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Cra DNA-binding transcriptional dual regulator

Synonyms: Cra, Cra-β-D-fructose 1,6-bisphosphate, Cra-β-D-fructofuranose 1-phosphate
Summary:
"Catabolite repressor activator," Cra, which was initially named "Fructose repressor," "FruR," is a dual transcriptional regulator that plays a pleiotropic role to modulate the direction of carbon flow through the different metabolic pathways of energy metabolism, but independently of the CRP regulator [7, 8, 13] For this reason the FruR regulator is implicated in the expression of a large number of operons that encode enzymes which comprise central pathways of carbon metabolism. FruR acts as an activator of genes encoding gluconeogenic [7, 24] Krebs cycle [] and glyoxylate shunt [1]enzymes, and the negative effect on genes encoding Entner-Doudoroff pathway [13]and glycolytic enzymes [7, 24] Cra carries out a glycolytic flux-dependent regulation.
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Transcription factor      
TF conformation(s):
Name Conformation Type TF-Effector Interaction Type Apo/Holo Conformation Evidence (Confirmed, Strong, Weak) References
Cra Functional   Apo [APPH], [BPP], [GEA], [HIFS] [1], [2], [3], [4], [5], [6], [7], [8], [9]
Cra-β-D-fructofuranose 1-phosphate Non-Functional Allosteric Holo [BPP], [GEA] [7], [8]
Cra-β-D-fructose 1,6-bisphosphate Non-Functional Allosteric Holo [BPP], [GEA] [7], [8]
Evolutionary Family: GalR/LacI
Sensing class: Using internal synthesized signals
Connectivity class: Local Regulator
Gene name: cra
  Genome position: 88028-89032
  Length: 1005 bp / 334 aa
Operon name: cra
TU(s) encoding the TF:
Transcription unit        Promoter
cra
fruRp8


Regulon       
Regulated gene(s) aceA, aceB, aceE, aceF, aceK, acnA, acnB, adhE, aroP, betA, betB, betI, betT, crp, crr, csgD, csgE, csgF, csgG, cydA, cydB, cyoA, cyoB, cyoC, cyoD, cyoE, cysG, eda, edd, eno, envC, epd, fbaA, fbaB, fruA, fruB, fruK, gapA, glcC, glk, gpmM, hypF, icd, lpd, manX, manY, manZ, marA, marB, marR, mpl, mtlA, mtlD, mtlR, nirB, nirC, nirD, pck, pdeL, pdhR, pfkA, pgk, poxB, ppc, ppsA,
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Multifun term(s) of regulated gene(s) MultiFun Term (List of genes associated to the multifun term)
carbon compounds (26)
membrane (19)
Transcription related (8)
repressor (8)
Phosphotransferase Systems (PEP-dependent PTS) (7)
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Regulated operon(s) aceBAK, acnA, acnB, adhE, aroP, betIBA, betT, crp, csgDEFG, cydAB, cyoABCDE, edd-eda, epd-pgk-fbaA, fbaB, fruBKA, gapA-yeaD, glcC, glk, gpmM-envC-yibQ, hcp-hcr-poxB-ltaE-ybjT, hydN-hypF, icd, manXYZ, marRAB, mpl, mtlADR, nirBDC-cysG, pck, pdeL, pdhR-aceEF-lpd, pfkA, ppc, ppsA, prpBCDE, ptsHI-crr, pykF, pyrG-eno, sgrST-setA, tpiA, zwf
First gene in the operon(s) aceB, acnA, acnB, adhE, aroP, aroP, betI, betT, crp, csgD, cydA, cyoA, edd, eno, eno, eno, epd, epd, fbaB, fruB, gapA, gapA, gapA, gapA, glcC, glk, gpmM, hypF, hypF, icd, manX, marR, mpl, mtlA, nirB, pck, pdeL, pdhR, pfkA, poxB, ppc, ppsA, prpB, ptsH, ptsH, pykF, sgrS, tpiA, zwf
Simple and complex regulons AcrR,CRP,CpxR,Cra,Fis,MarA,MarR,Rob,SoxS
ArcA,BetI,Cra
ArcA,CRP,Cra,CusR,FNR,Fis,Fur,GadE,HprR,PdhR
ArcA,CRP,Cra,FNR,MarA,Rob,SoxS
ArcA,CRP,Cra,Fis
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Simple and complex regulatory phrases Regulatory phrase (List of promoters regulated by the phrase)
[Cra,+](10)
[Cra,-](33)


Transcription factor regulation    


Transcription factor binding sites (TFBSs) arrangements
      

  Functional conformation Function Promoter Sigma factor Central Rel-Pos Distance to first Gene Genes Sequence LeftPos RightPos Evidence (Confirmed, Strong, Weak) References
  Cra activator aceBp Sigma70 -166.5 -242.5 aceB, aceA, aceK
tgatagtcgaTCGTTAAGCGATTCAGCAccttacctca
4215227 4215244 [BPP], [GEA], [SM] [1], [8], [10]
  Cra activator acnAp2 Sigma70 nd -1335831.0 acnA nd nd [GEA] [11]
  Cra repressor acnBp Sigma70 nd -131615.0 acnB nd nd [GEA] [11]
  Cra repressor adhEp1 Sigma38, Sigma70 36.5 -257.0 adhE
tttttgcaaaGCTGACACCTTTCAGCatcgcttttc
1298370 1298386 [BCE], [GEA], [ICWHO], [SM] [4], [12], [13]
  Cra activator aroPp1 Sigma70 -177.5 -297.5 aroP
taaaaatccgAATTGAACCGATTCACTTaccaattttg
121840 121857 [AIBSCS], [GEA] [13], [14]
  Cra activator aroPp2 Sigma70 -198.5 -297.5 aroP
taaaaatccgAATTGAACCGATTCACTTaccaattttg
121840 121857 [AIBSCS], [GEA], [ICWHO] [13], [14]
  Cra activator betIp Sigma70 nd 329334.0 betI, betB, betA nd nd [GEA] [10]
  Cra activator betTp Sigma70 nd -329463.0 betT nd nd [GEA] [10]
  Cra activator crpp1 Sigma70 -45.5 -212.0 crp
cctgggtcatGCTGAAGCGAGACACCaggagacaca
3485900 3485916 [BPP], [GEA], [SM] [15]
  Cra activator csgDp1 Sigma38, Sigma70 -563.5 -711.5 csgD, csgE, csgF, csgG
gttaacaaccAAGTTGAAATGATTTAATttcttaaatg
1103899 1103916 [AIBSCS], [BPP], [GEA] [16]
  Cra activator csgDp1 Sigma38, Sigma70 -227.5 -375.5 csgD, csgE, csgF, csgG
cctttcgtcaTGTAAAACGTTCTTGTTTtttctccaca
1103563 1103580 [AIBSCS], [BPP], [GEA], [SM] [16]
  Cra activator csgDp1 Sigma38, Sigma70 -196.5 -344.5 csgD, csgE, csgF, csgG
ataatgcataTAAAAAATATTTCGGTGTagtcctttcg
1103532 1103549 [AIBSCS], [BPP], [GEA], [SM] [16]
  Cra activator csgDp1 Sigma38, Sigma70 46.5 -102.5 csgD, csgE, csgF, csgG
actgctgtgtGTAGTAATAAATCAGCCCtaaatgggta
1103290 1103307 [AIBSCS], [BPP], [GEA], [SM] [16]
  Cra activator cydAp5 Sigma70 -134.5 -224.5 cydA, cydB
cgtcgaaaaaTGCAAATTTGCTTCAACAaaaacctgtt
771225 771242 [AIBSCS], [BPP], [GEA] [17]
  Cra activator cydAp5 Sigma70 -105.5 -195.5 cydA, cydB
aaacctgtttATTGTAAGGATTTTGCGGcgtaatatat
771254 771271 [AIBSCS], [BPP], [GEA] [17]
  Cra repressor cyoAp Sigma70 -268.5 -311.5 cyoA, cyoB, cyoC, cyoD, cyoE
aagactcattAGCTGAATCGTGTAAGAAaaatagaatt
451913 451930 [AIBSCS], [GEA] [14]
  Cra repressor eddp1 Sigma70 4.5 -105.5 edd, eda
gagctcatagGGCAAAAACGTTTCAGTCagtgtaataa
1934701 1934718 [BPP], [GEA], [HIBSCS], [SM] [7]
  Cra repressor enop1 nd -205.5 -938.5 eno
gggcagtttaAGCTGAATCGTTTACAAAtataatcgtc
2908871 2908888 [BPP], [GEA], [HIBSCS] [18]
  Cra repressor enop1 nd -8.5 -741.5 eno
tttcaacatcTTGTGAATCGATCAGTTTgatgttgacg
2908674 2908691 [BPP], [GEA] [18]
  Cra repressor enop2 nd -177.5 -938.5 eno
gggcagtttaAGCTGAATCGTTTACAAAtataatcgtc
2908871 2908888 [BPP], [GEA], [HIBSCS] [18]
  Cra repressor enop2 nd 20.5 -741.5 eno
tttcaacatcTTGTGAATCGATCAGTTTgatgttgacg
2908674 2908691 [BPP], [GEA] [18]
  Cra repressor enop3 nd -33.5 -938.5 eno
gggcagtttaAGCTGAATCGTTTACAAAtataatcgtc
2908871 2908888 [BPP], [GEA], [HIBSCS] [18]
  Cra repressor enop3 nd 164.5 -741.5 eno
tttcaacatcTTGTGAATCGATCAGTTTgatgttgacg
2908674 2908691 [BPP], [GEA] [18]
  Cra repressor epdp Sigma70, Sigma38 3.5 -129.5 epd, pgk, fbaA
acatttaatcGACTGAAACGCTTCAGCTaggataagcg
3073812 3073829 [BPP], [GEA], [HIBSCS], [SM] [7]
  Cra repressor fbaBp Sigma38 -35.5 -138.5 fbaB
cagcgttttcTGTTGGCTCGATTCATCAgaaaaaatgt
2178694 2178711 [AIBSCS], [GEA] [14]
  Cra repressor fruBp Sigma70 7.5 -95.5 fruB, fruK, fruA
acaagatcgcGCTGAAACGTTTCAAGAAagcataatac
2263582 2263599 [AIBSCS], [GEA], [NTAS] [8], [14]
  Cra repressor fruBp Sigma70 73.5 -29.5 fruB, fruK, fruA
tcttgctgaaTTGAAACGATTCAGCCTCtatgagaaaa
2263516 2263533 [AIBSCS], [GEA] [14]
  Cra repressor gapAp1 Sigma70, Sigma38 -725.5 -761.5 gapA
tggaataagtTGCTGAATCGTTTTTTCAaccgttgatt
1862001 1862018 [BPP], [GEA], [HIBSCS] [18]
  Cra repressor gapAp1 Sigma70, Sigma38 -43.5 -79.5 gapA
ttgtcgcaatGATTGACACGATTCCGCTtgacgctgcg
1862683 1862700 [AIBSCS], [BPP], [GEA] [18]
  Cra repressor gapAp2 Sigma32 -608.5 -761.5 gapA, yeaD
tggaataagtTGCTGAATCGTTTTTTCAaccgttgatt
1862001 1862018 [BPP], [GEA], [HIBSCS] [18]
  Cra repressor gapAp2 Sigma32 74.5 -79.5 gapA, yeaD
ttgtcgcaatGATTGACACGATTCCGCTtgacgctgcg
1862683 1862700 [AIBSCS], [BPP], [GEA] [18]
  Cra repressor gapAp3 Sigma70 -587.5 -761.5 gapA
tggaataagtTGCTGAATCGTTTTTTCAaccgttgatt
1862001 1862018 [BPP], [GEA], [HIBSCS] [18]
  Cra repressor gapAp3 Sigma70 95.5 -79.5 gapA
ttgtcgcaatGATTGACACGATTCCGCTtgacgctgcg
1862683 1862700 [AIBSCS], [BPP], [GEA] [18]
  Cra repressor gapAp4 Sigma70 -516.5 -761.5 gapA
tggaataagtTGCTGAATCGTTTTTTCAaccgttgatt
1862001 1862018 [BPP], [GEA], [HIBSCS] [18]
  Cra repressor gapAp4 Sigma70 166.5 -79.5 gapA
ttgtcgcaatGATTGACACGATTCCGCTtgacgctgcg
1862683 1862700 [AIBSCS], [BPP], [GEA] [18]
  Cra repressor glcCp Sigma70 -4.5 -70.5 glcC
aattgatgtaACATAATCACTTACGTGAtgtgcgtgtt
3128193 3128210 [AIBSCS], [GEA] [14]
  Cra repressor glkp Sigma70, Sigma38 -91.5 -126.5 glk
gtcacacaatTACTTTATCGTTTCAGCAccaattgcag
2509544 2509561 [AIBSCS], [GEA], [HIBSCS] [14], [19]
  Cra repressor gpmMp nd -31.5 -56.5 gpmM, envC, yibQ
cttcagaggcTATTTTATCGATTCAGCTgtagtaaaat
3785195 3785212 [AIBSCS], [GEA] [14]
  Cra repressor hypFp2 Sigma70 -84.5 -673.5 hypF
gcgtcagcaaTGATGAAACGGTTCATCAaattctccag
2838090 2838107 [BPP], [GEA], [HIBSCS] [18]
  Cra repressor hypFp3 Sigma70 7.5 -673.5 hypF
gcgtcagcaaTGATGAAACGGTTCATCAaattctccag
2838090 2838107 [BPP], [GEA], [HIBSCS] [18]
  Cra activator icdAp2 Sigma70 -76.5 -238.5 icd
gaacgttgcgAGCTGAATCGCTTAACCTggtgatttct
1194876 1194893 [BPP] [20]
  Cra repressor manXp Sigma70 40.5 -75.0 manX, manY, manZ
tgtagcccttATCTGAATCGATTCGAttgtggacga
1901965 1901981 [AIBSCS], [GEA] [13]
  Cra repressor marRp Sigma70 -72.5 -99.5 marR, marA, marB
attgaacaaaACTTGAACCGATTTAGCAaaacgtggca
1619012 1619029 [AIBSCS], [GEA] [14]
  Cra repressor mplp1 Sigma38, Sigma70 -36.5 -93.5 mpl
tgttgcgctcAGGTGAATCGCGCCAGCAaattacggat
4455683 4455700 [AIBSCS], [GEA] [14]
  Cra repressor mtlAp Sigma70 -22.5 -115.5 mtlA, mtlD, mtlR
gtcgactggaCAGTTAACCGATTCAGTGccagatttcg
3772157 3772174 [BPP], [GEA] [7]
  Cra repressor nirBp Sigma70 -15.5 -39.5 nirB, nirD, nirC, cysG
taagcggggtTGCTGAATCGTTAAGGTAggcggtaata
3493963 3493980 [BCE], [GEA], [HIBSCS] [21], [22]
  Cra activator pckp Sigma70 -139.5 -278.5 pck
aaagttagcgTGGTGAATCGATACTTTAccggttgaat
3532531 3532548 [BPP], [GEA], [HIBSCS], [SM] [7]
  Cra repressor pdeLp1 Sigma70 -159.5 -491.5 pdeL
ggttatcattTGCTGAATGGATTCAGTCttaatgagtg
331871 331888 [BPP], [GEA], [HIBSCS] [18]
  Cra repressor pdeLp1 Sigma70 -13.5 -345.5 pdeL
cagttgatctGGATTGTTAAATTCATATaatgcgcctt
332017 332034 [AIBSCS], [GEA] [18]
  Cra repressor pdhRp Sigma70, Sigma38 -185.5 -243.5 pdhR, aceE, aceF, lpd
caaaattggtAAGTGAATCGGTTCAATTcggattttta
121840 121857 [AIBSCS], [GEA] [13], [14]
  Cra repressor pfkAp2 Sigma70, Sigma38 -35.5 -113.5 pfkA
atttggcctgACCTGAATCAATTCAGCAggaagtgatt
4107430 4107447 [BPP] [5]
  Cra activator poxBp Sigma38, Sigma70 nd 911049.0 poxB nd nd [GEA] [10]
  Cra repressor ppcp nd -46.5 -138.5 ppc
cgtcaaatgcTGCGAAATCGCTTCAGCAaacgaataaa
4153228 4153245 [AIBSCS], [GEA] [14]
  Cra activator ppsp Sigma70 -45.5 -95.0 ppsA
tgaaaaaaacGGTGAATCGTTCAAGCaaatatattt
1787199 1787215 [AIBSCS], [BPP], [GEA], [ICWHO] [8], [10], [13], [14]
  Cra repressor prpBp Sigma54 -101.5 -137.5 prpB, prpC, prpD, prpE
ctgaaacgttAACTGAAACGCATATTTGcggattagtt
348536 348553 [AIBSCS], [GEA] [14]
  Cra repressor ptsHp1 nd -55.5 -211.5 ptsH, ptsI, crr
gaaaagccaaAGCTGAATCGATTTTATGatttggttca
2533544 2533561 [AIBSCS], [BPP], [GEA], [HIBSCS] [14], [23]
  Cra activator ptsHp2 Sigma70 -48.5 -211.5 ptsH, ptsI, crr
gaaaagccaaAGCTGAATCGATTTTATGatttggttca
2533544 2533561 [AIBSCS], [BPP], [GEA], [HIBSCS] [14], [23]
  Cra repressor pykFp Sigma70 18.5 -213.5 pykF
gccaattgacTCTTGAATGGTTTCAGCActttggactg
1755476 1755493 [BCE], [SM] [24]
  Cra repressor sgrSp Sigma70, Sigma38 -332.5 -332.5 sgrS, sgrT, setA
tgcaccagttGATCGATACGATCCTGCTccagcaggtc
77026 77043 [AIBSCS], [GEA] [14]
  Cra repressor tpiAp2 Sigma70, Sigma38 -91.5 -153.5 tpiA
agatagcgccAGCTTAATCGGTTCAACAgcgaaggtca
4111652 4111669 [AIBSCS], [GEA] [14]
  Cra repressor zwfp Sigma70 -35.5 -97.5 zwf
tttacgagctTGCGAAAACTGTAAACGCttatccaccc
1936403 1936420 [AIBSCS], [GEA] [14]


Alignment and PSSM for Cra TFBSs    

Aligned TFBS of Cra   
  Sequence
  ATCGATACGATCCTGC
  ATTGAACCGATTCACT
  ACTGAATCCATTCAGC
  AATTTAACAATCCAGA
  AAATATGCGTTTCAGT
  TCTTACACGATTCAGC
  GTTGAAGCAAATTTGC
  ATTGTAAGGATTTTGC
  GTAGTAATAAATCAGC
  ACCGAAATATTTTTTA
  ACAAGAACGTTTTACA
  ATTAAATCATTTCAAC
  GGTTAAGCGATTCAGC
  GCTGACACCTTTCAGC
  CTTGAACCGATTTAGC
  CTTGAATGGTTTCAGC
  GCTTGAACGATTCACC
  GAAAAAACGATTCAGC
  ATTGACACGATTCCGC
  ATCGAATCGATTCAGA
  ACTGAAACGTTTTTGC
  CGTTTACAGTTTTCGC
  GTTGGCTCGATTCATC
  ATTGAAACGATTCAGC
  CTTGAAACGTTTCAGC
  ACTTTATCGTTTCAGC
  ATAAAATCGATTCAGC
  GATGAACCGTTTCATC
  AACTGATCGATTCACA
  TTGTAAACGATTCAGC
  ACTGAAACGCTTCAGC
  ACATAATCACTTACGT
  GGTGTCTCGCTTCAGC
  ACCTTAACGATTCAGC
  AAAGTATCGATTCACC
  AGTTAACCGATTCAGT
  ATTTTATCGATTCAGC
  CCTGAATCAATTCAGC
  GTTGAACCGATTAAGC
  GCTGAAGCGATTTCGC
  CGTTAAGCGATTCAGC
  GCTGGCGCGATTCACC

Position weight matrix (PWM).   
A	22	6	7	4	28	34	17	1	7	28	2	0	2	33	1	5
C	6	14	5	0	0	6	6	37	2	3	0	2	32	4	6	33
G	12	5	1	24	5	0	6	2	33	0	0	0	0	0	32	0
T	2	17	29	14	9	2	13	2	0	11	40	40	8	5	3	4

PWM logo   


 


Evolutionary conservation of regulatory elements    
     Note: Evolutionary conservation of regulatory interactions and promoters is limited to gammaproteobacteria.
TF-target gene evolutionary conservation
Promoter-target gene evolutionary conservation


Evidence    

 [APPH] Assay of protein purified to homogeneity

 [BPP] Binding of purified proteins

 [GEA] Gene expression analysis

 [HIFS] Human inference of function from sequence

 [SM] Site mutation

 [BCE] Binding of cellular extracts

 [ICWHO] Inferred computationally without human oversight

 [AIBSCS] Automated inference based on similarity to consensus sequences

 [HIBSCS] Human inference based on similarity to consensus sequences

 [NTAS] Non-traceable author statement



Reference(s)    

 [1] Cortay JC., Negre D., Scarabel M., Ramseier TM., Vartak NB., Reizer J., Saier MH., Cozzone AJ., 1994, In vitro asymmetric binding of the pleiotropic regulatory protein, FruR, to the ace operator controlling glyoxylate shunt enzyme synthesis., J Biol Chem 269(21):14885-91

 [2] Jahreis K., Postma PW., Lengeler JW., 1991, Nucleotide sequence of the ilvH-fruR gene region of Escherichia coli K12 and Salmonella typhimurium LT2., Mol Gen Genet 226(1-2):332-6

 [3] Leclerc G., Noel G., Drapeau GR., 1990, Molecular cloning, nucleotide sequence, and expression of shl, a new gene in the 2-minute region of the genetic map of Escherichia coli., J Bacteriol 172(8):4696-700

 [4] Mikulskis A., Aristarkhov A., Lin EC., 1997, Regulation of expression of the ethanol dehydrogenase gene (adhE) in Escherichia coli by catabolite repressor activator protein Cra., J Bacteriol 179(22):7129-34

 [5] Negre D., Bonod-Bidaud C., Geourjon C., Deleage G., Cozzone AJ., Cortay JC., 1996, Definition of a consensus DNA-binding site for the Escherichia coli pleiotropic regulatory protein, FruR., Mol Microbiol 21(2):257-66

 [6] Penin F., Geourjon C., Montserret R., Bockmann A., Lesage A., Yang YS., Bonod-Bidaud C., Cortay JC., Negre D., Cozzone AJ., Deleage G., 1997, Three-dimensional structure of the DNA-binding domain of the fructose repressor from Escherichia coli by 1H and 15N NMR., J Mol Biol 270(3):496-510

 [7] Ramseier TM., Bledig S., Michotey V., Feghali R., Saier MH., 1995, The global regulatory protein FruR modulates the direction of carbon flow in Escherichia coli., Mol Microbiol 16(6):1157-69

 [8] Ramseier TM., Negre D., Cortay JC., Scarabel M., Cozzone AJ., Saier MH., 1993, In vitro binding of the pleiotropic transcriptional regulatory protein, FruR, to the fru, pps, ace, pts and icd operons of Escherichia coli and Salmonella typhimurium., J Mol Biol 234(1):28-44

 [9] Yura T., Mori H., Nagai H., Nagata T., Ishihama A., Fujita N., Isono K., Mizobuchi K., Nakata A., 1992, Systematic sequencing of the Escherichia coli genome: analysis of the 0-2.4 min region., Nucleic Acids Res 20(13):3305-8

 [10] Son YJ., Phue JN., Trinh LB., Lee SJ., Shiloach J., 2011, The role of Cra in regulating acetate excretion and osmotic tolerance in E. coli K-12 and E. coli B at high density growth., Microb Cell Fact 10:52

 [11] Cunningham L., Gruer MJ., Guest JR., 1997, Transcriptional regulation of the aconitase genes (acnA and acnB) of Escherichia coli., Microbiology 143 ( Pt 12):3795-805

 [12] Kaga N., Umitsuki G., Clark DP., Nagai K., Wachi M., 2002, Extensive overproduction of the AdhE protein by rng mutations depends on mutations in the cra gene or in the Cra-box of the adhE promoter., Biochem Biophys Res Commun 295(1):92-7

 [13] Sarkar D., Siddiquee KA., Arauzo-Bravo MJ., Oba T., Shimizu K., 2008, Effect of cra gene knockout together with edd and iclR genes knockout on the metabolism in Escherichia coli., Arch Microbiol 190(5):559-71

 [14] Shimada T., Yamamoto K., Ishihama A., 2011, Novel Members of the Cra Regulon Involved in Carbon Metabolism in Escherichia coli., J Bacteriol 193(3):649-59

 [15] Zhang Z., Aboulwafa M., Saier MH., 2014, Regulation of crp Gene Expression by the Catabolite Repressor/Activator, Cra, in Escherichia coli., J Mol Microbiol Biotechnol 24(3):135-41

 [16] Reshamwala SM., Noronha SB., 2011, Biofilm formation in Escherichia coli cra mutants is impaired due to down-regulation of curli biosynthesis., Arch Microbiol 193(10):711-22

 [17] Ramseier TM., Chien SY., Saier MH., 1996, Cooperative interaction between Cra and Fnr in the regulation of the cydAB operon of Escherichia coli., Curr Microbiol 33(4):270-4

 [18] Shimada T., Fujita N., Maeda M., Ishihama A., 2005, Systematic search for the Cra-binding promoters using genomic SELEX system., Genes Cells 10(9):907-18

 [19] Meyer D., Schneider-Fresenius C., Horlacher R., Peist R., Boos W., 1997, Molecular characterization of glucokinase from Escherichia coli K-12., J Bacteriol 179(4):1298-306

 [20] Prost JF., Negre D., Oudot C., Murakami K., Ishihama A., Cozzone AJ., Cortay JC., 1999, Cra-dependent transcriptional activation of the icd gene of Escherichia coli., J Bacteriol 181(3):893-8

 [21] Browning DF., Cole JA., Busby SJ., 2004, Transcription activation by remodelling of a nucleoprotein assembly: the role of NarL at the FNR-dependent Escherichia coli nir promoter., Mol Microbiol 53(1):203-15

 [22] Tyson K., Busby S., Cole J., 1997, Catabolite regulation of two Escherichia coli operons encoding nitrite reductases: role of the Cra protein., Arch Microbiol 168(3):240-4

 [23] Ryu S., Ramseier TM., Michotey V., Saier MH., Garges S., 1995, Effect of the FruR regulator on transcription of the pts operon in Escherichia coli., J Biol Chem 270(6):2489-96

 [24] Bledig SA., Ramseier TM., Saier MH., 1996, Frur mediates catabolite activation of pyruvate kinase (pykF) gene expression in Escherichia coli., J Bacteriol 178(1):280-3



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