Extracellular proteins secreted by probiotic bacteria as mediators of effects that promote mucosa-bacteria interactions

Microbiology

Volumn 156, Issue 11, 2010, Pages 3232-3242

  Sánchez, Borja ^a   Urdaci, María C ^b   Margolles, Abelardo ^a  

^a DEPARTMENT OF MICROBIOLOGY AND BIOCHEMISTRY OF DAIRY PRODUCTS   (Spain)

^b UNIVERSITÉ DE BORDEAUX   (France)

Author keywords

[No Author keywords available]

Indexed keywords

BACTERIAL PROTEIN; FLAGELLIN; PROBIOTIC AGENT;

BIFIDOBACTERIUM; CYTOKINE PRODUCTION; DIARRHEA; ELECTROLYTE TRANSPORT; ENTEROPATHY; ESCHERICHIA; EXTRACELLULAR SPACE; GASTROINTESTINAL TRACT; HOMEOSTASIS; HOST CELL; HUMAN; IMMUNOMODULATION; INTESTINE FLORA; INTESTINE MUCOSA; LACTOBACILLUS; NONHUMAN; PERMEABILITY BARRIER; PRIORITY JOURNAL; PROTEIN INTERACTION; SHORT SURVEY;

BACTERIA; BACTERIAL PROTEINS; FLAGELLIN; HOMEOSTASIS; HUMANS; IMMUNOMODULATION; INTESTINAL MUCOSA; PROBIOTICS; RECEPTORS, CELL SURFACE;

BACTERIA (MICROORGANISMS); BIFIDOBACTERIUM; ESCHERICHIA; LACTOBACILLUS;

EID: 78049494627     PISSN: 13500872     EISSN: None     Source Type: Journal    
DOI: 10.1099/mic.0.044057-0     Document Type: Short Survey

References (89)

1. Abdelaziz,D.H., Amr, K. & Am, A. O. (2010). Nlrc4/Ipaf/CLAN/CARD12: more than a flagellin sensor. Int J Biochem Cell Biol 42, 789-791.
2. Adams, R. J., Heazlewood, S. P., Gilshenan, K. S., O'Brien, M., McGuckin, M. A. & Florin, T. H. (2008). IgG antibodies against common gut bacteria are more diagnostic for Crohn's disease than IgG against mannan or flagellin. Am J Gastroenterol 103, 386-396. (Pubitemid 351248712)
3. Altenhoefer, A., Oswald, S., Sonnenborn, U., Enders, C., Schulze, J., Hacker, J. & Oelschlaeger, T. A. (2004). The probiotic Escherichia coli strain Nissle 1917 interferes with invasion of human intestinal epithelial cells by different enteroinvasive bacterial pathogens. FEMS Immunol Med Microbiol 40, 223-229. (Pubitemid 38368186)
4. Anokhina, I. V., Kravtsov, E. G., Yashina, N. V., Ermolaev, A. V., Chesnokova, V. L. & Dalin, M. V. (2006a). Characterization of surface adhesins of lactobacilli used in production of probiotic preparations. Bull Exp Biol Med 141, 716-719.
5. Anokhina, I. V., Kravtsov, E. G., Yashina, N. V., Ermolaev, A. V., Chesnokova, V. L. & Dalin, M. V. (2006b). Surface adhesins of lactobacilli loosely connected to the cell wall and eliminated into the environment during culturing in liquid nutrient media. Bull Exp Biol Med 142, 609-613.
6. Antikainen, J., Anton, L., Sillanpaa, J. & Korhönen, T. K. (2002). Domains in the S-layer protein CbsA of Lactobacillus crispatus involved in adherence to collagens, laminin and lipoteichoic acids and in self-assembly. Mol Microbiol 46, 381-394. (Pubitemid 35340951)
7. Araya, M., Morelli, L., Reid, G., Sanders, M. E. & Stanton, C. (2002). Guidelines for the evaluation of probiotics in foods. FAO/WHO report. http://www.who.int/foodsafety/fs-management/en/probiotic-guidelines.pdf.
8. Arvola, T., Laiho, K., Torkkeli, S., Mykkanen, H., Salminen, S., Maunula, L. & Isolauri, E. (1999). Prophylactic Lactobacillus GG reduces antibiotic-associated diarrhea in children with respiratory infections: a randomized study. Pediatrics 104, e64.
9. Asong, J., Wolfert, M. A., Maiti, K. K., Miller, D. & Boons, G. J. (2009). Binding and cellular activation studies reveal that Toll-like receptor 2 can differentially recognize peptidoglycan from Gram-positive and Gram-negative bacteria. J Biol Chem 284, 8643-8653.
10. Bansal, T., Alaniz, R. C., Wood, T. K. & Jayaraman, A. (2010). The bacterial signal indole increases epithelial-cell tight-junction resistance and attenuates indicators of inflammation. Proc Natl Acad Sci U S A 107, 228-233.
11. Barinov, A., Loux, V., Hammani, A., Nicolas, P., Langella, P., Ehrlich, D., Maguin, E. & van de Gutche, M. (2009). Prediction of surface exposed proteins in Streptococcus pyogenes, with a potential application to other Gram-positive bacteria. Proteomics 9, 61-73.
12. Båth, K., Roos, S., Wall, T. & Jonsson, H. (2005). The cell surface of Lactobacillus reuteri ATCC 55730 highlighted by identification of 126 extracellular proteins from the genome sequence. FEMS Microbiol Lett 253, 75-82. (Pubitemid 41605397)
13. Beck, H. C., Madsen, S. M., Glenting, J., Petersen, J., Israelsen, H., Norrelykke, M. R., Antonsson, M. & Hansen, A. M. (2009). Proteomic analysis of cell surface-associated proteins from probiotic Lactobacillus plantarum. FEMS Microbiol Lett 297, 61-66.
14. Benz, I. & Schmidt, M. A. (2002). Never say never again: protein glycosylation in pathogenic bacteria. Mol Microbiol 45, 267-276. (Pubitemid 34779823)
15. Berks, B. C. (1996). A common export pathway for proteins binding complex redox cofactors? Mol Microbiol 22, 393-404.
16. Binet, R., Letoffe, S., Ghigo, J. M., Delepelaire, P. & Wandersman, C. (1997). Protein secretion by Gram-negative bacterial ABC exporters - a review. Gene 192, 7-11. (Pubitemid 27267473)
17. Boesten, R. J. & de Vos, W. M. (2008). Interactomics in the human intestine: lactobacilli and bifidobacteria make a difference. J Clin Gastroenterol 42, S163-S167.
18. Borody, T. J. (2000). "Flora power" - fecal bacteria cure chronic C. difficile diarrhea. Am J Gastroenterol 95, 3028-3029.
19. - exchange activity in Caco-2 cells via PI-3 kinase mediated pathway. J Nutr 138, 1355-1359.
20. Buist, G., Ridder, A. N., Kok, J. & Kuipers, O. P. (2006). Different subcellular locations of secretome components of Gram-positive bacteria. Microbiology 152, 2867-2874. (Pubitemid 44542368)
21. Caballero-Franco, C., Keller, K., De Simone, C. & Chadee, K. (2007). The VSL#3 probiotic formula induces mucin gene expression and secretion in colonic epithelial cells. Am J Physiol Gastrointest Liver Physiol 292, G315-G322. (Pubitemid 46090273)
22. Candela, M., Biagi, E., Centanni, M., Turroni, S., Vici, M., Musiani, F., Vitali, B., Bergmann, S., Hammerschmidt, S. & Brigidi, P. (2009). Bifidobacterial enolase, a cell surface receptor for human plasminogen involved in the interaction with the host. Microbiology 155, 3294-3303.
23. Castaldo, C., Vastano, V., Siciliano, R. A., Candela, M., Vici, M., Muscariello, L., Marasco, R. & Sacco, M. (2009). Surface displaced alfa-enolase of Lactobacillus plantarum is a fibronectin binding protein. Microb Cell Fact 8, 14.
24. Chen, X., Xu, J., Shuai, J., Chen, J., Zhang, Z. & Fang, W. (2007). The S-layer proteins of Lactobacillus crispatus strain ZJ001 is responsible for competitive exclusion against Escherichia coli O157:H7 and Salmonella typhimurium. Int J Food Microbiol 115, 307-312. (Pubitemid 46400887)
25. Choi, C. H., Kim, T. I., Lee, S. K., Yang, K. M. & Kim, W. H. (2008). Effect of Lactobacillus GG and conditioned media on IL-1beta-induced IL-8 production in Caco-2 cells. Scand J Gastroenterol 43, 938-947.
26. Driessen, A. J. & Nouwen, N. (2008). Protein translocation across the bacterial cytoplasmic membrane. Annu Rev Biochem 77, 643-667.
27. Ewaschuk, J. B., Díaz, H., Meddings, L., Diederichs, B., Dmytrash, A., Backer, J., Looijer-van Langen, M. & Madsen, K. L. (2008). Secreted bioactive factors from Bifidobacterium infantis enhance epithelial cell barrier function. Am J Physiol Gastrointest Liver Physiol 295, G1025-G1034.
28. Favier, C. F., de Vos, W. M. & Akkermans, A. D. (2003). Development of bacterial and bifidobacterial communities in feces of newborn babies. Anaerobe 9, 219-229.
29. Furrie, E., Macfarlane, S., Cummings, J. H. & Macfarlane, G. T. (2004). Systemic antibodies towards mucosal bacteria in ulcerative colitis and Crohn's disease differentially activate the innate immune response. Gut 53, 91-98.
30. Gewirtz, A. T. (2006). Flag in the crossroads: flagellin modulates innate and adaptive immunity. Curr Opin Gastroenterol 22, 8-12.
31. Gill, S. R., Pop, M., Deboy, R. T., Eckburg, P. B., Turnbaugh, P. J., Samuel, B. S., Gordon, J. I., Relman, D. A., Fraser-Liggett, C. M. & Nelson, K. E. (2006). Metagenomic analysis of the human distal gut microbiome. Science 312, 1355-1359.
32. Grozdanov, L., Raasch, C., Schulze, J., Sonnenborn, U., Gottschalk, G., Hacker, J. & Dobrindt,U. (2004). Analysis of the genome structure of the nonpathogenic probiotic Escherichia coli strain Nissle 1917. J Bacteriol 186, 5432-5441.
33. Gueimonde, M., Ouwehand, A., Huhtinen, H., Salminen, E. & Salminen, S. (2007). Qualitative and quantitative analyses of the bifidobacterial microbiota in the colonic mucosa of patients with colorectal cancer, diverticulitis and inflammatory bowel disease. World J Gastroenterol 13, 3985-3989.
34. Hoarau, C., Martin, L., Faugaret, D., Baron, C., Dauba, A., Aubert- Jacquin, C., Velge-Roussel, F. & Lebranchu, Y. (2008). Supernatant from Bifidobacterium differentially modulates transduction signaling pathways for biological functions of human dendritic cells. PLoS ONE 3, e2753.
35. Hooper, L. V. & Gordon, J. I. (2001). Commensal host-bacterial relationships in the gut. Science 292, 1115-1118.
36. Hooper, L. V., Wong, M. H., Thelin, A., Hansson, L., Falk, P. G. & Gordon, J. I. (2001). Molecular analysis of commensal host-microbial relationships in the intestine. Science 291, 881-884.
37. Hurmalainen, V., Edelman, S., Antikainen, J., Baumann, M., Lahteenmaki, K. & Korhonen, T. K. (2007). Extracellular proteins of Lactobacillus crispatus enhance activation of human plasminogen. Microbiology 153, 1112-1122.
38. Iltanen, S., Tervo, L., Halttunen, T., Wei, B., Braun, J., Rantala, I., Honkanen, T., Kronenberg, M., Cheroutre, H. & other authors (2006). Elevated serum anti-I2 and anti-OmpW antibody levels in children with IBD. Inflamm Bowel Dis 12, 389-394.
39. Ivanov, D., Emonet, C., Foata, F., Affolter, M., Delley, M., Fisseha, M., Blum-Sperisen, S., Kochhar, S. & Arigoni, F. (2006). A serpin from the gut bacterium Bifidobacterium longum inhibits eukaryotic elastase-like serine proteases. J Biol Chem 281, 17246-17252. (Pubitemid 43909533)
40. Ivison, S. M. & Steiner, T. S. (2008). Anti-flagellin antibodies in irritable bowel syndrome: another attack on our commensals? Neurogastroenterol Motil 20, 1081-1085.
41. Iwamori, M., Shibagaki, T., Nakata, Y., Adachi, S. & Nomura, T. (2009). Distribution of receptor glycolipids for lactobacilli in murine digestive tract and production of antibodies crossreactive with them by immunization of rabbits with lactobacilli. J Biochem 146, 185-191.
42. Jeffery, C. J. (2003). Moonlighting proteins: old proteins learning new tricks. Trends Genet 19, 415-417. (Pubitemid 36930022)
43. Jongbloed, J. D., van der Ploeg, R. & van Dijl, J. M. (2006). Bifunctional TatA subunits in minimal Tat protein translocases. Trends Microbiol 14, 2-4.
44. Kankainen, M., Paulin, L., Tynkkynen, S., von Ossowski, I., Reunanen, J., Partanen, P., Satokari, R., Vesterlund, S., Hendrickx, A. P. & other authors (2009). Comparative genomic analysis of Lactobacillus rhamnosus GG reveals pili containing a human- mucus binding protein. Proc Natl Acad Sci U S A 106, 17193-17198.
45. Kinoshita, H., Uchida, H., Kawai, Y., Kawasaki, T., Wakahara, N., Matsuo, H., Watanabe, M., Kitazawa, H., Ohnuma, S. & other authors (2008a). Cell surface Lactobacillus plantarum LA 318 glyceraldehyde-3-phosphate dehydrogenase (GAPDH) adheres to human colonic mucin. J Appl Microbiol 104, 1667-1674.
46. Kinoshita, H., Wakahara, N., Watanabe, M., Kawasaki, T., Matsuo, H., Kawai, Y., Kitazawa, H., Ohnuma, S., Miura, K. & other authors (2008b). Cell surface glyceraldehyde-3-phosphate dehydrogenase (GAPDH) of Lactobacillus plantarum LA 318 recognizes human A and B blood group antigens. Res Microbiol 159, 685-691.
47. Kleerebezem, M., Hols, P., Bernard, E., Rolain, T., Zhou, M., Siezen, R. J. & Bron, P. A. (2010). The extracellular biology of the lactobacilli. FEMS Microbiol Rev 34, 199-230.
48. Konstantinov, S. R., Smidt, H., de Vos, W. M., Bruijns, S. C., Singh, S. K., Valence, F., Molle, D., Lortal, S., Altermann, E. & other authors (2008). S layer protein A of Lactobacillus acidophilus NCFM regulates immature dendritic cell and T cell functions. Proc Natl Acad Sci U S A 105, 19474-19479.
49. Kruis, W., Fric, P., Pokrotnieks, J., Lukas, M., Fixa, B., Kascak, M., Kamm, M. A., Weismueller, J., Beglinger, C. & other authors (2004). Maintaining remission of ulcerative colitis with the probiotic Escherichia coli Nissle 1917 is as effective as with standard mesalazine. Gut 53, 1617-1623.
50. Lasaro, M. A., Salinger, N., Zhang, J., Wang, Y., Zhong, Z., Goulian, M. & Zhu, J. (2009). F1C fimbriae play an important role in biofilm formation and intestinal colonization by the Escherichia coli commensal strain Nissle 1917. Appl Environ Microbiol 75, 246-251.
51. Lebeer, S., Ceuppens, J., Geboes, K., Van Assche, G., Rutgeerts, P., Vanderleyden, J. & De Keersmaecker, S. C. (2007a). Mechanisms of probiotic-host interaction with IBD as a case study: a role for exopolysaccharides? Commun Agric Appl Biol Sci 72, 41-45.
52. Lebeer, S., Verhoeven, T. L., Perea, V. M., Vanderleyden, J. & De Keersmaecker, S. C. (2007b). Impact of environmental and genetic factors on biofilm formation by the probiotic strain Lactobacillus rhamnosus GG. Appl Environ Microbiol 73, 6768-6775.
53. Lebeer, S., Vanderleyden, J. & De Keersmaecker, S. C. (2008). Genes and molecules of lactobacilli supporting probiotic action. Microbiol Mol Biol Rev 72, 728-764.
54. Lebeer, S., Vanderleyden, J. & De Keersmaecker, S. C. (2010). Host interactions of probiotic bacterial surface molecules: comparison with commensals and pathogens. Nat Rev Microbiol 8, 171-184.
55. López, P., Gueimonde, M., Margolles, A. & Suarez, A. (2010). Distinct Bifidobacterium strains drive different immune responses in vitro. Int J Food Microbiol 138, 157-165.
56. Lu, R., Fasano, S., Madayiputhiya, N., Morin, N. P., Nataro, J. & Fasano, A. (2009). Isolation, identification, and characterization of small bioactive peptides from Lactobacillus GG conditional media that exert both anti-Gram-negative and Gram-positive bactericidal activity. J Pediatr Gastroenterol Nutr 49, 23-30.
57. Mitsuma, T., Odajima, H., Momiyama, Z., Watanabe, K., Masuguchi, M., Sekine, T., Shidara, S. & Hirano, S. (2008). Enhancement of gene expression by a peptide p(CHWPR) produced by Bifidobacterium lactis BB-12. Microbiol Immunol 52, 144-155.
58. Müller, S., Schaffer, T., Schoepfer, A. M., Hilty, A., Bodmer, T. & Seibold, F. (2008). Partial overlap of anti-mycobacterial, and anti-Saccharomyces cerevisiae mannan antibodies in Crohn's disease. World J Gastroenterol 14, 3650-3661.
59. Natale, P., Bruser, T. & Driessen, A. J. (2008). Sec- and Tat-mediated protein secretion across the bacterial cytoplasmic membrane - distinct translocases and mechanisms. Biochim Biophys Acta 1778, 1735-1756.
60. Pant, A. R., Graham, S. M., Allen, S. J., Harikul, S., Sabchareon, A., Cuevas, L. & Hart, C. A. (1996). Lactobacillus GG and acute diarrhoea in young children in the tropics. J Trop Pediatr 42, 162-165.
61. Ramiah, K., van Reenen, C. A. & Dicks, L. M. (2008). Surface-bound proteins of Lactobacillus plantarum 423 that contribute to adhesion of Caco-2 cells and their role in competitive exclusion and displacement of Clostridium sporogenes and Enterococcus faecalis. Res Microbiol 159, 470-475.
62. Ren, T., Zamboni, D. S., Roy, C. R., Dietrich, W. F. & Vance, R. E. (2006). Flagellin-deficient Legionella mutants evade caspase-1- and Naip5-mediated macrophage immunity. PLoS Pathog 2, e18.
63. Salminen, S. J., Gueimonde, M. & Isolauri, E. (2005). Probiotics that modify disease risk. J Nutr 135, 1294-1298.
64. Salminen, S., Collado, M. C., Isolauri, E. & Gueimonde, M. (2009). Microbial-host interactions: selecting the right probiotics and prebiotics for infants. Nestle Nutr Workshop Ser Pediatr Program 64, 201-213.
65. Sánchez, B., Bressollier, P. & Urdaci, M. C. (2008a). Exported proteins in probiotic bacteria: adhesion to intestinal surfaces, host immunomodulation and molecular cross-talking with the host. FEMS Immunol Med Microbiol 54, 1-17.
66. Sánchez, B., Champomier-Vergès, M. C., Anglade, P., Baraige, F., de los Reyes-Gavilan, C. G., Margolles, A. & Zagorec, M. (2008b). A preliminary analysis of Bifidobacterium longum exported proteins by two-dimensional electrophoresis. J Mol Microbiol Biotechnol 14, 74-79.
67. Sánchez, B., Arias, S., Chaignepain, S., Denayrolles, M., Schmitter, J. M., Bressollier, P. & Urdaci, M. C. (2009a). Identification of surface proteins involved in the adhesion of a probiotic Bacillus cereus strain to mucin and fibronectin. Microbiology 155, 1708-1716.
68. Sánchez, B., Chaignepain, S., Schmitter, J. M. & Urdaci, M. C. (2009b). A method for the identification of proteins secreted by lactic acid bacteria grown in complex media. FEMS Microbiol Lett 295, 226-229.
69. Sánchez, B., Schmitter, J. M. & Urdaci, M. C. (2009c). Identification of novel proteins secreted by Lactobacillus plantarum that bind to mucin and fibronectin. J Mol Microbiol Biotechnol 17, 158-162.
70. Sánchez, B., Schmitter, J. M. & Urdaci, M. C. (2009d). Identification of novel proteins secreted by Lactobacillus rhamnosus GG grown in de Man-Rogosa-Sharpe broth. Lett Appl Microbiol 48, 618-622.
71. Schlee, M., Wehkamp, J., Altenhoefer, A., Oelschlaeger, T. A., Stange, E. F. & Fellermann, K. (2007). Induction of human beta-defensin 2 by the probiotic Escherichia coli Nissle 1917 is mediated through flagellin. Infect Immun 75, 2399-2407.
72. Schlee, M., Harder, J., Köten, B., Stange, E. F., Wehkamp, J. & Fellermann, K. (2008). Probiotic lactobacilli and VSL#3 induce enterocyte beta-defensin 2. Clin Exp Immunol 151, 528-535.
73. Schoepfer, A. M., Schaffer, T., Seibold-Schmid, B., Muller, S. & Seibold, F. (2008). Antibodies to flagellin indicate reactivity to bacterial antigens in IBS patients. Neurogastroenterol Motil 20, 1110-1118.
74. Seth, A., Yan, F., Polk, D. B. & Rao, R. K. (2008). Probiotics ameliorate the hydrogen peroxide-induced epithelial barrier disruption by a PKC- and MAP kinase-dependent mechanism. Am J Physiol Gastrointest Liver Physiol 294, G1060-G1069. (Pubitemid 351520059)
75. Sibbald, M. J. J. B. & van Dijl, J. M. (2009). Bacterial Secreted Proteins: Secretory Mechanisms and Role in Pathogenesis. Edited by K. Wooldridge. Norwich, UK: Caister Academic Press. Horizon Scientific Press.
76. Stappenbeck, T. S., Hooper, L. V. & Gordon, J. I. (2002). Developmental regulation of intestinal angiogenesis by indigenous microbes via Paneth cells. Proc Natl Acad Sci U S A 99, 15451-15455.
77. Stringer, C. (2003). Human evolution: out of Ethiopia. Nature 423, 692-693.
78. Tao, Y., Drabik, K. A., Waypa, T. S., Musch, M. W., Alverdy, J. C., Schneewind, O., Chang, E. B. & Petrof, E. O. (2006). Soluble factors from Lactobacillus GG activate MAPKs and induce cytoprotective heat shock proteins in intestinal epithelial cells. Am J Physiol Cell Physiol 290, C1018-C1030.
79. Turnbaugh, P. J., Ley, R. E., Hamady, M., Fraser-Liggett, C. M., Knight, R. & Gordon, J. I. (2007). The human microbiome project. Nature 449, 804-810.
80. Turner, M. S., Timms, P., Hafner, L. M. & Giffard, P. M. (1997). Identification and characterization of a basic cell surface-located protein from Lactobacillus fermentum BR11. J Bacteriol 179, 3310-3316.
81. Turner, M. S., Hafner, L. M., Walsh, T. & Giffard, P. M. (2004). Identification and characterization of the novel LysM domain-containing surface protein Sep from Lactobacillus fermentum BR11 and its use as a peptide fusion partner in Lactobacillus and Lactococcus. Appl Environ Microbiol 70, 3673-3680.
82. Turroni, F., Marchesi, J. R., Foroni, E., Gueimonde, M., Shanahan, F., Margolles, A., van Sinderen, D. & Ventura, M. (2009). Microbiomic analysis of the bifidobacterial population in the human distal gut. ISME J 3, 745-751.
83. van Pijkeren, J. P., Canchaya, C., Ryan, K. A., Li, Y., Claesson, M. J., Sheil, B., Steidler, L., O'Mahony, L., Fitzgerald, G. F. & other authors (2006). Comparative and functional analysis of sortase-dependent proteins in the predicted secretome of Lactobacillus salivarius UCC118. Appl Environ Microbiol 72, 4143-4153.
84. van Wely, K. H. M., Swaving, J., Freudl, R. & Driessen, A. J. M. (2001). Translocation of proteins across the cell envelope of Gram-positive bacteria. FEMS Microbiol Rev 25, 437-454.
85. Vélez, M. P., De Keersmaecker, S. C. & Vanderleyden, J. (2007). Adherence factors of Lactobacillus in the human gastrointestinal tract. FEMS Microbiol Lett 276, 140-148.
86. von Heijne, G. (1989). The structure of signal peptides from bacterial lipoproteins. Protein Eng 2, 531-534.
87. von Heijne, G. & Abrahmsen, L. (1989). Species-specific variation in signal peptide design. Implications for protein secretion in foreign hosts. FEBS Lett 244, 439-446.
88. Willing, B., Halfvarson, J., Dicksved, J., Rosenquist, M., Järnerot, G., Engstrand, L., Tysk, C. & Jansson, J. K. (2009). Twin studies reveal specific imbalances in the mucosa-associated microbiota of patients with ileal Crohn's disease. Inflamm Bowel Dis 15, 653-660.
89. Yan, F., Cao, H., Cover, T. L., Whitehead, R., Washington, M. K. & Polk, D. B. (2007). Soluble proteins produced by probiotic bacteria regulate intestinal epithelial cell survival and growth. Gastroenterology 132, 562-575.