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. 2012 May 16;21(7):929–948. doi: 10.1002/pro.2093

Table I.

Representative High-Resolution Structures of c-di-GMP Metabolizing Enzymes and Protein Receptors

Protein Organism Function Conserved domains PDB code and nucleotide-bound state References
C-di-GMP turnover enzymes
 Diguanylate cyclases
  PleD Caulobacter crescentus C-di-GMP synthesis; polar differentiation Two CheY-like REC domains; 1W25 (c-di-GMP/Mg complex); 41,44
GGDEF domain 2WB4 (c-di-GMP/ Mg/BeF3 complex);
2V0N (c-di-GMP/Mg/BeF3/ GTP-α-S complex)
  WspR Pseudomonas aeruginosa C-di-GMP synthesis; biofilm formation and persistence CheY-like REC domain; 3BRE (c-di-GMP/Mg complex); 39,45
GGDEF domain 3I5B (GGDEF domain; apoprotein)
 Phosphodiesterases
  YkuI Bacillus subtilis C-di-GMP hydrolysis EAL domain; 2BAS (apoprotein) 46
PAS-like domain 2W27 (c-di-GMP/Ca complex)
  BlrP1 Klebsiella pneumoniae Light-regulated c-di-GMP hydrolysis BLUF sensor domain; 3GFX 3GG1 (c-di-GMP/ Ca/FMN complex); 47
EAL domain 3GFY (c-di-GMP/FMN complex);
3GFZ, 3GG0 (c-di-GMP/ Mn/FMN complex)
  Bd1817 Bdellovibrio bacteriovorus Unknown HD-GYP domain 3TM8, 3TMB, 3TMC, 3TMD (apoprotein) 48
C-di-GMP protein receptors
 PilZ domain proteins
  VCA0042/PlzD Vibrio cholerae Potential role in virulence YcgR-N* domain; 1YLN (apoprotein); 28
PilZ domain 2RDE (c-di-GMP complex)
  PP4397 Pseudomonas putida unknown YcgR-N* domain; 3KYF (c-di-GMP complex) 31
PilZ domain
  PA4608 Pseudomonas aeruginosa Potential role in biofilm formation and/or quorum sensing PilZ domain 1YWU (apoprotein); 30,32
2L74 (c-di-GMP complex)
 Transcription factors
  VpsT Vibrio cholerae Global transcription control; biofilm formation Noncanonical REC domain; 3KLN (apoprotein); 37
HTH DNA-binding domain 3KLO (c-di-GMP complex)
  Clp Xanthomonas campestris Global transcription control; virulence cNMP-binding domain; 3IWZ (apoprotein) 40
HTH DNA-binding domain
 Degenerate EAL domains
  FimX Pseudomonas aeruginosa Type IV pilus-mediated twitching motility CheY-like REC domain; 3HV9 (EAL domain; apoprotein); 35
PAS domain; GGDEF domain; EAL domain 3HV8 (EAL domain; c-di-GMP complex);
3HVA (GGDEF domain);
3HVB (GGDEF-EAL module; apoprotein)
  LapD Pseudomonas fluorescens Adhesin maintenance; biofilm formation and dispersal Periplasmic output domain; 3PJV (periplasmic domain); 49
HAMP domain; GGDEF domain; EAL domain 3PJT, 3PJU (EAL domain; c-di-GMP complex);
3PJW, 3PJX (GGDEF-EAL module; apoprotein)
 Others
  PNPase Escherichia coli O2-dependent RNA degradation Two RNase PH domains; 3CDI (apoprotein); 50
KH RNA-binding domain; 3CDJ (PH1-PH2 module; apoprotein)
S1 RNA-binding domain

Conserved domains responsible for c-di-GMP synthesis, degradation, or recognition are underlined.