KEGG   PATHWAY: ko05111
Entry
ko05111                     Pathway                                
Name
Biofilm formation - Vibrio cholerae
Description
Surface colonization and subsequent biofilm formation and development provide numerous advantages to microorganisms. On the other hand, biofilm formation is an energetically costly process and therefore must be tightly regulated and plastic, enabling biofilm bacteria to be responsive to the various environmental cues. The quorum sensing (QS) pathway of Vibrio cholerae activates the expression of VpsR, VpsT and AphA (the main activators of biofilm formation) at low cell density and HapR (the main repressor) at high cell density. At low cell density, biofilm genes, including Vibrio polysaccharide (VPS) biosynthesis genes and the major extracellular matrix genes RbmA, RbmC, and Bap1, are expressed. The QS pathway also leads to induction of virulence factors such as toxin coregulated pilus (TCP) and cholera toxin (CTX), essential for colonization of the host and enterotoxicity, respectively. The VarS/VarA signaling system responds to an unknown environmental cue and represses biofilm production by process which involves the regulatory sRNAs CsrB, CsrC and CsrD and the RNA-binding protein CsrA, interfering with LuxO of the QS pathway. The second messenger cAMP acts as a repressor of biofilm formation. cAMP is generated by adenylate cyclase CyaA, and in complex with cAMP receptor protein (CRP), this nucleotide has been shown to upregulate HapR production through the positive regulation of the CAI-I synthase CqsA and the negative regulation of Fis. The second messenger c-di-GMP is a key signalling molecule controlling motility and biofilm matrix production. High cellular levels of c-di-GMP promote enhanced transcription of genes involved in biofilm formation, possibly by promoting VpsT-mediated transcriptional activation of vps genes.
Class
Cellular Processes; Cellular community - prokaryotes
Pathway map
ko05111  Biofilm formation - Vibrio cholerae
ko05111

Module
M00852  Vibrio cholerae pathogenicity signature, toxin coregulated pilus [PATH:ko05111]
Orthology
K00640  cysE; serine O-acetyltransferase [EC:2.3.1.30]
K01791  wecB; UDP-N-acetylglucosamine 2-epimerase (non-hydrolysing) [EC:5.1.3.14]
K01912  paaK; phenylacetate-CoA ligase [EC:6.2.1.30]
K02405  fliA, whiG; RNA polymerase sigma factor FliA
K02452  gspC; general secretion pathway protein C
K02453  gspD; general secretion pathway protein D
K02454  gspE; general secretion pathway protein E [EC:7.4.2.8]
K02455  gspF; general secretion pathway protein F
K02456  gspG; general secretion pathway protein G
K02457  gspH; general secretion pathway protein H
K02458  gspI; general secretion pathway protein I
K02459  gspJ; general secretion pathway protein J
K02460  gspK; general secretion pathway protein K
K02461  gspL; general secretion pathway protein L
K02462  gspM; general secretion pathway protein M
K02463  gspN; general secretion pathway protein N
K02472  wecC; UDP-N-acetyl-D-mannosaminuronic acid dehydrogenase [EC:1.1.1.336]
K02777  crr; sugar PTS system EIIA component [EC:2.7.1.-]
K02779  ptsG; glucose PTS system EIICB or EIICBA component [EC:2.7.1.199]
K03087  rpoS; RNA polymerase nonessential primary-like sigma factor
K03092  rpoN; RNA polymerase sigma-54 factor
K03557  fis; Fis family transcriptional regulator, factor for inversion stimulation protein
K03563  csrA; carbon storage regulator
K03606  wcaJ; undecaprenyl-phosphate glucose phosphotransferase [EC:2.7.8.31]
K03666  hfq; host factor-I protein
K05851  cyaA; adenylate cyclase, class 1 [EC:4.6.1.1]
K07173  luxS; S-ribosylhomocysteine lyase [EC:4.4.1.21]
K07181  cdgJ; c-di-GMP phosphodiesterase [EC:3.1.4.52]
K07678  barA, gacS, varS; two-component system, NarL family, sensor histidine kinase BarA [EC:2.7.13.3]
K07689  uvrY, gacA, varA; two-component system, NarL family, invasion response regulator UvrY
K08604  hap, nprV; vibriolysin [EC:3.4.24.25]
K08720  ompU; outer membrane protein OmpU
K10909  luxQ; two-component system, autoinducer 2 sensor kinase/phosphatase LuxQ [EC:2.7.13.3 3.1.3.-]
K10910  luxP; autoinducer 2-binding periplasmic protein LuxP
K10911  luxU; two-component system, phosphorelay protein LuxU
K10912  luxO; two-component system, repressor protein LuxO
K10913  hapR, luxR, litR; TetR/AcrR family transcriptional regulator, hemagglutinin/protease regulatory protein
K10914  crp; CRP/FNR family transcriptional regulator, cyclic AMP receptor protein
K10915  cqsA; CAI-1 autoinducer synthase [EC:2.3.-.-]
K10916  cqsS; two-component system, CAI-1 autoinducer sensor kinase/phosphatase CqsS [EC:2.7.13.3 3.1.3.-]
K10917  aphA; PadR family transcriptional regulator, regulatory protein AphA
K10918  aphB; LysR family transcriptional regulator, transcriptional activator AphB
K10919  tcpH; toxin coregulated pilus biosynthesis protein H
K10920  tcpP; toxin coregulated pilus biosynthesis protein P
K10921  toxR; cholera toxin transcriptional activator
K10922  toxS; transmembrane regulatory protein ToxS
K10923  tcpN, toxT; AraC family transcriptional regulator, TCP pilus virulence regulatory protein
K10924  mshA; MSHA pilin protein MshA
K10925  mshB; MSHA pilin protein MshB
K10926  mshC; MSHA pilin protein MshC
K10927  mshD; MSHA pilin protein MshD
K10928  ctxA; cholera enterotoxin subunit A [EC:2.4.2.36]
K10929  ctxB; cholera enterotoxin subunit B
K10930  tcpA; toxin coregulated pilin
K10931  tcpB; toxin coregulated pilus biosynthesis protein B
K10932  tcpC; toxin coregulated pilus biosynthesis outer membrane protein C
K10933  tcpD; toxin coregulated pilus biosynthesis protein D
K10934  tcpE; toxin coregulated pilus biosynthesis protein E
K10935  tcpF; toxin coregulated pilus biosynthesis protein F
K10936  acfA; accessory colonization factor AcfA
K10937  acfB; accessory colonization factor AcfB
K10938  acfC; accessory colonization factor AcfC
K10939  acfD; accessory colonization factor AcfD
K10940  ompT; outer membrane protein OmpT
K10941  flrA, fleQ, flaK; sigma-54 dependent transcriptional regulator, flagellar regulatory protein
K10942  flrB, fleS, flaL; two-component system, sensor histidine kinase FlrB [EC:2.7.13.3]
K10943  flrC, fleR, flaM; two-component system, response regulator FlrC
K10961  tcpI; toxin coregulated pilus biosynthesis protein I
K10962  tcpQ; toxin coregulated pilus biosynthesis protein Q
K10963  tcpR; toxin coregulated pilus biosynthesis protein R
K10964  tcpS; toxin coregulated pilus biosynthesis protein S
K10965  tcpT; toxin coregulated pilus biosynthesis protein T
K12276  mshE; MSHA biogenesis protein MshE
K13246  vieA; c-di-GMP phosphodiesterase [EC:3.1.4.52]
K15851  qrr; quorum regulatory RNA Qrr
K16554  exoP, vpsO; polysaccharide biosynthesis transport protein [EC:2.7.10.3]
K18515  csrB; small regulatory RNA CsrB
K20918  vpsT; LuxR family transcriptional regulator, positive regulator of biofilm formation
K20919  vpsR, cpsR; sigma-54 dependent transcriptional regulator, positive regulator of biofilm formation
K20920  vpsM; polysaccharide biosynthesis protein VpsM
K20921  vpsD, epsF; polysaccharide biosynthesis protein VpsD
K20922  vpsI; polysaccharide biosynthesis protein VpsI
K20945  vpsU; tyrosine-protein phosphatase [EC:3.1.3.48]
K20946  vpsE; polysaccharide biosynthesis protein VpsE
K20947  vpsF; polysaccharide biosynthesis protein VpsF
K20948  vpsJ; polysaccharide biosynthesis protein VpsJ
K20949  vpsP; polysaccharide biosynthesis protein VpsP
K20950  vpsQ; polysaccharide biosynthesis protein VpsQ
K20951  bap1; extracellular matrix protein
K20952  rbmC; rugosity and biofilm structure modulator C
K20953  rbmA; rugosity and biofilm structure modulator A
K20954  acgB; diguanylate cyclase [EC:2.7.7.65]
K20955  cdgK; diguanylate cyclase [EC:2.7.7.65]
K20956  cdgA; diguanylate cyclase [EC:2.7.7.65]
K20957  cdgH; diguanylate cyclase [EC:2.7.7.65]
K20958  cdgL; diguanylate cyclase [EC:2.7.7.65]
K20959  cdgM; diguanylate cyclase [EC:2.7.7.65]
K20960  K20960; diguanylate cyclase [EC:2.7.7.65]
K20961  vpvC; diguanylate cyclase [EC:2.7.7.65]
K20962  cdgC; c-di-GMP phosphodiesterase [EC:3.1.4.52]
K20963  acgA; c-di-GMP phosphodiesterase [EC:3.1.4.52]
K20964  cdpA; c-di-GMP phosphodiesterase [EC:3.1.4.52]
K20965  rocS; c-di-GMP phosphodiesterase [EC:3.1.4.52]
K20966  mbaA; c-di-GMP phosphodiesterase [EC:3.1.4.52]
K20988  vpsN; polysaccharide biosynthesis/export protein VpsN
K25886  vpsK; polysaccharide biosynthesis protein VpsK
Compound
C00031  D-Glucose
C00044  GTP
C00092  D-Glucose 6-phosphate
C00575  3',5'-Cyclic AMP
C16421  AI-2
C16463  3',5'-Cyclic diGMP
C16640  CAI-1
C18076  5'-Phosphoguanylyl(3'->5')guanosine
Reference
PMID:25895940
  Authors
Teschler JK, Zamorano-Sanchez D, Utada AS, Warner CJ, Wong GC, Linington RG, Yildiz FH
  Title
Living in the matrix: assembly and control of Vibrio cholerae biofilms.
  Journal
Nat Rev Microbiol 13:255-68 (2015)
DOI:10.1038/nrmicro3433
Reference
PMID:26700108
  Authors
Dang H, Lovell CR
  Title
Microbial Surface Colonization and Biofilm Development in Marine Environments.
  Journal
Microbiol Mol Biol Rev 80:91-138 (2016)
DOI:10.1128/MMBR.00037-15
Reference
PMID:19287449
  Authors
Hengge R
  Title
Principles of c-di-GMP signalling in bacteria.
  Journal
Nat Rev Microbiol 7:263-73 (2009)
DOI:10.1038/nrmicro2109
Reference
PMID:25666534
  Authors
Liang ZX
  Title
The expanding roles of c-di-GMP in the biosynthesis of exopolysaccharides and secondary metabolites.
  Journal
Nat Prod Rep 32:663-83 (2015)
DOI:10.1039/c4np00086b
Reference
PMID:9371445
  Authors
Sandkvist M, Michel LO, Hough LP, Morales VM, Bagdasarian M, Koomey M, DiRita VJ, Bagdasarian M
  Title
General secretion pathway (eps) genes required for toxin secretion and outer membrane biogenesis in Vibrio cholerae.
  Journal
J Bacteriol 179:6994-7003 (1997)
DOI:10.1128/JB.179.22.6994-7003.1997
Reference
PMID:16262799
  Authors
Lenz DH, Miller MB, Zhu J, Kulkarni RV, Bassler BL
  Title
CsrA and three redundant small RNAs regulate quorum sensing in Vibrio cholerae.
  Journal
Mol Microbiol 58:1186-202 (2005)
DOI:10.1111/j.1365-2958.2005.04902.x
Reference
PMID:17981973
  Authors
Houot L, Watnick PI
  Title
A novel role for enzyme I of the Vibrio cholerae phosphoenolpyruvate phosphotransferase system in regulation of growth in a biofilm.
  Journal
J Bacteriol 190:311-20 (2008)
DOI:10.1128/JB.01410-07
Reference
PMID:17921282
  Authors
Liang W, Silva AJ, Benitez JA
  Title
The cyclic AMP receptor protein modulates colonial morphology in Vibrio cholerae.
  Journal
Appl Environ Microbiol 73:7482-7 (2007)
DOI:10.1128/AEM.01564-07
Reference
PMID:17181781
  Authors
Lenz DH, Bassler BL
  Title
The small nucleoid protein Fis is involved in Vibrio cholerae quorum sensing.
  Journal
Mol Microbiol 63:859-71 (2007)
DOI:10.1111/j.1365-2958.2006.05545.x
Related
pathway
ko02024  Quorum sensing
ko02040  Flagellar assembly
ko03070  Bacterial secretion system
ko05110  Vibrio cholerae infection

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