View conserved binding sites


Gene
GM15G06560
Description
Transcription factor GTE3, chloroplastic
Alias
Glyma15g06560;Glyma15g06560.1;PAC:26331776;Glyma15g06560.1
Binding siteNameDescriptionLocationPosition
Matrix_101ERF5Not availableUpstream-146
Matrix_104PINot availableUpstream-424
Matrix_109GBF3Not availableUpstream-423
Upstream-424
Matrix_110ATABI4;AT3G57600Not availableUpstream-146
Upstream-158
Matrix_113ABI5Not availableUpstream-423
Upstream-424
Upstream-426
Matrix_118PIF3_2Direct targeting of light signals to a promoter element-bound transcription factorUpstream-421
Upstream-541
Upstream-542
Matrix_119RRTF1Not availableUpstream-147
Matrix_120BEE2Not availableUpstream-424
Upstream-425
Matrix_122ABF1;AREB2Not availableUpstream-425
Matrix_129ABF1Not availableUpstream-424
Upstream-425
Matrix_130TCP16Not availableUpstream-539
Upstream-540
Matrix_134ABF1Not availableUpstream-424
Upstream-425
Matrix_138RRTF1Not availableUpstream-147
Matrix_143GATA14;GATA6;GATA5Not availableUpstream-388
Matrix_144AT5G08330;AT5G23280Not availableUpstream-539
Upstream-540
Matrix_145GBF4;AT5G44080Not availableUpstream-425
Upstream-426
Matrix_146ORA47Not availableUpstream-146
Matrix_147ERF3;AT1G80580Not availableUpstream-148
Matrix_15PIF3_1Direct targeting of light signals to a promoter element-bound transcription factorUpstream-421
Matrix_155RAP2.6;ERF110;ABR1Not availableUpstream-147
Matrix_156POC1Not availableUpstream-423
Upstream-424
Matrix_16AT3G04450;PHL1Not availableUpstream-28
Matrix_162AtPHR1Not availableUpstream-28
Matrix_166TGA4Not availableUpstream-552
Upstream-553
Matrix_169E2F1Not availableUpstream-145
Matrix_171LBD3;LBD4Not availableUpstream-530
Upstream-531
Matrix_179AtMYB84;MYB36;MYB68;O49746_ARATHNot availableUpstream-534
Upstream-535
Matrix_181Dof5.7Not availableUpstream-249
Matrix_183BES1A brassinosteroid transcriptional network revealed by genome-wide identification of BESI target genes in Arabidopsis thalianaUpstream-121
Upstream-122
Upstream-141
Upstream-142
Matrix_186FHY3Not availableUpstream-121
Upstream-122
Upstream-142
Upstream-584
Matrix_19PIF5Phytochrome interacting factors 4 and 5 control seedling growth in changing light conditions by directly controlling auxin signalingUpstream-287
Upstream-288
Matrix_190ATERF1Not availableUpstream-145
Upstream-146
Matrix_192FHY3/FAR1Not availableUpstream-421
Upstream-422
Upstream-426
Matrix_194HYH;HY5Not availableUpstream-422
Matrix_195GATA2;GATA4Not availableUpstream-387
Matrix_196TCP20;AT5G41030Not availableUpstream-537
Upstream-538
Upstream-541
Upstream-542
Matrix_200PIL5;AT4G28790;AT4G28800;AT4G28811;AT4G28815Not availableUpstream-424
Upstream-425
Matrix_214AP1Not availableUpstream-423
Matrix_216TCP16Not availableUpstream-539
Upstream-540
Matrix_217BES1A brassinosteroid transcriptional network revealed by genome-wide identification of BESI target genes in Arabidopsis thalianaUpstream-421
Upstream-422
Matrix_224ERF1Not availableUpstream-126
Matrix_232TCP23Not availableUpstream-538
Upstream-539
Upstream-540
Upstream-541
Matrix_234RAP2.3Not availableUpstream-125
Upstream-126
Upstream-146
Matrix_24POC1Not availableUpstream-421
Matrix_243RAP2.12;RAP2.2Not availableUpstream-147
Matrix_246ARR10Molecular structure of the GARP family of plant Myb-related DNA binding motifs of the Arabidopsis response regulatorsUpstream-278
Upstream-279
Upstream-1251
Matrix_247PIF3Not availableUpstream-424
Upstream-425
Matrix_250ADAP;AT1G79700;WRI1;ANTNot availableUpstream-295
Matrix_252RAP2.6Not availableUpstream-125
Upstream-126
Upstream-146
Matrix_261ATERF-1Not availableUpstream-17
Upstream-147
Matrix_264ATAREB1Not availableUpstream-421
Upstream-422
Upstream-425
Matrix_266TCP16Determinants of the DNA binding specificity of class I and class II TCP transcription factorsUpstream-539
Upstream-540
Matrix_272DEAR4Not availableUpstream-125
Upstream-126
Upstream-146
Matrix_278AtbZIP44Not availableUpstream-552
Upstream-553
Matrix_279HRS1Not availableUpstream-29
Matrix_280TCP24;TCP1;BRC2;ATTCP18Not availableUpstream-158
Upstream-538
Upstream-539
Matrix_281TCP13Not availableUpstream-539
Upstream-540
Upstream-541
Upstream-542
Matrix_288RAP2.3Not availableUpstream-147
Matrix_294MEE35Not availableUpstream-539
Upstream-540
Upstream-541
Upstream-542
Matrix_295ERF1Not availableUpstream-148
Matrix_296GBF2Not availableUpstream-424
Upstream-425
Upstream-426
Matrix_297TCP15Not availableUpstream-538
Upstream-539
Upstream-540
Upstream-541
Matrix_300bZIP68;bZIP16Not availableUpstream-424
Upstream-425
Upstream-426
Matrix_301PIL5Not availableUpstream-426
Matrix_305PIF4Interaction between BZR1 and PIF4 integrates brassinosteroid and environmental responsesUpstream-380
Upstream-381
Matrix_306TGA1TGA1 and G-box binding factors: two distinct classes of Arabidopsis leucine zipper proteins compete for the G-box-like element TGACGTGGUpstream-550
Upstream-551
Matrix_324AT2G01060Not availableUpstream-28
Matrix_326AT5G07310;Rap2.6L;AT5G61890Not availableUpstream-126
Upstream-127
Upstream-147
Matrix_331GBF1Not availableUpstream-423
Upstream-424
Matrix_332SPT;ALCNot availableUpstream-288
Upstream-289
Upstream-424
Upstream-425
Matrix_333GATA3Not availableUpstream-1252
Matrix_335HSFB2ANot availableUpstream-299
Matrix_338AP2Not availableUpstream-422
Matrix_339bHLH104Not availableUpstream-424
Matrix_343AT2G33710Not availableUpstream-125
Upstream-126
Upstream-146
Upstream-149
Matrix_345POC1Not availableUpstream-421
Upstream-541
Upstream-542
Matrix_348AT5G51910Not availableUpstream-539
Upstream-540
Matrix_355ERF10;ERF11Not availableUpstream-126
Upstream-127
Upstream-147
Matrix_356PRR5Not availableUpstream-418
Upstream-422
Matrix_360ORA59Not availableUpstream-148
Matrix_361AT1G25550Not availableUpstream-28
Matrix_363RAP2.3Not availableUpstream-147
Matrix_374AT5G07580;AT5G61590Not availableUpstream-147
Matrix_377AT1G75490;DREB2C;AT2G40350;AT5G18450Not availableUpstream-126
Matrix_378ATERF1Not availableUpstream-125
Upstream-126
Upstream-146
Matrix_380ATMYR1Not availableUpstream-29
Upstream-30
Matrix_389ILR3Not availableUpstream-424
Upstream-425
Matrix_396AP3Not availableUpstream-208
Upstream-520
Upstream-521
Matrix_399TGA1Not availableUpstream-551
Upstream-552
Matrix_40TCP2Not availableUpstream-539
Upstream-540
Matrix_401MYB55Not availableUpstream-533
Matrix_403BZR1Not availableUpstream-423
Upstream-542
Upstream-543
Matrix_406ATERF-7Not availableUpstream-148
Matrix_409DEAR3Not availableUpstream-126
Matrix_42AT2G45680Not availableUpstream-539
Upstream-540
Matrix_426CRF1;CRF2Not availableUpstream-126
Upstream-127
Upstream-147
Matrix_438AtbZIP63Not availableUpstream-551
Upstream-552
Matrix_443AGL15Not availableUpstream-423
Matrix_45DRNNot availableUpstream-148
Matrix_456bZIP60Not availableUpstream-553
Upstream-554
Matrix_457TGA2Not availableUpstream-551
Upstream-552
Matrix_462ATERF-8Not availableUpstream-148
Matrix_473RRTF1Not availableUpstream-125
Upstream-126
Upstream-146
Matrix_475AT5G64220Not availableUpstream-124
Upstream-125
Upstream-145
Matrix_480BES1Not availableUpstream-423
Upstream-424
Matrix_484ATERF13Not availableUpstream-148
Matrix_488ABF1Not availableUpstream-417
Matrix_491AT1G68670;AT3G25790Not availableUpstream-28
Matrix_493AT1G22985;AT1G71130Not availableUpstream-126
Upstream-127
Upstream-147
Matrix_506DRNL;ATERF-4Not availableUpstream-147
Matrix_507TCP3Not availableUpstream-538
Upstream-539
Upstream-540
Upstream-541
Matrix_508APL;AT3G12730;AT3G24120;UNE16Not availableUpstream-28
Matrix_517ERF12Not availableUpstream-148
Matrix_53MYC3Not availableUpstream-123
Upstream-124
Upstream-144
Upstream-422
Upstream-423
Matrix_55PIF3Not availableUpstream-424
Matrix_56BZIP17;BZIP28;BZIP49Not availableUpstream-552
Upstream-553
Matrix_57WIN1;SHN3;SHN2Not availableUpstream-126
Upstream-127
Upstream-147
Upstream-150
Matrix_60AT1G01260;AT5G57150Not availableUpstream-425
Matrix_65POC1;PIL1Not availableUpstream-425
Matrix_69AT2G03500Not availableUpstream-28
Matrix_7PIF4Not availableUpstream-424
Upstream-425
Upstream-426
Upstream-427
Matrix_77PRR5Not availableUpstream-121
Upstream-141
Upstream-424
Upstream-584
Matrix_80BIM1Not availableUpstream-121
Upstream-140
Upstream-141
Matrix_82TCP17Not availableUpstream-539
Upstream-540
Upstream-541
Upstream-542
Matrix_86CRF5;CRF6;CRF4Not availableUpstream-126
Upstream-127
Upstream-147
Matrix_91CRF3Not availableUpstream-126
Matrix_94TCP5Not availableUpstream-539
Upstream-541
Matrix_97APRR2Not availableUpstream-28
Motif_116INRNTPSADBInr (initiator) elements found in the tobacco psaDb gene promoter without TATA boxes; Light-responsive transcription of psaDb depends on Inr, but not TATA boxUpstream-488
Motif_139RHERPATEXPA7Right part of RHEs (Root Hair-specific cis-Elements) conserved among the Arabidopsis thaliana A7 (AtEXPA7) orthologous (and paralogous) genes from diverse angiosperm species with different hair distribution patternsUpstream-414
Motif_141BP5OSWXOsBP-5 (a MYC protein) binding site in Wx promoterUpstream-546
Motif_150RBCSCONSENSUSrbcS general consensus sequence; AATCCAA or AATCCAACUpstream-474
Upstream-480
Motif_179CACGTGMOTIF;BES1;PIF4;PIF5Phytochrome interacting factors 4 and 5 control seedling growth in changing light conditions by directly controlling auxin signaling;A brassinosteroid transcriptional network revealed by genome-wide identification of BESI target genes in Arabidopsis thaliana; CACGTG motif; G-box; Binding site of Arabidopsis GBF4; C. roseus G-box binding factor 1 (CrGBF1) and 1 (CrGBF2) can act as transcriptional repressors of the Str promoter via direct interaction with the G-box; Essential for expression of beta-phaseolin gene during embryogenesis in bean, tobacco, Arabidopsis; Tomato Pti4 (ERF) regulates defense-related gene expression via GCC box and non-GCC box cis-element (Myb1 (GTTAGTT) and G-box (CACGTG)); Isolation and characterization of a fourth Arabidopsis thaliana G-box-binding factor, which has similarities to Fos oncoprotein; Interaction between BZR1 and PIF4 integrates brassinosteroid and environmental responsesUpstream-426
Motif_189CMSRE1IBSPOACMSRE-1 (Carbohydrate Metabolite Signal Responsive Element 1) found in the promoter of sweet potato sporamin A geneUpstream-19
Motif_194EBOXBNNAPAE-box of napA storage-protein gene of Brassica napus;This sequence is also known as RRE (R response element); MYC recognition site found in the promoters of the dehydration-responsive gene rd22 and many other genes in Arabidopsis; Binding site of ATMYC2 (previously known as rd22BP1); see E-box and MYCATRD22; MYC recognition sequence in CBF3 promoter; Binding site of ICE1 (inducer of CBF expression 1) that regulates the transcription of CBF/DREB1 genes in the cold in Arabidopsis; ICE1Upstream-426
Motif_200GBOXLERBCSG box; Conserved sequence upstream of light-regulated genes; Sequence found in the promoter region of rbcS of tomato and Arabidopsis; Binding with GBFUpstream-424
Motif_218ABRERATCALABRE-related sequence or Repeated sequence motifs identified in the upstream regions of 162 Ca(2+)-responsive upregulated genes; see also ABREUpstream-104
Upstream-123
Upstream-143
Upstream-425
Upstream-426
Upstream-545
Upstream-602
Upstream-603
Motif_221MYBCOREBinding site for all animal MYB and at least two plant MYB proteins ATMYB1 and ATMYB2, both isolated from Arabidopsis; ATMYB2 is involved in regulation of genes that are responsive to water stress in Arabidopsis; A petunia MYB protein (MYB.Ph3) is involved in regulation of flavonoid biosynthesisUpstream-182
Upstream-260
Motif_223LTRE1HVBLT49LTRE-1 (low-temperature-responsive element) in barley (H.v.) blt4.9 gene promoter; A new LTRE; A previously known LTRE is CCGACUpstream-374
Motif_24CAMTA3Roles for Arabidopsis CAMTA transcription factors in cold-regulated gene expression and freezing toleranceUpstream-602
Motif_243TELOBOXATEEF1AA1telo-box (telomere motif) found in the Arabidopsis (A.t.) eEF1AA1 gene promoter; Conserved in all known plant eEF1A gene promoters; Found in the 5' region of numerous genes encoding components of the translational apparatus; Required for the activation of expression in root primordia; Acts co-operatively with tef-box; Binding site of AtPur alpha-1; Plant interstitial telomere mitifs participate in the control of gene expression in root meristemsUpstream-208
Upstream-521
Motif_244ABRE-like binding site motifNot availableUpstream-424
Upstream-426
Upstream-555
Motif_249DPBF1&2 binding site motifA novel class of bZIP transcription factors, DPBF-1 and 2 (Dc3 promoter-binding factor-1 and 2) binding core sequence; Found in the carrot Dc3 gene promoter; Dc3 expression is normally embryo-specific, and also can be induced by ABA; The Arabidopsis abscisic acid response gene ABI5 encodes a bZIP transcription factor; abi5 mutant have a pleiotropic defects in ABA response; ABI5 regulates a subset of late embryogenesis-abundant genes; GIA1 (growth-insensitivity to ABA) is identical to ABI5; Isolation of a novel class of bZIP transcription factors that interact with ABA-responsive and embryo-specification elements in the Dc3 promoter using a modified yeast one-hybrid systemUpstream-426
Upstream-604
Motif_291ABFs binding site motifBinding site of trans-acting factor EMBP-1; wheat Em gene;Binding site of ABFs; ABFs (ABRE binding factors) were isolated from Arabidopsis by a yeast one-hybrid screening system; Expression ABFs is induced by ABA and various stress treatment; ABFs belongs to a distinct subfamily of bZIP proteins; Involved in ABA-mediated stress-signaling pathway;A plant leucine zipper protein that recognizes an abscisic acid response elementUpstream-424
Motif_310ANAERO3CONSENSUSOne of 16 motifs found in silico in promoters of 13 anaerobic genes involved in the fermentative pathway (anaerobic set 1); Arbitrary named ANAERO3CONSENSUS by the PLACEdb curatorUpstream-405
Motif_318CGCGBOXATCGCG box recognized by AtSR1-6 (Arabidopsis thaliana signal-responsive genes); Multiple CGCG elements are found in promoters of many genes; Ca2+/calmodulin binds to all AtSRsUpstream-105
Upstream-124
Upstream-144
Upstream-603
Motif_339ABRE-like binding site motifMolecular responses to dehydration and low temperatureUpstream-426
Upstream-555
Motif_346SND1;VND6;VND7;NST1;NST2Global analysis of direct targets of secondary wall NAC master switches in ArabidopsisUpstream-235
Motif_347OPAQUE2ZMB32opaque-2 binding site of maize b-32 (type I ribosome-inactivating protein gene); O2; O2S; O2S and GARE form a gibberellin response complex (GARC)Upstream-552
Motif_353HEXAMERATH4hexamer motif of Arabidopsis thaliana histone H4 promoter;Identification of cis-elements regulating the expression of an Arabidopsis histone H4 geneUpstream-345
Motif_358UP2ATMSDUp2 motif found in 193 of the 1184 up-regulated genes after main stem decapitation in ArabidopsisUpstream-209
Upstream-522
Motif_371DRE-like promoter motifThe Expression Profile Matrix of Arabidopsis Transcription Factor Genes Suggests Their Putative Functions in Response to Environmental StressesUpstream-67
Upstream-343
Motif_383UPRE-III(bZIP60)The plant-specific transcription factor NAC103 is induced by bZIP60 through a new cis-regulatory element to modulate the unfolded protein response in ArabidopsisUpstream-277
Motif_392PALBOXAPCBox A; Consensus; One of three putative cis-acting elements (boxes P, A, and L) of phenylalanine ammonia-lyase (PAL; EC 4.3.1.5) genes in parsley (P.c.); None of these elements (boxes P, A, and L) alone, or the promoter region containing all of them together, conferred elicitor or light responsiveness. These elements appear to be necessary but not sufficient for elicitor- or light-mediated PAL gene activation; See also Box P, Box LUpstream-19
Upstream-369
Motif_399UPRMOTIFIATMotif I in the conserved UPR (unfolded protein response) cis-acting element in Arabidopsis genes coding for SAR1B, HSP-90, SBR-like, Ca-ATPase 4, CNX1, PDI, etc.Upstream-554
Motif_411PRECONSCRHSP70AConsensus sequence of PRE (plastid response element) in the promoters of HSP70A in Chlamydomonas; Involved in induction of HSP70A gene by both MgProto and lightUpstream-344
Motif_440TGA1 binding site motifHex motif; Binding site of Arabidopsis bZIP protein TGA1 and G box binding factor GBF1; TGA1 and members of the GBF family differ in their DNA binding properties; G-Box-like element;TGA1 and G-box binding factors: two distinct classes of Arabidopsis leucine zipper proteins compete for the G-box-like element TGACGTGGUpstream-554
Motif_448IRO2OSOsIRO2-binding core sequence; G-box plus G; Transcription factor OsIRO2 is induced exclusively by Fe deficiencyUpstream-425
Motif_450E2FCONSENSUSE2F consensus sequence of all different E2F-DP-binding motifs that were experimentally verified in plantsUpstream-107
Motif_453HEXMOTIFTAH3H4hexamer motif found in promoter of wheat histone genes H3 and H4; CaMV35S; NOS; Binding with HBP-1A and HBP-1B; Binding site of wheat nuclear protein HBP-1 (histone DNA binding protein-1); HBP-1 has a leucine zipper motif; hexamer motif in type 1 element may play important roles in regulation of replication- dependent but not of replication-independent expression of the wheat histone H3 gene;Rice OBF1-homodimer-binding siteUpstream-554
Motif_48BES1A brassinosteroid transcriptional network revealed by genome-wide identification of BESI target genes in Arabidopsis thalianaUpstream-587
Motif_542ABI5;AtMYC2;HY5A basic helix-loop-helix transcription factor in Arabidopsis, MYC2, acts as a repressor of blue light-mediated photomorphogenic growth. Arabidopsis bZIP protein HY5 directly interacts with light-responsive promoters in mediating light control of gene expression. The homologous ABI5 and EEL transcription factors function antagonistically to fine-tune gene expression during late embryogenesisUpstream-424
Motif_558BOXIIPCCHSCore of Box II/G box found in the parsley chs genes; Essential for light regulationUpstream-424
Motif_567T/GBOXATPIN2T/G-box found in tomato proteinase inhibitor II (pin2) and leucine aminopeptidase (LAP) genes; Involved in jasmonate (JA) induction of these genes; bHLH-Leu zipper JAMYC2 and JAMYC10 proteins specifically recognize this motifUpstream-546
Motif_57ABREOSRAB21ABA responsive element (ABRE) of wheat Em and rice rab21 genes; Proposed consensus sequence for the repeated motif (Em1a and Em1b) of wheat Em geneUpstream-543
Motif_576TGACGTVMAMYTGACGT motif found in the Vigna mungo alpha-Amylase (Amy) gene promoter; Located between -128 and -123; Required for high level expression of alpha-Amylase in the cotyledons of the germinated seedsUpstream-554
Motif_580L1BOXATPDF1L1 box found in promoter of Arabidopsis thaliana PROTODERMAL FACTOR1 (PDF1) gene; Located between -134 and -127; Involved in L1 layer-specific expression; L1-specific homeodomain protein ATML can bind to the L1 box; Y=C/T; A cotton fiber gene, RD22-like 1 (RDL1), contains a homeodomain binding L1 box and a MYB binding motif ; HDZip IV; Identification of a cis-regulatory element for L1 layer-specific gene expression, which is targeted by an L1-specific homeodomain proteinUpstream-516
Motif_584UPRE-III(bZIP60)The plant-specific transcription factor NAC103 is induced by bZIP60 through a new cis-regulatory element to modulate the unfolded protein response in ArabidopsisUpstream-277
Motif_591CBFHVBinding site of barley CBF1, and also of barley CBF2; CBF = C-repeat (CRT) binding factors; CBFs are also known as dehydration-responsive element (DRE) binding proteins (DREBs)Upstream-356
Upstream-609
Motif_603SITEIIATCYTCSite II element found in the promoter regions of cytochrome genes (Cytc-1, Cytc-2) in Arabidopsis; Located between -147 and -156 from the translational starts sites;Overrepresented in the promoters of nuclear genes encoding components of the oxidative phosphorylation (OxPhos) machinery from both Arabidopsis and riceUpstream-540
Upstream-542
Motif_606NAPINMOTIFBNSequence found in 5' upstream region (-6, -95, -188) of napin (2S albumin) gene in Brassica napus; Interact with a protein present in crude nuclear extracts from developing B. napus seedsDownstream2217
Motif_627ACGTABREMOTIFA2OSEMExperimentally determined sequence requirement of ACGT-core of motif A in ABRE of the rice gene, OSEM; DRE and ABRE are interdependent in the ABA-responsive expression of the rd29A in ArabidopsisUpstream-424
Upstream-427
Motif_630WBOXNTCHN48W box identified in the region between -125 and -69 of a tobacco class I basic chitinase gene CHN48; NtWRKY1, NtWRKY2 and NtWRKY4 bound to W box; NtWRKYs possibly involved in elicitor-respsonsive transcription of defense genes in tobaccoUpstream-83
Upstream-381
Motif_638ABRE binding site motifNot availableUpstream-424
Motif_641LRENPCABELRE; A positive light regulatory element in tobacco CAB (cab-E) gene; Located at -241Upstream-423
Motif_658GT1CONSENSUSConsensus GT-1 binding site in many light-regulated genes, e.g., RBCS from many species, PHYA from oat and rice, spinach RCA and PETA, and bean CHS15; GT-1 can stabilize the TFIIA-TBP-DNA (TATA box) complex; The activation mechanism of GT-1 may be achieved through direct interaction between TFIIA and GT-1; Binding of GT-1-like factors to the PR-1a promoter influences the level of SA-inducible gene expressionUpstream-598
Motif_72GADOWNATSequence present in 24 genes in the GA-down regulated d1 cluster (106 genes) found in Arabidopsis seed germination; This motif is similar to ABREUpstream-427
Motif_79UPRMOTIFIIATMotif II in the conserved UPR (unfolded protein response) cis-acting element in Arabidopsis genes coding for SAR1B, HSP-90, SBR-like, Ca-ATPase 4, CNX1, PDI, etcUpstream-386
Motif_92ABREATRD22ABRE (ABA responsive element) in Arabidopsis dehydration-responsive gene rd22Upstream-423
Motif_95UPRE2ATXBP1 binding site-like sequence found in the plant UPRE (unfolded protein response element) in Arabidopsis thaliana;Either of ERSEII or XBP1 binding sites is essential and sufficient for the UPRUpstream-552