Molecular mechanisms involved in Bacillus subtilis biofilm formation

Environmental Microbiology

Volumn 17, Issue 3, 2015, Pages 555-565

  Mielich Süss, Benjamin ^a   Lopez, Daniel ^a  

^a UNIVERSITY OF WÜRZBURG   (Germany)

Author keywords

[No Author keywords available]

Indexed keywords

BACILLUS SUBTILIS; BACTERIA (MICROORGANISMS);

BACILLUS SUBTILIS; BIOFILM; GENETICS; GROWTH, DEVELOPMENT AND AGING; PHYSIOLOGY; SIGNAL TRANSDUCTION;

BACILLUS SUBTILIS; BIOFILMS; SIGNAL TRANSDUCTION;

EID: 84925136038     PISSN: 14622912     EISSN: 14622920     Source Type: Journal    
DOI: 10.1111/1462-2920.12527     Document Type: Review

References (113)

1. Aguilar, C., Vlamakis, H., Guzman, A., Losick, R., and Kolter, R. (2010) KinD is a checkpoint protein linking spore formation to extracellular-matrix production in Bacillus subtilis biofilms. MBio 1: e00035-10.
2. Amati, G., Bisicchia, P., and Galizzi, A. (2004) DegU-P represses expression of the motility fla-che operon in Bacillus subtilis. J Bacteriol 186: 6003-6014.
3. Anderson, G.G., and O'Toole, G.A. (2008) Innate and induced resistance mechanisms of bacterial biofilms. Curr Top Microbiol Immunol 322: 85-105.
4. Angelini, T.E., Roper, M., Kolter, R., Weitz, D.A., and Brenner, M.P. (2009) Bacillus subtilis spreads by surfing on waves of surfactant. Proc Natl Acad Sci USA 106: 18109-18113.
5. Asally, M., Kittisopikul, M., Rue, P., Du, Y., Hu, Z., Cagatay, T., etal. (2012) Localized cell death focuses mechanical forces during 3D patterning in a biofilm. Proc Natl Acad Sci USA 109: 18891-18896.
6. Bach, J.N., and Bramkamp, M. (2013) Flotillins functionally organize the bacterial membrane. Mol Microbiol 88: 1205-1217.
7. Banse, A.V., Chastanet, A., Rahn-Lee, L., Hobbs, E.C., and Losick, R. (2008) Parallel pathways of repression and antirepression governing the transition to stationary phase in Bacillus subtilis. Proc Natl Acad Sci USA 105: 15547-15552.
8. Baty, A.M., 3rd, Eastburn, C.C., Techkarnjanaruk, S., Goodman, A.E., and Geesey, G.G. (2000a) Spatial and temporal variations in chitinolytic gene expression and bacterial biomass production during chitin degradation. Appl Environ Microbiol 66: 3574-3585.
9. Baty, A.M., 3rd, Eastburn, C.C., Diwu, Z., Techkarnjanaruk, S., Goodman, A.E., and Geesey, G.G. (2000b) Differentiation of chitinase-active and non-chitinase-active subpopulations of a marine bacterium during chitin degradation. Appl Environ Microbiol 66: 3566-3573.
10. Beauregard, P.B., Chai, Y., Vlamakis, H., Losick, R., and Kolter, R. (2013) Bacillus subtilis biofilm induction by plant polysaccharides. Proc Natl Acad Sci USA 110: E1621-E1630.
11. Blair, K.M., Turner, L., Winkelman, J.T., Berg, H.C., and Kearns, D.B. (2008) A molecular clutch disables flagella in the Bacillus subtilis biofilm. Science 320: 1636-1638.
12. Branda, S.S., Gonzalez-Pastor, J.E., Ben-Yehuda, S., Losick, R., and Kolter, R. (2001) Fruiting body formation by Bacillus subtilis. Proc Natl Acad Sci USA 98: 11621-11626.
13. Branda, S.S., Chu, F., Kearns, D.B., Losick, R., and Kolter, R. (2006) A major protein component of the Bacillus subtilis biofilm matrix. Mol Microbiol 59: 1229-1238.
14. Bryers, J.D. (2008) Medical biofilms. Biotechnol Bioeng 100: 1-18.
15. Cairns, L.S., Marlow, V.L., Bissett, E., Ostrowski, A., and Stanley-Wall, N.R. (2013) A mechanical signal transmitted by the flagellum controls signalling in Bacillus subtilis. Mol Microbiol 90: 6-21.
16. Cava, F., Lam, H., de Pedro, M.A., and Waldor, M.K. (2011) Emerging knowledge of regulatory roles of D-amino acids in bacteria. Cell Mol Life Sci 68: 817-831.
17. Cazorla, F.M., Romero, D., Perez-Garcia, A., Lugtenberg, B.J., Vicente, A., and Bloemberg, G. (2007) Isolation and characterization of antagonistic Bacillus subtilis strains from the avocado rhizoplane displaying biocontrol activity. J Appl Microbiol 103: 1950-1959.
18. Chai, Y., Chu, F., Kolter, R., and Losick, R. (2008) Bistability and biofilm formation in Bacillus subtilis. Mol Microbiol 67: 254-263.
19. Chen, Y., Cao, S., Chai, Y., Clardy, J., Kolter, R., Guo, J.H., and Losick, R. (2012) A Bacillus subtilis sensor kinase involved in triggering biofilm formation on the roots of tomato plants. Mol Microbiol 85: 418-430.
20. Chen, Y., Yan, F., Chai, Y., Liu, H., Kolter, R., Losick, R., and Guo, J.H. (2013) Biocontrol of tomato wilt disease by Bacillus subtilis isolates from natural environments depends on conserved genes mediating biofilm formation. Environ Microbiol 15: 848-864.
21. Chu, F., Kearns, D.B., McLoon, A., Chai, Y., Kolter, R., and Losick, R. (2008) A novel regulatory protein governing biofilm formation in Bacillus subtilis. Mol Microbiol 68: 1117-1127.
22. Chung, J.D., Stephanopoulos, G., Ireton, K., and Grossman, A.D. (1994) Gene expression in single cells of Bacillus subtilis: evidence that a threshold mechanism controls the initiation of sporulation. J Bacteriol 176: 1977-1984.
23. Cosby, W.M., Vollenbroich, D., Lee, O.H., and Zuber, P. (1998) Altered srf expression in Bacillus subtilis resulting from changes in culture pH is dependent on the Spo0K oligopeptide permease and the ComQX system of extracellular control. J Bacteriol 180: 1438-1445.
24. Costerton, J.W., Cheng, K.J., Geesey, G.G., Ladd, T.I., Nickel, J.C., Dasgupta, M., and Marrie, T.J. (1987) Bacterial biofilms in nature and disease. Annu Rev Microbiol 41: 435-464.
25. Cutting, S., Anderson, M., Lysenko, E., Page, A., Tomoyasu, T., Tatematsu, K., etal. (1997) SpoVM, a small protein essential to development in Bacillus subtilis, interacts with the ATP-dependent protease FtsH. J Bacteriol 179: 5534-5542.
26. Dahl, M.K., Msadek, T., Kunst, F., and Rapoport, G. (1992) The phosphorylation state of the DegU response regulator acts as a molecular switch allowing either degradative enzyme synthesis or expression of genetic competence in Bacillus subtilis. J Biol Chem 267: 14509-14514.
27. Davies, D.G., Parsek, M.R., Pearson, J.P., Iglewski, B.H., Costerton, J.W., and Greenberg, E.P. (1998) The involvement of cell-to-cell signals in the development of a bacterial biofilm. Science 280: 295-298.
28. Dietrich, L.E., Teal, T.K., Price-Whelan, A., and Newman, D.K. (2008) Redox-active antibiotics control gene expression and community behavior in divergent bacteria. Science 321: 1203-1206.
29. Dong, Y.H., and Zhang, L.H. (2005) Quorum sensing and quorum-quenching enzymes. J Microbiol 43 Spec No: 101-109.
30. Dong, Y.H., Wang, L.H., Xu, J.L., Zhang, H.B., Zhang, X.F., and Zhang, L.H. (2001) Quenching quorum-sensing-dependent bacterial infection by an N-acyl homoserine lactonase. Nature 411: 813-817.
31. Donlan, R.M. (2002) Biofilms: microbial life on surfaces. Emerg Infect Dis 8: 881-890.
32. Donlan, R.M. (2008) Biofilms on central venous catheters: is eradication possible? Curr Top Microbiol Immunol 322: 133-161.
33. Fall, R., Kinsinger, R.F., and Wheeler, K.A. (2004) A simple method to isolate biofilm-forming Bacillus subtilis and related species from plant roots. Syst Appl Microbiol 27: 372-379.
34. Flemming, H.C., and Wingender, J. (2010) The biofilm matrix. Nat Rev Microbiol 8: 623-633.
35. Fletcher, M. (1977) The effects of culture concentration and age, time, and temperature on bacterial attachment to polystyrene. Can J Microbiol 23: 1-6.
36. Freese, E. (1972) Sporulation of bacilli, a model of cellular differentiation. Curr Top Dev Biol 7: 85-124.
37. Fujita, M., Gonzalez-Pastor, J.E., and Losick, R. (2005) High- and low-threshold genes in the Spo0A regulon of Bacillus subtilis. J Bacteriol 187: 1357-1368.
38. Garcia-Gutierrez, L., Zeriouh, H., Romero, D., Cubero, J., de Vicente, A., and Perez-Garcia, A. (2013) The antagonistic strain Bacillus subtilis UMAF6639 also confers protection to melon plants against cucurbit powdery mildew by activation of jasmonate- and salicylic acid-dependent defence responses. Microb Biotechnol 6: 264-274.
39. Gaur, N.K., Dubnau, E., and Smith, I. (1986) Characterization of a cloned Bacillus subtilis gene that inhibits sporulation in multiple copies. J Bacteriol 168: 860-869.
40. Gonzalez-Pastor, J.E., Hobbs, E.C., and Losick, R. (2003) Cannibalism by sporulating bacteria. Science 301: 510-513.
41. Guttenplan, S.B., Blair, K.M., and Kearns, D.B. (2010) The EpsE flagellar clutch is bifunctional and synergizes with EPS biosynthesis to promote Bacillus subtilis biofilm formation. PLoS Genet 6: e1001243.
42. Hahn, J., Roggiani, M., and Dubnau, D. (1995) The major role of Spo0A in genetic competence is to downregulate abrB, an essential competence gene. J Bacteriol 177: 3601-3605.
43. Hall-Stoodley, L., Costerton, J.W., and Stoodley, P. (2004) Bacterial biofilms: from the natural environment to infectious diseases. Nat Rev Microbiol 2: 95-108.
44. Hobley, L., Ostrowski, A., Rao, F.V., Bromley, K.M., Porter, M., Prescott, A.R., etal. (2013) BslA is a self-assembling bacterial hydrophobin that coats the Bacillus subtilis biofilm. Proc Natl Acad Sci USA 110: 13600-13605.
45. Hobley, L., Kim, S.H., Maezato, Y., Wyllie, S., Fairlamb, A.H., Stanley-Wall, N.R., and Michael, A.J. (2014) Norspermidine is not a self-produced trigger for biofilm disassembly. Cell 156: 844-854.
46. Hochbaum, A.I., Kolodkin-Gal, I., Foulston, L., Kolter, R., Aizenberg, J., and Losick, R. (2011) Inhibitory effects of D-amino acids on Staphylococcus aureus biofilm development. J Bacteriol 193: 5616-5622.
47. Hofer, U. (2014) Biofilms: biofilm disassembly revisited. Nat Rev Microbiol 12: 234.
48. Hsueh, Y.H., Cozy, L.M., Sham, L.T., Calvo, R.A., Gutu, A.D., Winkler, M.E., and Kearns, D.B. (2011) DegU-phosphate activates expression of the anti-sigma factor FlgM in Bacillus subtilis. Mol Microbiol 81: 1092-1108.
49. Jiang, M., Shao, W., Perego, M., and Hoch, J.A. (2000) Multiple histidine kinases regulate entry into stationary phase and sporulation in Bacillus subtilis. Mol Microbiol 38: 535-542.
50. Karatan, E., and Watnick, P. (2009) Signals, regulatory networks, and materials that build and break bacterial biofilms. Microbiol Mol Biol Rev 73: 310-347.
51. Kearns, D.B., Chu, F., Branda, S.S., Kolter, R., and Losick, R. (2005) A master regulator for biofilm formation by Bacillus subtilis. Mol Microbiol 55: 739-749.
52. Kobayashi, K. (2007a) Bacillus subtilis pellicle formation proceeds through genetically defined morphological changes. J Bacteriol 189: 4920-4931.
53. Kobayashi, K. (2007b) Gradual activation of the response regulator DegU controls serial expression of genes for flagellum formation and biofilm formation in Bacillus subtilis. Mol Microbiol 66: 395-409.
54. Kobayashi, K. (2008) SlrR/SlrA control the initiation of biofilm formation in Bacillus subtilis. Mol Microbiol 69: 1399-1410.
55. Kobayashi, K., and Iwano, M. (2012) BslA(YuaB) forms a hydrophobic layer on the surface of Bacillus subtilis biofilms. Mol Microbiol 85: 51-66.
56. Kolodkin-Gal, I., Romero, D., Cao, S., Clardy, J., Kolter, R., and Losick, R. (2010) D-amino acids trigger biofilm disassembly. Science 328: 627-629.
57. Kolodkin-Gal, I., Cao, S., Chai, L., Bottcher, T., Kolter, R., Clardy, J., and Losick, R. (2012) A self-produced trigger for biofilm disassembly that targets exopolysaccharide. Cell 149: 684-692.
58. Kolodkin-Gal, I., Elsholz, A.K., Muth, C., Girguis, P.R., Kolter, R., and Losick, R. (2013) Respiration control of multicellularity in Bacillus subtilis by a complex of the cytochrome chain with a membrane-embedded histidine kinase. Genes Dev 27: 887-899.
59. Kunst, F., Msadek, T., Bignon, J., and Rapoport, G. (1994) The DegS/DegU and ComP/ComA two-component systems are part of a network controlling degradative enzyme synthesis and competence in Bacillus subtilis. Res Microbiol 145: 393-402.
60. Latasa, C., Garcia, B., Echeverz, M., Toledo-Arana, A., Valle, J., Campoy, S., etal. (2012) Salmonella biofilm development depends on the phosphorylation status of RcsB. J Bacteriol 194: 3708-3722.
61. LeDeaux, J.R., Yu, N., and Grossman, A.D. (1995) Different roles for KinA, KinB, and KinC in the initiation of sporulation in Bacillus subtilis. J Bacteriol 177: 861-863.
62. Lopez, D., and Kolter, R. (2010a) Extracellular signals that define distinct and coexisting cell fates in Bacillus subtilis. FEMS Microbiol Rev 34: 134-149.
63. Lopez, D., and Kolter, R. (2010b) Functional microdomains in bacterial membranes. Genes Dev 24: 1893-1902.
64. Lopez, D., Vlamakis, H., and Kolter, R. (2009a) Generation of multiple cell types in Bacillus subtilis. FEMS Microbiol Rev 33: 152-163.
65. Lopez, D., Vlamakis, H., Losick, R., and Kolter, R. (2009b) Cannibalism enhances biofilm development in Bacillus subtilis. Mol Microbiol 74: 609-618.
66. Lopez, D., Vlamakis, H., Losick, R., and Kolter, R. (2009c) Paracrine signaling in a bacterium. Genes Dev 23: 1631-1638.
67. Lopez, D., Fischbach, M.A., Chu, F., Losick, R., and Kolter, R. (2009d) Structurally diverse natural products that cause potassium leakage trigger multicellularity in Bacillus subtilis. Proc Natl Acad Sci USA 106: 280-285.
68. Magnuson, R., Solomon, J., and Grossman, A.D. (1994) Biochemical and genetic characterization of a competence pheromone from B.subtilis. Cell 77: 207-216.
69. Marlow, V.L., Cianfanelli, F.R., Porter, M., Cairns, L.S., Dale, J.K., and Stanley-Wall, N.R. (2014) The prevalence and origin of exoprotease-producing cells in the Bacillus subtilis biofilm. Microbiology 160: 56-66.
70. McLoon, A.L., Guttenplan, S.B., Kearns, D.B., Kolter, R., and Losick, R. (2011) Tracing the domestication of a biofilm-forming bacterium. J Bacteriol 193: 2027-2034.
71. Mielich-Süss, B., Schneider, J., and Lopez, D. (2013) Overproduction of flotillin influences cell differentiation and shape in Bacillus subtilis. MBio 4: e00719-13.
72. Mirani, Z.A., Aziz, M., Khan, M.N., Lal, I., Hassan, N.U., and Khan, S.I. (2013) Biofilm formation and dispersal of Staphylococcus aureus under the influence of oxacillin. Microb Pathog 61-62C: 66-72.
73. Morris, C.E., and Monier, J.M. (2003) The ecological significance of biofilm formation by plant-associated bacteria. Annu Rev Phytopathol 41: 429-453.
74. Morris, V.K., Ren, Q., Macindoe, I., Kwan, A.H., Byrne, N., and Sunde, M. (2011) Recruitment of class I hydrophobins to the air:water interface initiates a multi-step process of functional amyloid formation. J Biol Chem 286: 15955-15963.
75. Murray, E.J., Kiley, T.B., and Stanley-Wall, N.R. (2009) A pivotal role for the response regulator DegU in controlling multicellular behaviour. Microbiology 155: 1-8.
76. Nadell, C.D., Xavier, J.B., and Foster, K.R. (2009) The sociobiology of biofilms. FEMS Microbiol Rev 33: 206-224.
77. Nakano, M.M., Xia, L.A., and Zuber, P. (1991) Transcription initiation region of the srfA operon, which is controlled by the comP-comA signal transduction system in Bacillus subtilis. J Bacteriol 173: 5487-5493.
78. Ogura, M., and Tsukahara, K. (2010) Autoregulation of the Bacillus subtilis response regulator gene degU is coupled with the proteolysis of DegU-P by ClpCP. Mol Microbiol 75: 1244-1259.
79. Ogura, M., Shimane, K., Asai, K., Ogasawara, N., and Tanaka, T. (2003) Binding of response regulator DegU to the aprE promoter is inhibited by RapG, which is counteracted by extracellular PhrG in Bacillus subtilis. Mol Microbiol 49: 1685-1697.
80. O'Toole, G.A., and Kolter, R. (1998) Flagellar and twitching motility are necessary for Pseudomonas aeruginosa biofilm development. Mol Microbiol 30: 295-304.
81. Pandey, A., and Palni, L.M. (1997) Bacillus species: the dominant bacteria of the rhizosphere of established tea bushes. Microbiol Res 152: 359-365.
82. Perego, M., and Hoch, J.A. (1996) Cell-cell communication regulates the effects of protein aspartate phosphatases on the phosphorelay controlling development in Bacillus subtilis. Proc Natl Acad Sci USA 93: 1549-1553.
83. Perego, M., Glaser, P., and Hoch, J.A. (1996) Aspartyl-phosphate phosphatases deactivate the response regulator components of the sporulation signal transduction system in Bacillus subtilis. Mol Microbiol 19: 1151-1157.
84. Pereira, P.M., Filipe, S.R., Tomasz, A., and Pinho, M.G. (2007) Fluorescence ratio imaging microscopy shows decreased access of vancomycin to cell wall synthetic sites in vancomycin-resistant Staphylococcus aureus. Antimicrob Agents Chemother 51: 3627-3633.
85. Pottathil, M., and Lazazzera, B.A. (2003) The extracellular Phr peptide-Rap phosphatase signaling circuit of Bacillus subtilis. Front Biosci 8: d32-d45.
86. Pottathil, M., Jung, A., and Lazazzera, B.A. (2008) CSF, a species-specific extracellular signaling peptide for communication among strains of Bacillus subtilis and Bacillus mojavensis. J Bacteriol 190: 4095-4099.
87. Ramamurthi, K.S., Clapham, K.R., and Losick, R. (2006) Peptide anchoring spore coat assembly to the outer forespore membrane in Bacillus subtilis. Mol Microbiol 62: 1547-1557.
88. Romero, D., Aguilar, C., Losick, R., and Kolter, R. (2010) Amyloid fibers provide structural integrity to Bacillus subtilis biofilms. Proc Natl Acad Sci USA 107: 2230-2234.
89. Romero, D., Vlamakis, H., Losick, R., and Kolter, R. (2011) An accessory protein required for anchoring and assembly of amyloid fibres in B.subtilis biofilms. Mol Microbiol 80: 1155-1168.
90. Serra, D.O., Richter, A.M., Klauck, G., Mika, F., and Hengge, R. (2013) Microanatomy at cellular resolution and spatial order of physiological differentiation in a bacterial biofilm. MBio 4: e103-e113.
91. Shemesh, M., and Chai, Y. (2013) A combination of glycerol and manganese promotes biofilm formation in Bacillus subtilis via histidine kinase KinD signaling. J Bacteriol 195: 2747-2754.
92. Stoodley, P., Sauer, K., Davies, D.G., and Costerton, J.W. (2002) Biofilms as complex differentiated communities. Annu Rev Microbiol 56: 187-209.
93. Stragier, P., and Losick, R. (1996) Molecular genetics of sporulation in Bacillus subtilis. Annu Rev Genet 30: 297-341.
94. Strauch, M., Webb, V., Spiegelman, G., and Hoch, J.A. (1990) The SpoOA protein of Bacillus subtilis is a repressor of the abrB gene. Proc Natl Acad Sci USA 87: 1801-1805.
95. Strauch, M.A., Bobay, B.G., Cavanagh, J., Yao, F., Wilson, A., and Le Breton, Y. (2007) Abh and AbrB control of Bacillus subtilis antimicrobial gene expression. J Bacteriol 189: 7720-7732.
96. Sutherland, I. (2001a) Biofilm exopolysaccharides: a strong and sticky framework. Microbiology 147: 3-9.
97. Sutherland, I.W. (2001b) The biofilm matrix - an immobilized but dynamic microbial environment. Trends Microbiol 9: 222-227.
98. Sutherland, I.W. (2001c) Exopolysaccharides in biofilms, flocs and related structures. Water Sci Technol 43: 77-86.
99. Tortosa, P., Logsdon, L., Kraigher, B., Itoh, Y., Mandic-Mulec, I., and Dubnau, D. (2001) Specificity and genetic polymorphism of the Bacillus competence quorum-sensing system. J Bacteriol 183: 451-460.
100. Tran, L.S., Nagai, T., and Itoh, Y. (2000) Divergent structure of the ComQXPA quorum-sensing components: molecular basis of strain-specific communication mechanism in Bacillus subtilis. Mol Microbiol 37: 1159-1171.
101. Trejo, M., Douarche, C., Bailleux, V., Poulard, C., Mariot, S., Regeard, C., and Raspaud, E. (2013) Elasticity and wrinkled morphology of Bacillus subtilis pellicles. Proc Natl Acad Sci USA 110: 2011-2016.
102. Veening, J.W., Hamoen, L.W., and Kuipers, O.P. (2005) Phosphatases modulate the bistable sporulation gene expression pattern in Bacillus subtilis. Mol Microbiol 56: 1481-1494.
103. Veening, J.W., Igoshin, O.A., Eijlander, R.T., Nijland, R., Hamoen, L.W., and Kuipers, O.P. (2008) Transient heterogeneity in extracellular protease production by Bacillus subtilis. Mol Syst Biol 4: 184.
104. Verhamme, D.T., Kiley, T.B., and Stanley-Wall, N.R. (2007) DegU co-ordinates multicellular behaviour exhibited by Bacillus subtilis. Mol Microbiol 65: 554-568.
105. Vlamakis, H., Aguilar, C., Losick, R., and Kolter, R. (2008) Control of cell fate by the formation of an architecturally complex bacterial community. Genes Dev 22: 945-953.
106. Vlamakis, H., Chai, Y., Beauregard, P., Losick, R., and Kolter, R. (2013) Sticking together: building a biofilm the Bacillus subtilis way. Nat Rev Microbiol 11: 157-168.
107. Walker, T.S., Bais, H.P., Deziel, E., Schweizer, H.P., Rahme, L.G., Fall, R., and Vivanco, J.M. (2004) Pseudomonas aeruginosa - plant root interactions. Pathogenicity, biofilm formation, and root exudation. Plant Physiol 134: 320-331.
108. Watnick, P., and Kolter, R. (2000) Biofilm, city of microbes. J Bacteriol 182: 2675-2679.
109. Whitchurch, C.B., Tolker-Nielsen, T., Ragas, P.C., and Mattick, J.S. (2002) Extracellular DNA required for bacterial biofilm formation. Science 295: 1487.
110. Wilking, J.N., Zaburdaev, V., De Volder, M., Losick, R., Brenner, M.P., and Weitz, D.A. (2013) Liquid transport facilitated by channels in Bacillus subtilis biofilms. Proc Natl Acad Sci USA 110: 848-852.
111. Yepes, A., Schneider, J., Mielich, B., Koch, G., Garcia-Betancur, J.C., Ramamurthi, K.S., etal. (2012) The biofilm formation defect of a Bacillus subtilis flotillin-defective mutant involves the protease FtsH. Mol Microbiol 86: 457-471.
112. Zeigler, D.R., Pragai, Z., Rodriguez, S., Chevreux, B., Muffler, A., Albert, T., etal. (2008) The origins of 168, W23, and other Bacillus subtilis legacy strains. J Bacteriol 190: 6983-6995.
113. Zeriouh, H., de Vicente, A., Perez-Garcia, A., and Romero, D. (2013) Surfactin triggers biofilm formation of Bacillus subtilis in melon phylloplane and contributes to the biocontrol activity. Environ Microbiol. doi:10.1111/1462-2920.12271.