Plant Transcription Factor Database
Previous version: v3.0
Transcription Factor Information
Basic Information | Signature Domain | Sequence | 
Basic Information? help Back to Top
TF ID AT1G42990.1
Common NameATBZIP60, BZIP60, F13A11.5
Taxonomic ID
Taxonomic Lineage
cellular organisms; Eukaryota; Viridiplantae; Streptophyta; Streptophytina; Embryophyta; Tracheophyta; Euphyllophyta; Spermatophyta; Magnoliophyta; Mesangiospermae; eudicotyledons; Gunneridae; Pentapetalae; rosids; malvids; Brassicales; Brassicaceae; Camelineae; Arabidopsis
Family bZIP
Protein Properties Length: 295aa    MW: 33174.1 Da    PI: 4.3626
Description basic region/leucine zipper motif 60
Gene Model
Gene Model ID Type Source Coding Sequence
AT1G42990.1genomeTAIRView CDS
Signature Domain? help Back to Top
Signature Domain
No. Domain Score E-value Start End HMM Start HMM End
       bZIP_1   4 lkrerrkqkNReAArrsRqRKkaeieeLeekvkeLeaeNkaLk 46 
                   k+ rr+ +NR AA rsR+RKk++++ Le+k+k Le e  +L 
                  699*********************************9976665 PP

Protein Features ? help Back to Top
3D Structure
Database Entry ID E-value Start End InterPro ID Description
Gene3DG3DSA: hitNo description
SMARTSM003385.5E-5138202IPR004827Basic-leucine zipper domain
PROSITE profilePS5021710.944140197IPR004827Basic-leucine zipper domain
PfamPF077162.5E-9142184IPR004827Basic-leucine zipper domain
SuperFamilySSF579592.94E-11142200No hitNo description
PROSITE patternPS000360145160IPR004827Basic-leucine zipper domain
CDDcd147047.00E-13156194No hitNo description
Gene Ontology ? help Back to Top
GO Term GO Category GO Description
GO:0002376Biological Processimmune system process
GO:0006355Biological Processregulation of transcription, DNA-templated
GO:0010200Biological Processresponse to chitin
GO:0030968Biological Processendoplasmic reticulum unfolded protein response
GO:0005634Cellular Componentnucleus
GO:0005789Cellular Componentendoplasmic reticulum membrane
GO:0016021Cellular Componentintegral component of membrane
GO:0003700Molecular Functiontranscription factor activity, sequence-specific DNA binding
GO:0043565Molecular Functionsequence-specific DNA binding
Plant Ontology ? help Back to Top
PO Term PO Category PO Description
PO:0000003anatomywhole plant
PO:0000013anatomycauline leaf
PO:0000014anatomyrosette leaf
PO:0000037anatomyshoot apex
PO:0000056anatomyflower bud
PO:0000084anatomyplant sperm cell
PO:0000230anatomyinflorescence meristem
PO:0000293anatomyguard cell
PO:0008019anatomyleaf lamina base
PO:0009006anatomyshoot system
PO:0009009anatomyplant embryo
PO:0009025anatomyvascular leaf
PO:0009052anatomyflower pedicel
PO:0020137anatomyleaf apex
PO:0025022anatomycollective leaf structure
PO:0025195anatomypollen tube cell
PO:0001016developmental stageL mature pollen stage
PO:0001017developmental stageM germinated pollen stage
PO:0001054developmental stagevascular leaf senescent stage
PO:0001078developmental stageplant embryo cotyledonary stage
PO:0001081developmental stagemature plant embryo stage
PO:0001185developmental stageplant embryo globular stage
PO:0004507developmental stageplant embryo bilateral stage
PO:0007064developmental stageLP.12 twelve leaves visible stage
PO:0007095developmental stageLP.08 eight leaves visible stage
PO:0007098developmental stageLP.02 two leaves visible stage
PO:0007103developmental stageLP.10 ten leaves visible stage
PO:0007115developmental stageLP.04 four leaves visible stage
PO:0007123developmental stageLP.06 six leaves visible stage
PO:0007131developmental stageseedling development stage
PO:0007611developmental stagepetal differentiation and expansion stage
PO:0007616developmental stageflowering stage
Sequence ? help Back to Top
Protein Sequence    Length: 295 aa     Download sequence    Send to blast
Nucleic Localization Signal ? help Back to Top
No. Start End Sequence
Expression -- UniGene ? help Back to Top
UniGene ID E-value Expressed in
At.212710.0flower| root| seed
Expression -- Microarray ? help Back to Top
Source ID E-value
Expression AtlasAT1G42990-
Expression -- Description ? help Back to Top
Source Description
UniprotTISSUE SPECIFICITY: Expressed in seedlings, rosette and cauline leaves, stems, buds, flowers, siliques, immature seeds, anthers and pollen grains. {ECO:0000269|PubMed:19017746}.
Functional Description ? help Back to Top
Source Description
TAIRAtbZIP60 consists of a bZIP DNA binding domain followed by a putative transmembrane domain. GFP fusions containing the first 260 amino acids (AtbZIP60deltaC) are nuclear-localized. AtbZIP60 is upregulated by the addition of tunicamycin (ER stress response inductor), DTT (inhibitor of disulfide bond formation) and azetin-2-carboxylate (proline analog perturbing protein structure). Upon ER stress the protein is proteolyzed and the soluble part is translocalized into the nucleus. AtbZIP60deltaC can activate the promoters of the ER chaperones BiP1, BiP2 and BiP3 and CNX1 and CNX2 via binding to the ER stress response element (ERSE) and the plant unfolded protein response element(P-UPRE). It can also activate its own transcription.
UniProtTranscription factor involved in the unfolded protein response (UPR). Acts during endoplasmic reticulum stress (ER) by activating unfolded protein response (UPR) target genes via direct binding to the UPR element (UPRE). Plays a role in plant immunity and abiotic stress responses. {ECO:0000269|PubMed:15781873, ECO:0000269|PubMed:18574595, ECO:0000269|PubMed:19017746, ECO:0000269|PubMed:21482766, ECO:0000269|PubMed:22359644}.
Function -- GeneRIF ? help Back to Top
  1. AtPDI gene expression was investigated in different tissues, in response to chemically induced unfolded protein response (UPR), and in null mutants of UPR signaling mediators (AtIRE1-2 and AtbZIP60).
    [PMID: 18574595]
  2. bZIP60 is involved in the plant endoplasmic reticulum stress response and is upregulated during normal development in anther cells.
    [PMID: 19017746]
  3. possible mechanisms of AtbZIP60's action in the nucleus are discussed
    [PMID: 19352029]
  4. observed that the active form of bZIP60 protein detected in the wild-type was missing in ire1a/ire1b
    [PMID: 22355548]
  5. the IRE1/bZIP60 branch of unfolded protein response is part of the plant response to pathogens for which two AT IRE1 isoforms play only partially overlapping roles; and IRE1 has both bZIP60-dependent and bZIP60-independent functions in plant immunity.
    [PMID: 22359644]
  6. overexpression of the AtbZIP60 gene in transgenic cell lines improved salt, drought, and cold stress tolerances by regulating expression of Ca(2+)-dependent protein kinase genes.
    [PMID: 23275191]
  7. bZIP60 binds directly to UPRE-III-containing DNA, and regulates NAC103.
    [PMID: 23869562]
  8. Results indicate that the ER-located transcription factors bZIP17 and bZIP60 play a role in salt stress but this response goes through a signaling pathway that is different to that triggered by the unfolded protein response.
    [PMID: 25704669]
  9. bZIP28 and bZIP60 interact with Ash2 and WDR5a, both of which are core COMPASS-like components.
    [PMID: 25730865]
  10. RLG1 proteins derived from transgene were mainly found in the cytoplasm; however, some were in the microsomal fraction, possibly on the ER membrane.
    [PMID: 26820526]
Binding Motif ? help Back to Top
Motif ID Method Source Motif file
Motif logo
Cis-element ? help Back to Top
Regulation -- Description ? help Back to Top
Source Description
UniProtINDUCTION: By tunicamycin, DTT and azetidine-2-carboxylate. Isoform 2 is induced by heat. {ECO:0000269|PubMed:15781873, ECO:0000269|PubMed:21482766}.
Regulation -- PlantRegMap ? help Back to Top
Source Upstream Regulator Target Gene
Regulation -- ATRM (Manually Curated Target Genes) ? help Back to Top
Source Target Gene (A: Activate/R: Repress)
ATRM AT1G09080(A), AT2G31955(A), AT5G20990(A), AT5G28540(A), AT5G42020(A)
Phenotype -- Disruption Phenotype ? help Back to Top
Source Description
UniProtDISRUPTION PHENOTYPE: No visible phenotype, but reduced induction of ER stress-responsive genes. Shows enhanced susceptibility to a bacterial pathogen. {ECO:0000269|PubMed:18574595, ECO:0000269|PubMed:19017746, ECO:0000269|PubMed:22359644}.
Phenotype -- Mutation ? help Back to Top
Source ID
T-DNA ExpressAT1G42990
Annotation -- Nucleotide ? help Back to Top
Source Hit ID E-value Description
GenBankAY0459640.0AY045964.1 Arabidopsis thaliana putative bZIP transcription factor (At1g42990) mRNA, complete cds.
GenBankAY1338050.0AY133805.1 Arabidopsis thaliana clone U10320 putative bZIP transcription factor (At1g42990) mRNA, complete cds.
Annotation -- Protein ? help Back to Top
Source Hit ID E-value Description
RefseqNP_174998.10.0bZIP transcription factor 60
SwissprotQ9C7S00.0BZP60_ARATH; bZIP transcription factor 60
TrEMBLD7KMV41e-142D7KMV4_ARALL; Putative uncharacterized protein
STRINGAT1G42990.10.0(Arabidopsis thaliana)
Orthologous Group ? help Back to Top
LineageOrthologous Group IDTaxa NumberGene Number
Representative plantOGRP54681518
Publications ? help Back to Top
  1. Riechmann JL, et al.
    Arabidopsis transcription factors: genome-wide comparative analysis among eukaryotes.
    Science, 2000. 290(5499): p. 2105-10
  2. Jakoby M, et al.
    bZIP transcription factors in Arabidopsis.
    Trends Plant Sci., 2002. 7(3): p. 106-11
  3. Yamada K, et al.
    Empirical analysis of transcriptional activity in the Arabidopsis genome.
    Science, 2003. 302(5646): p. 842-6
  4. Deppmann CD, et al.
    Dimerization specificity of all 67 B-ZIP motifs in Arabidopsis thaliana: a comparison to Homo sapiens B-ZIP motifs.
    Nucleic Acids Res., 2004. 32(11): p. 3435-45
  5. Casson S,Spencer M,Walker K,Lindsey K
    Laser capture microdissection for the analysis of gene expression during embryogenesis of Arabidopsis.
    Plant J., 2005. 42(1): p. 111-23
  6. Iwata Y,Koizumi N
    An Arabidopsis transcription factor, AtbZIP60, regulates the endoplasmic reticulum stress response in a manner unique to plants.
    Proc. Natl. Acad. Sci. U.S.A., 2005. 102(14): p. 5280-5
  7. Deppmann CD,Alvania RS,Taparowsky EJ
    Cross-species annotation of basic leucine zipper factor interactions: Insight into the evolution of closed interaction networks.
    Mol. Biol. Evol., 2006. 23(8): p. 1480-92
  8. Wright SI, et al.
    Testing for effects of recombination rate on nucleotide diversity in natural populations of Arabidopsis lyrata.
    Genetics, 2006. 174(3): p. 1421-30
  9. Kaplan B, et al.
    Rapid transcriptome changes induced by cytosolic Ca2+ transients reveal ABRE-related sequences as Ca2+-responsive cis elements in Arabidopsis.
    Plant Cell, 2006. 18(10): p. 2733-48
  10. Liu JX,Srivastava R,Che P,Howell SH
    Salt stress responses in Arabidopsis utilize a signal transduction pathway related to endoplasmic reticulum stress signaling.
    Plant J., 2007. 51(5): p. 897-909
  11. Libault M,Wan J,Czechowski T,Udvardi M,Stacey G
    Identification of 118 Arabidopsis transcription factor and 30 ubiquitin-ligase genes responding to chitin, a plant-defense elicitor.
    Mol. Plant Microbe Interact., 2007. 20(8): p. 900-11
  12. Peng M,Bi YM,Zhu T,Rothstein SJ
    Genome-wide analysis of Arabidopsis responsive transcriptome to nitrogen limitation and its regulation by the ubiquitin ligase gene NLA.
    Plant Mol. Biol., 2007. 65(6): p. 775-97
  13. Ramel F, et al.
    Genome-wide interacting effects of sucrose and herbicide-mediated stress in Arabidopsis thaliana: novel insights into atrazine toxicity and sucrose-induced tolerance.
    BMC Genomics, 2007. 8: p. 450
  14. Lu DP,Christopher DA
    Endoplasmic reticulum stress activates the expression of a sub-group of protein disulfide isomerase genes and AtbZIP60 modulates the response in Arabidopsis thaliana.
    Mol. Genet. Genomics, 2008. 280(3): p. 199-210
  15. Ascencio-Ib
    Global analysis of Arabidopsis gene expression uncovers a complex array of changes impacting pathogen response and cell cycle during geminivirus infection.
    Plant Physiol., 2008. 148(1): p. 436-54
  16. Wang Y, et al.
    Transcriptome analyses show changes in gene expression to accompany pollen germination and tube growth in Arabidopsis.
    Plant Physiol., 2008. 148(3): p. 1201-11
  17. Mitsuya Y, et al.
    Spermine signaling plays a significant role in the defense response of Arabidopsis thaliana to cucumber mosaic virus.
    J. Plant Physiol., 2009. 166(6): p. 626-43
  18. Iwata Y,Fedoroff NV,Koizumi N
    Arabidopsis bZIP60 is a proteolysis-activated transcription factor involved in the endoplasmic reticulum stress response.
    Plant Cell, 2008. 20(11): p. 3107-21
  19. Iwata Y,Yoneda M,Yanagawa Y,Koizumi N
    Characteristics of the nuclear form of the Arabidopsis transcription factor AtbZIP60 during the endoplasmic reticulum stress response.
    Biosci. Biotechnol. Biochem., 2009. 73(4): p. 865-9
  20. Iwata Y,Fedoroff NV,Koizumi N
    The Arabidopsis membrane-bound transcription factor AtbZIP60 is a novel plant-specific endoplasmic reticulum stress transducer.
    Plant Signal Behav, 2009. 4(6): p. 514-6
  21. Liu JX,Howell SH
    bZIP28 and NF-Y transcription factors are activated by ER stress and assemble into a transcriptional complex to regulate stress response genes in Arabidopsis.
    Plant Cell, 2010. 22(3): p. 782-96
  22. Wang M,Xu Q,Yu J,Yuan M
    The putative Arabidopsis zinc transporter ZTP29 is involved in the response to salt stress.
    Plant Mol. Biol., 2010. 73(4-5): p. 467-79
  23. Iwata Y,Nishino T,Takayama S,Koizumi N
    Characterization of a plant-specific gene induced by endoplasmic reticulum stress in Arabidopsis thaliana.
    Biosci. Biotechnol. Biochem., 2010. 74(10): p. 2087-91
  24. Katoh H,Fujita K,Takuhara Y,Ogawa A,Suzuki S
    ER stress-induced protein, VIGG, disturbs plant cation homeostasis, which is correlated with growth retardation and robustness to ER stress.
    Biochem. Biophys. Res. Commun., 2011. 405(3): p. 514-20
  25. Deng Y, et al.
    Heat induces the splicing by IRE1 of a mRNA encoding a transcription factor involved in the unfolded protein response in Arabidopsis.
    Proc. Natl. Acad. Sci. U.S.A., 2011. 108(17): p. 7247-52
  26. Moreno AA,Orellana A
    The physiological role of the unfolded protein response in plants.
    Biol. Res., 2011. 44(1): p. 75-80
  27. Iwata Y,Lee MH,Koizumi N
    Analysis of a transcription factor using transient assay in Arabidopsis protoplasts.
    Methods Mol. Biol., 2011. 754: p. 107-17
  28. Hayashi S,Wakasa Y,Takahashi H,Kawakatsu T,Takaiwa F
    Signal transduction by IRE1-mediated splicing of bZIP50 and other stress sensors in the endoplasmic reticulum stress response of rice.
    Plant J., 2012. 69(6): p. 946-56
  29. Wang B,Zheng J,Liu Y,Wang J,Wang G
    Cloning and characterization of the stress-induced bZIP gene ZmbZIP60 from maize.
    Mol. Biol. Rep., 2012. 39(5): p. 6319-27
  30. Qiang X,Zechmann B,Reitz MU,Kogel KH,Sch
    The mutualistic fungus Piriformospora indica colonizes Arabidopsis roots by inducing an endoplasmic reticulum stress-triggered caspase-dependent cell death.
    Plant Cell, 2012. 24(2): p. 794-809
  31. Nagashima Y, et al.
    Arabidopsis IRE1 catalyses unconventional splicing of bZIP60 mRNA to produce the active transcription factor.
    Sci Rep, 2011. 1: p. 29
  32. Moreno AA, et al.
    IRE1/bZIP60-mediated unfolded protein response plays distinct roles in plant immunity and abiotic stress responses.
    PLoS ONE, 2012. 7(2): p. e31944
  33. Humbert S,Zhong S,Deng Y,Howell SH,Rothstein SJ
    Alteration of the bZIP60/IRE1 pathway affects plant response to ER stress in Arabidopsis thaliana.
    PLoS ONE, 2012. 7(6): p. e39023
  34. Iwata Y,Koizumi N
    Plant transducers of the endoplasmic reticulum unfolded protein response.
    Trends Plant Sci., 2012. 17(12): p. 720-7
  35. Liu Y, et al.
    Degradation of the endoplasmic reticulum by autophagy during endoplasmic reticulum stress in Arabidopsis.
    Plant Cell, 2012. 24(11): p. 4635-51
  36. Wakasa Y,Hayashi S,Ozawa K,Takaiwa F
    Multiple roles of the ER stress sensor IRE1 demonstrated by gene targeting in rice.
    Sci Rep, 2012. 2: p. 944
  37. Tang W,Page M
    Transcription factor AtbZIP60 regulates expression of Ca2+ -dependent protein kinase genes in transgenic cells.
    Mol. Biol. Rep., 2013. 40(3): p. 2723-32
  38. Mishiba K, et al.
    Defects in IRE1 enhance cell death and fail to degrade mRNAs encoding secretory pathway proteins in the Arabidopsis unfolded protein response.
    Proc. Natl. Acad. Sci. U.S.A., 2013. 110(14): p. 5713-8
  39. Sun L, et al.
    The lumen-facing domain is important for the biological function and organelle-to-organelle movement of bZIP28 during ER stress in Arabidopsis.
    Mol Plant, 2013. 6(5): p. 1605-15
  40. Sun L, et al.
    The plant-specific transcription factor gene NAC103 is induced by bZIP60 through a new cis-regulatory element to modulate the unfolded protein response in Arabidopsis.
    Plant J., 2013. 76(2): p. 274-86
  41. Henriquez-Valencia C, et al.
    bZIP17 and bZIP60 Regulate the Expression of BiP3 and Other Salt Stress Responsive Genes in an UPR-Independent Manner in Arabidopsis thaliana.
    J. Cell. Biochem., 2015. 116(8): p. 1638-45
  42. Song ZT, et al.
    Transcription factor interaction with COMPASS-like complex regulates histone H3K4 trimethylation for specific gene expression in plants.
    Proc. Natl. Acad. Sci. U.S.A., 2015. 112(9): p. 2900-5
  43. Jin J, et al.
    An Arabidopsis Transcriptional Regulatory Map Reveals Distinct Functional and Evolutionary Features of Novel Transcription Factors.
    Mol. Biol. Evol., 2015. 32(7): p. 1767-73
  44. Nagashima Y,Iwata Y,Mishiba K,Koizumi N
    Arabidopsis tRNA ligase completes the cytoplasmic splicing of bZIP60 mRNA in the unfolded protein response.
    Biochem. Biophys. Res. Commun., 2016. 470(4): p. 941-6