Vanderleyden J, De Moor B. (2004) In silico identification and experimental validation of PmrAB targets in *Salmonella typhimurium* by regulatory motif detection. Genome Biol. 5:R9. [28] Heinrichs DE, Yethon JA, Whitfield C. (1998) Molecular basis for structural diversity in the core regions of the lipopolysaccharides of *Escherichia coli* and *Salmonella enterica*.

[29] Rietschel ET, Kirikae T, Schade FU, Mamat U, Schmidt G, Loppnow H, Ulmer AJ, Zähringer U, Seydel U, Di Padova F. (1994) Bacterial endotoxin: molecular relationships

[30] Bailey MJ, Hughes C, Koronakis V. (1997) RfaH and the ops element, components of a novel system controlling bacterial transcription elongation. Mol. Microbiol. 26:845–51. [31] Geerlof A, Lewendon A, Shaw WV. (1999) Purification and characterization of phosphopantetheine adenylyltransferase from *Escherichia coli*. J. Biol. Chem. 274:27105–11. [32] Feng L, Han W, Wang Q, Bastin DA, Wang L. (2005) Characterization of *Escherichia coli*  O86 O-antigen gene cluster and identification of O86-specific genes. Vet. Microbiol.

[33] Clark CG, Kropinski AM, Parolis H, Grant CCR, Trout-Yakel KM, Franklin K, Ng L-K, Paramonov NA, Parolis LAS, Rahn K, Tabor H. (2009) *Escherichia coli* O123 O antigen genes and polysaccharide structure are conserved in some *Salmonella enterica*

[34] Sozhamannan S, Deng YK, Li M, Sulakvelidze A, Kaper JB, Johnson JA, Nair GB, Morris JG Jr. (1999) Cloning and sequencing of the genes downstream of the *wbf* gene cluster of *Vibrio cholerae* serogroup O139 and analysis of the junction genes in other serogroups.

[35] Keenleyside WJ, Whitfield C. (1996) A Novel Pathway for O-Polysaccharide Biosynthesis in *Salmonella enterica* Serovar Borreze. J. Biol. Chem. 271:28581–92. [36] Wang L, Jensen S, Hallman R, Reeves PR. (1998) Expression of the O antigen gene cluster is regulated by RfaH through the JUMPstart sequence. FEMS Microbiol. Lett. 165:201–6. [37] Liu B, Knirel YA, Feng L, Perepelov AV, Senchenkova SN, Wang Q, Reeves PR, Wang L. (2008) Structure and genetics of *Shigella* O antigens. FEMS Microbiol. Rev. 32:627–53. [38] Samuel G, Reeves P. (2003) Biosynthesis of O-antigens: genes and pathways involved in nucleotide sugar precursor synthesis and O-antigen assembly. Carbohydr. Res. 338:2503–19. [39] Olson MV, Kas A, Bubb K, Qui R, Smith EE, Raymond CK, Kaul R. (2004) Hypervariability, suppressed recombination and the genetics of individuality. Philos.

[40] Dixon DR, Darveau RP. (2005) Lipopolysaccharide heterogeneity: innate host responses

[41] Lawson AJ, Chart H, Dassama MU, Threlfall EJ. (2002) Heterogeneity in expression of lipopolysaccharide by strains of *Salmonella enterica* serotype Typhimurium definitive

[42] Tso MD, Dooley JS. (1995) Temperature-dependent protein and lipopolysaccharide

to bacterial modification of lipid a structure. J. Dent. Res. 84:584–95.

expression in clinical *Aeromonas* isolates. J. Med. Microbiol. 42:32–8.

phage type 104 and related phage types. Lett. Appl. Microbiol. 34:428–32.

antimicrobial peptide resistance and more. Trends Microbiol. 16:284–90.

of structure to activity and function. FASEB J. 8:217–25.

serogroups. J. Med. Microbiol. 58:884–94.

Trans. R. Soc. Lond., B, Biol. Sci .359:129–40.

Infect. Immun. 67:5033–40.

