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. 2014 Mar 6;9(3):e90447. doi: 10.1371/journal.pone.0090447

Table 1. Constitutive Promoters (Type-A Spacers).

Map Promoter sequence Right Operon Left Gene D RpoD D Right Gene Left Operon Promoter sequence
0.87 TTAACG-16-AATAAT caiTABCDE caiT < HNS > fixA fixABCX GTGACA-17-TAAAGT
1.82 TTGATT-18-TGAAAT leuLABCD leuL < > leuO* leuO TTCGCA-17-TATTAT
2.43 TTGTCT-18-CATAAA coaED coaE < > guaC guaC TTTATA-18-GATTAT
2.99 ATGACG-18-TATAAT gcd gcd < HNS > hpt hpt TTAATA-18-TATAGG
3.06 TTTAAA-17-TATATT can can < > yadG yadGH
5.16 ATAACA-17-GATATT yafV yafV < > ivy ivy TTGGAA-17-TATCGT
5.19 TTGCTA-17-TACAAC fadE fadE < > lpcA lpcA CTGACC-17-TGTAGT
5.31 TTGTCG-17-TACAAT dinJ-yafQ dinJ < HNS > yafL yafL TTTACA-17-TATGTT
5.57 TTAAGA-17-TATATC phoE phoE < HNS > proB proBA TTGTTA-17-AATAAT
5.99 CTGCCA-16-TATGAT insAB-2 insA2 < > insI-1 insI-1
6.04 TCTACA-17-TTTATT yagA--ykgN yagA* < > yagE yagEF TTTACC-16-CATAGT
6.22 TTGTGA-16-AATAAT argF argF < > ykgS ykgS TTGAAT-18-TGTAAT
6.25 CTGCCA-16-TATGAT insAB-3 insA3 < > yagJ yagJ
6.36 TTTATA-16-TATAGC intF intF < > ptwF ptwF TTGCTA-18-TATAAA
7.07 TTGAAC-17-TTTAAT betIBA betI* < HNS > betT betT TGGACG-17-CTTAAT
7.43 yahN yahN < > yahO yahO
8.26 TTGACA-18-TACAAT yaiS yaiS < HNS > tauA tauABCD TTGAGA-16-TACAAT
11.08 qmcA-ybbJ qmcA < > ybbL ybbLM
12.53 TTGATT-17-TATAAC ybcY ybcY < HNS > tfaX tfaX ATGGCA-15-TTAAAT
13.05 TTGCCT-17-TACCAT ybdK ybdK < > sokE sokE ATGGCA-16-AAGAAT
13.14 TTGATT-18-TATTAT fepA fepA < HNS > fes fes- -entFE AAGACA-17-AATAAT
13.37 TCGATA-17-TATCAT fepDGC fepD < > entS entS ATGAAA-17-TCTTAT
13.42 ATGATA-17-TATCAT fepB fepB < > entC entCEBAH ATGATA-17-TATCAT
13.73 ATGACA-16-TTTACT dsbG dsbG < > ahpC ahpCF TTGCCC-17-TGTAAT
14.09 ATGACA-17-AATTAT dcuC dcuC < HNS > pagP pagP TTAAGA-17-TAAAAA
14.54 CTGACG-17-AATAAG ybeQ ybeQ < HNS > ybeR ybeR-djiB
15.13 TTTACT-17-TATTTT nagB nagB < > nagE nagE GTGACA-19-TTTAAT
15.69 ATGAAA-16-TGTAAT kdpFABC kdpF < HNS > ybfA ybfA TTCGCA-17-TGTAAA
16.22 TTGACA-18-TACAAA gltA gltA < HNS > sdhC sdhCDAB TTGTAA-17-TATACT
16.46 TTGAAA-18-TATTAT mngR mngR* < HNS > mngA mngAB TTAATA-19-GATAAA
17.93 CTGAAA-18-TATTGT ybiA ybiA < > dinG dinG ATGCCA-16-TACAGT
18.29 TTCAAA-18-TATATC rhtA rhtA < HNS > ompX ompX TTGACT-18-TGGAAT
18.37 TTAACG-16-TATAAT mntS mntS < > mntR* mntR-ybiR ATTACA-18-TATATT
19.13 TTGATA-19-TAAAAT ybjL ybjL < HNS > ybjM ybjM TTGAAA-15-GATAAT
21.47 ATGACA-17-AATAAT ssuEADCB ssuE < > elfA elfAD TTTAAA-17-TATTAT
21.98 TTAACT-16-TATTCT sulA sulA < HNS > sxy sxy TTGCCC-17-TATTTT
22.59 TGGAGA-18-TACACT gfcA gfcA < > insA4 insAB-4 CTGCCA-16-TATGAT
23.75 TTATCA-17-TAAAAT csgDEFG csgD* < HNS > csgB csgBAC CTGACA-17-TGTAGT
23.86 TGGATA-17-CAGAAT mdoC mdoC < > mdoG mboGH GTGAAA-17-CTTAAT
24.03 TTAACA-16-TACATT waaM waaM < > yceA yceA TTGCCG-15-AATAAT
25.16 TTTATA-17-TAAAAA comR comR* < HNS > bhsA bhsA TTCACC-17-AATAAT
25.79 TTGTAA-18-TATAAT ymfED ymfE < > lit lit TTGATA-15-AATAAT
26.15 TTTGCA-17-TAAAAC bluF bluF* < HNS > ycgZ ycgZ-ymgC TTGTCA-18-TTTTAT
26.43 TTTACC-17-TGGAAT pliG pliG < > ycgL ycgLM TTCAGA-18-TATAAT
26.48 ATGACA-19-TATCAT hlyE hlyE < > umuD umuDC CTGGCA-16-TATAAA
26.56 CTGACC-19-CATAAA nhaB nhaB < > fadR* fadR TTGATA-19-GAAAAT
26.62 TAGATA-17-TCTAAT ycgB ycgB < > dadA dadAX TTATCA-17-TATTCT
26.79 ycgR ycgR < > ymgE ymgE TTGCCT-16-TGTAAG
26.92 TTGAAC-16-CGTAAT dhaKLM dhaK < > dhaR* dhaR TTGCGA-17-CATAAG
27.10 TTGTCA-17-TAAACT pth-ychF pth < > ychH ychH TTGTAA-17-CATAAC
27.37 TCGACA-16-TAAAAT chaA chaA < > chaB chaBC TTGACC-18-TGTAAA
27.84 ATGAAA-17-TATCAT hns hns* < > tdk tdk TTACCA-17-TATAAC
27.90 GTGACG-16-TCTAAT adhE adhE < > ychE ychE TTTAAA-16-TCTAAT
29.94 ATGATA-19-TCTAAT ycjY-ymjDC ycjY < > pgrR* pgrR TAGACC-18-TATCAT
31.03 TTGAAG-17-CATAGT ldhA ldhA < > ydbH ydbH--ydbL TTGAAA-15-AAAAAT
31.90 azoR azoR < > hrpA hrpA TAGAAA-17-TATATC
32.07 TTAACA-19-TTTCAT gapC gapC < HNS > cybB cybB TTGAGA-19-GAAAAT
32.16 TTAATA-17-TATCAT ydcI ydcI* < HNS > ydcJ ydcJ CTGACA-16-TATGAT
32.48 TAAACA-18-TGTAAT yncJ yncJ < > hicA hicAB TAAACA-19-TATACT
33.50 TTGCCA-15-TATAAA bdm-sra bdm < HNS > osmC osmC TTGATA-17-TATATT
34.73 TTTAAA-17-TATCTT sad sad < > yneJ* yneJ TTTACT-17-AATAAT
34.84 CTGGCA-17-GATAAT marC marC < > marR* marRAB TTGACT-17-TATTAT
35.08 ATGCCA-17-TTTAAG ydfI ydfI < > ydfK ydfK TTCCCA-16-GATAAT
35.43 relBE-hokD relB < > ydfV ydfV
37.33 TTGCCG-17-AATAAT grxD grxD < HNS > ydhO ydhO TTAACT-18-TAGAAT
37.44 GTGAAA-17-TAATAT ydhB ydhB* < > ydhC ydhC TTCACA-17-TACACT
37.78 TTGTCT-18-TTTAAT ydhZ ydhZ < HNS > pykF pykF TTAACT-18-TATATT
38.77 TCGTCA-17-TAAAAT thrS- -infA thrS < > arpB arpB ATGATG-16-TATAAC
41.11 TTGCGA-17-TATAAT kdgR kdgR* < > yebQ yebQ TTATCA-17-TATAAA
41.39 TTAACA-15-TGTGAT pphA pphA < > ryeA ryeA ATCACA-18-TAAAAA
41.57 TTGATT-17-TATACT yebG yebG < HNS > purT purT AAGACA-18-TATACT
41.81 ATGAAA-17-TCTCAT znuA znuA < > znuC znuCB ATGAGA-17-TTTCAT
41.90 TTGTCA-15-TACAAA yobI yobI < > yebB yebB CTGAAA-17-TATTAC
41.96 TTTCCA-17-TTTCAT aspS aspS < > yecD yecDEN GACACA-19-TATTAT
42.61 TTGAAA-17-TATGAC insAB-5 insA5 < > uspC uspC TTGGCA-16-TATAAG
43.11 CTAACA-19-AAAAAT fliC fliC < > fliD fliCST
43.23 TTGAAA-18-TTTAAT yedD yedD < > yedE yedEF TTCAGA-17-TATTAT
43.32 CTGACA-16-ACTAAT yedM yedM < > intG intG TTGCCC-16-CACAAT
43.89 TTGTAA-16-TATAAC yedWV yedW* < > hluH hluH TAGAAA-15-CAAAAT
44.99 CTGACA-16-ACTAAT yefM-yoeB yefM* < > hisL hisLG--AFI TAAACA-19-TATAAA
46.28 TTGTTA-18-TATAAC yegI yegI < HNS > yegJ yegJ ATGATA-17-TACAAT
46.68 TAAACA-18-TATATT yegRZ yegR < HNS > yegS yegS TTTAAA-17-TATTAT
48.01 TCAACA-18-CATCAT dusC dusC < > yohJ yohJK
48.30 ATGAAA-15-AATAAT folE-yeiB folE < > yeiG yeiG CTCACA-17-TATGAT
48.42 TTAAAA-19-TATAAT yeiE yeiE* < > yeiH yeiH TTACCA-15-TATAAA
49.59 napF- -ccmH napF < > yojO yojO TTGAGA-16-CATAAT
49.78 ATGAAA-17-TATCAT ompC ompC < HNS > micF micF TTAAGA-17-AATAAG
50.41 TTGATC-16-CATAAT ygaL yfaL < HNS > nrdA nrdAB
51.81 CTGCCA-16-TATGAT lrhA lrhA* < > alaA alaA TTAAGA-17-TATAAC
51.96 ATGACA-17-CATCAT yfbV yfbV < > ackA ackA-pta CTGAAA-16-TAGACT
52.11 GTGACA-15-TATAGT yfcF yfcF < > yfcG yfcG TCGGCA-15-TATCAA
53.46 TTGATA-17-TTTCAT emrKV emrK < HNS > evgA* evgAS TTGACA-19-TATATG
53.73 TTGCCA-17-TACTAT yfdV yfdY < > lpxP lpxP TCGACA-18-AACAAT
54.02 TTTACA-17-TATTTT glk glk < > yfeO yfeO CTCACA-17-CATAAC
54.09 TTGATA-17-TAGCAT mntH mntH < > nupC nupC ATGACT-17-TTTAAT
54.40 yfeR yfeR* < > yfeH yfeH TTCTCA-17-TAGAAC
54.49 TTAATA-16-GGTAAT zipA zipA < > cysZ cysZK TTAACT-19-TGAAAT
54.96 TTGGCT-17-CATAAT ypeA-yfeZ ypeA < > amiA amiA-hemF TTGTCA-17-TAAAAC
55.80 TTCACC-17-TAGAAT ypfM ypfM < HNS > yffB yffB- -ypfN TTCATA-16-TTTAAT
55.97 dapA-bamC dapA < > gcvR* gcvR TTAAAA-18-TCTGAT
56.43 TTGCCG-16-TATAAT upp-uraA upp < HNS > purM purMN ATGATA-17-TATTTT
56.57 ATGACA-18-TATAGT yfgF yfgF < HNS > yfgG yfgG TTGACC-17-CTTAAT
56.70 guaBA guaB < > xseA xseA TCGACT-16-TAGAAT
57.34 TCGGCA-18-TAAGAT trmJ trmJ < > suhB suhB AAGACA-17-TATACT
57.81 TTGATG-16-TATAAA glyA glyA < > hmp hmp TTTACA-16-TATAAG
58.35 TAAACA-17-CATCAT rpoE--rseC rpoE* < > nadB nadB TAGACT-16-TATAAG
59.01 TTGAAA-16-TATCGT aroF-tyrA aroF < > yfiL yfiL TTTTCA-17-TTTTAT
59.32 ratAB ratA < > smpB smpB
60.00 TAGATA-16-TATCAT ileY ileY < HNS > ygaQ ygaQ_12 TTATCA-17-TGTAAT
60.24 TTAAGA-17-AACAAT yqaE yqaE < HNS > ygaV* ygaVF TTTAGA-17-AATACT
60.27 TTTACA-17-TTTAAT stpA stpA* < HNS > alaE alaE GTGATA-17-TCTAAT
60.83 TTGAAA-16-TTTGAT mltB mltB < HNS > srlA srlA- -gutM TTAACA-18-TATGGT
61.61 TTCACA-18-TATTTT ygbI ygbI* < HNS > ygbJ ygbJK TTCACA-16-GTTAAT
62.45 TTGACA-16-TGTGAT ygcW ygcW < HNS > yqcE yqcE-ygcE TTCTCA-18-GATAAT
62.56 TTTAAA-17-GTTAAT queE queE < HNS > yqcG ygcG TTAACA-18-GATAAT
63.94 TTGACG-18-TTTAGT rppH-ptsP rppH < > mutH mutH TCGGCA-18-TTTAAT
65.80 ygfB- -visC ygfB < > zapA zapA TTGTCT-17-TATAGT
68.75 TAGAGA-19-TTTAAT glgS glgS < HNS > yqiJ yqiJK TTTAAA-15-TATATT
69.27 TTAACA-18-TTTTAT yqjH yqjH < > yqjI* yqjI TTGCAA-16-TATAAA
69.31 TTGATC-18-TATAGT aer aer < HNS > patA ygjG TAAACA-19-TAAAAT
69.68 TTTTCA-18-TATCAT rlmG rlmG < > ygjP ygjP TTGCCC-18-TATACC
70.35 TTGATA-16-TGTAAT trcA- -G tdcA* < HNS > tdcR tdcR TTTAAA-16-TATAAA
71.48 TTGACA-18-TATAAT metY- -pnp metY < > argG argG ATGAAA-17-AAAAAT
71.93 TGGACT-16-TAAAAC mlaFEDCB mlaF < > yrbG yrbG- -lptAB TTTACT-17-CAAAAT
72.51 TTACCA-16-CATAAA insH-10 insH10 < > yhcF yfcF CTCACA-18-TTTAAG
72.99 ATAACA-18-TATATT aaeXAB aaeX < HNS > aaeR* aaeR TTGATA-19-TGTTAT
73.51 TTAAAA-17-TATATT envR envR* < HNS > acrE acrEF TTGAGT-19-AATAAT
73.94 CTGTCA-18-TAGAAT smf smf < > def def-fmt TTGCTA-19-GATAAG
74.39 TTAATA-17-TATGAT gspAB gspA < HNS > gspC gspD- -LMO TTGATT-17-TACTAT
76.72 TTTACG-19-CATAAT glpEGR glpE < > glpD glpD TTGAAA-19-TATAAC
77.61 GTGCCA-18-TGTAGT ftsYEX ftsY < > rsmD rsmD-yhhL TTCACA-18-TGTTAT
78.27 TTTACA-17-GATTAT yhiL yhiL < HNS > yhiM yhiM CTGAAA-16-TATAAA
78.33 TTGCCC-17-GATAAT yhiN yhiN < HNS > pitA pitA TTCACT-18-TATAAT
78.58 ATGACG-17-TATAAA dinQ dinQ < > arsR* arsRBC ATGACG-17-TATAAA
78.67 TTGTCA-18-TATAAT insH-11 insH11 < HNS > slp slp-dctR TTTACG-17-TAAAAT
78.93 TTGAAA-16-TATAAG gadW gadW* < HNS > gadY gadY TTCGCA-18-TATAAA
80.49 TTCACA-17-TCAAAT bax bax < > malS malS TTGATA-18-TCAAAT
80.81 TTGTAA-19-TTTTAT yiaT yiaT < HNS > yiaU* yiaU TTCATA-18-TAAAAT
81.24 TATACA-18-TATAAT yibIH yibI < > mtlA mltADR TCTACA-19-TACAAT
81.71 TATACA-18-TATAAT yibB htrL < HNS > hldD rfaD-waaCL TTAATA-17-CATAAA
82.01 TCTACA-17-TTTATT waaQ- -waaK waaQ < HNS > waaA waaA-coaD CTGACA-16-TTTTAT
82.28 GTCACA-16-TAAAAG ligB ligB < > gmk gmk GTCACA-16-AATAAG
82.60 TTGTAA-19-TATAAT yicJI yicJ < HNS > selC selC TTATCA-18-TATAAA
83.00 TGGACA-17-GATACT istR istR < HNS > tisB tisA TTGTCC-17-TATACA
84.59 TTTATA-18-TATGAT asnC--mnmG asnC* < > asnA asnA TTGATT-16-TAAAAT
85.06 TTGGCC-18-AATAAT yifB yifB < HNS > ilvL ilvLXGM TTGGCC-18-AATAAT
85.39 TTCACC-19-AATAGT rhlB rhlB < > trxA trxA TTTACG-16-AATAAA
85.83 TTAACA-18-TGTAAT aslA aslA < HNS > glmZ glmZ TTGAGA-17-GATGAT
85.95 CTCACA-15-TGTAAT hemCDKY hemC < > cyaA cyaA CTGACA-18-TAGGAT
87.95 TCGACA-15-TACATT fdoG--fdhE fdoG < > fdhD fdhD TCGACA-15-TACATT
88.43 TTTGCA-19-TATCGT cpxRA cpxR* < > cpxP cpxP ATGACG-19-TTTAAA
88.70 TGGACA-17-TACAAT glpFKX glpF < > zapB zapB TGGACA-17-TACAAT
88.86 TTTGCA-18-TATGAT priA priA < > rpmE rpmE CAGACA-17-TATAGC
88.93 TTGAGC-17-TAAAAT metJ metJ* < > metB metBL TTGAGC-17-TAAAAT
89.48 TTCATA-17-GATACT argE argE < > argC argCBH TTGACA-18-TATCAA
91.12 TTGGCT-16-TCAAAT pepE pepE < > rluF rluF ATAACA-17-TATTTT
91.17 TTGACA-16-TTTATT lysC lysC < > pgi pgi ATCACA-18-TACAAT
91.65 plsB plsB < > dgkA dgkA TTAACG-19-CATAAT
91.76 TTGATA-17-CATAAC zur zur* < > yjbL yjbLM TTGTCG-18-AATAAT
91.84 qorA qorA < > dnaB dnaB TCGTCA-17-TAAAGT
92.01 ATGCCA-15-TTTAAT uvrA uvrA < > ssb ssb TTGACC-18-TGGAAT
92.09 CTAACA-15-TATAGT yjcB yjcB < HNS > yjcC yjcC TTTTCA-16-TATAAA
94.13 GTGAAA-18-TTTCAT yjeH yjeH < > groS groSL TTTTCA-17-CAGAAT
95.19 ATCACA-18-TATCAT ulaG ulaG < HNS > ulaA ulaA--EF TTAACT-15-GATAAT
95.32 TTGATT-15-GATCAT yjfY yjfY < > rpsF rpsF- -rplI TTCAAA-17-TGTGAT
95.37 TTGATT-15-GATCAT yjfZ yjfZ < HNS > ytfA* ytfA TTCACA-16-AATAAA
95.51 TCGACA-15-TACATT qorB qorB < > ytfH* ytfH
96.37 TTGACA-17-TGTAAT bdcA bdcA < > bdcR* bdcR TTGATT-17-TACAAA
96.46 TTGCAA-15-TATAAA argI argI < > rraB rraB TTAAAA-16-GATTAT
96.50 TTGATA-19-TAAAAT yjgM yjgM < HNS > yjgN yjgN TTGCCA-18-TATTGT
96.99 TTAACA-17-GATAAA insG insG < > yjhB yjhBC AAGACA-17-TATTGT
97.13 TTAACG-17-TAGAAT insM insM < > yjhV yjhV CTGTCA-16-TATAAA
97.33 TTCTCA-17-GATAAT fecIR fecI** < > insA7 insA-7 TTAACA-17-TATAAG
97.78 TCAACA-17-TTTAAT nanCM nanC < HNS > fimB fimB TTGGCA-16-TATATT
98.46 TTCACA-19-TTTTAT yjiR yjiR* < > yjiS yjiS TTAACC-15-TAAAAG
99.15 ATGAAA-17-TTTAAT yjjP yjjP < HNS > yjjQ* yjjQ-bglJ CTGATA-17-GATAAT
99.21 TTGATA-19-GATAAT fhuF fhuF < > yjjZ yjjZ TTGCAA-16-TATGAT
99.96 arcA arcA* < > yjjY yjjY TTGCCA-19-TACAAA
300 genes (a) 178 (b) K HNS 178 (a) 291 genes (b)
1.68 (a/b) 26 TFs 64 19 TFs 1.63 (a/b)
63 Y-genes 36% 74 Y-genes
20 essential 19 essential

A total of 1,075 RpoD holoenzyme-binding sites were identified within spacers on the entire E. coli K-12 W3110 genome. The constitutive promoters were predicted within type-A and type-B intergenic spacers (see Fig. 1A for classficiation). A total of 178 RNA polymerase RpoD holoenzyme-binding sites were identified within type-A spacers, which direct bidirectional transcription. Based on the gene orientation around these promoters, the genes and operons under the control of these promoters were estimated, that are located on either left side (left gene column) or right side (right gene column) of the respective spacers. Genes encoding transcription factors are indicated by star symbols (*) and the operons are shown in the operon columns [note that only the first and the last genes are shown for polycitronic operons]. The directions of transcription for these flanking genes are shown by arrows in column D. The map positions of left-side and right-side genes are shown in the map columns. The essential genes listed in the PEC database are underlined. The promoter sequences were predicted according to the analysis procedure described in Materials and Methods. For some spacers, multiple promoters were identified, of which the best-match promoters with the highest scores are described. The promoter sequence with complete match with the canonical promoter (see Fig. 4) is shown in bold and italic while the promoter sequence with 5-out of-6 match is shown in bold. The spacer including H-NS binding sites are shown as HNS mark in the spacer column. The numbers of hitherto identified promoters are 121 and 133 for left-ward and right-ward transcription, respectively, which correspond to 68 and 75%. Total number of genes under the control of 178 promoters were 300 for left-ward transcription, and 291 for right-ward transcription. The average numbers of genes under one promoter are 1.68 and 1.63 for left-ward and right-ward transcription, respectively. Among the total of 178 RpoD holoenyme-binding sites, 64 (36%) overlap with the H-NS-binding sites.