PlantRegMap/PlantTFDB v5.0
Plant Transcription Factor Database
Previous version: v3.0 v4.0
Transcription Factor Information
Basic Information | Signature Domain | Sequence | 
Basic Information? help Back to Top
TF ID AT4G21330.1
Common NameBHLH22, DYT1, EN49, T6K22.60
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 bHLH
Protein Properties Length: 207aa    MW: 24021.6 Da    PI: 4.7997
Description bHLH family protein
Gene Model
Gene Model ID Type Source Coding Sequence
AT4G21330.1genomeTAIRView CDS
Signature Domain? help Back to Top
Signature Domain
No. Domain Score E-value Start End HMM Start HMM End
          HLH  7 erErrRRdriNsafeeLrellPkaskapskKlsKaeiLekAveYIksLq 55
                 e+ErrRR+++  ++  Lr+++P       + ++Ka+i+e A+ YI +Lq
                 79*********************8.....69****************98 PP

Protein Features ? help Back to Top
3D Structure
Database Entry ID E-value Start End InterPro ID Description
SuperFamilySSF474593.14E-142593IPR011598Myc-type, basic helix-loop-helix (bHLH) domain
CDDcd000831.04E-102682No hitNo description
PROSITE profilePS5088813.962877IPR011598Myc-type, basic helix-loop-helix (bHLH) domain
SMARTSM003537.6E-123483IPR011598Myc-type, basic helix-loop-helix (bHLH) domain
Gene3DG3DSA:, basic helix-loop-helix (bHLH) domain
PfamPF000107.6E-83578IPR011598Myc-type, basic helix-loop-helix (bHLH) domain
Gene Ontology ? help Back to Top
GO Term GO Category GO Description
GO:0006355Biological Processregulation of transcription, DNA-templated
GO:0048658Biological Processanther wall tapetum development
GO:0005634Cellular Componentnucleus
GO:0003700Molecular Functiontranscription factor activity, sequence-specific DNA binding
GO:0043565Molecular Functionsequence-specific DNA binding
GO:0046983Molecular Functionprotein dimerization activity
Plant Ontology ? help Back to Top
PO Term PO Category PO Description
PO:0000229anatomyflower meristem
PO:0006089anatomyanther primordium
PO:0009071anatomyanther wall tapetum
Sequence ? help Back to Top
Protein Sequence    Length: 207 aa     Download sequence    Send to blast
Nucleic Localization Signal ? help Back to Top
No. Start End Sequence
Expression -- Microarray ? help Back to Top
Source ID E-value
Expression AtlasAT4G21330-
Expression -- Description ? help Back to Top
Source Description
UniprotDEVELOPMENTAL STAGE: First observed in floral meristem and early anther primordia. Later detected in archesporial cells. From stage 4 to early stage 5, weakly expressed in precursors of the middle layer, tapetum and meiocytes. Strongly expressed in the tapetum from late anther stage 5 to early stage 6, and at a lower level in meiocytes. {ECO:0000269|PubMed:16831835}.
UniprotTISSUE SPECIFICITY: Mostly expressed in anthers, and, to a lower extent, in young inflorescences undergoing meiosis and siliques. {ECO:0000269|PubMed:16831835}.
Functional Description ? help Back to Top
Source Description
TAIREncodes a bHLH transcription factor strongly expressed in the tapetum from late anther stage 5 to early stage 6, and at a lower level in meiocytes. dyt1 mutant exhibits abnormal anther morphology beginning at anther stage 4. DYT1 acts downstream of SPL/NZZ and EMS1/EXS , and is required for normal expression of AMS, MS1 and other tapetum preferential genes.
UniProtTranscription factor. Involved in the control of tapetum development. Required for male fertility and pollen differentiation, especially during callose deposition. {ECO:0000269|PubMed:16831835}.
Function -- GeneRIF ? help Back to Top
  1. DYT1 encodes a putative bHLH transcription factor and is a component of a genetic network that controls anther development and function. [DYT1]
    [PMID: 16831835]
  2. DYT1, TDF1 and AMS function in early tapetum development, while MS188 and MS1 are important for late tapetum development.
    [PMID: 21957980]
  3. DYT1 positively regulates genes for lipid metabolism, cell-wall modification and secondary metabolism that are important for pollen development.
    [PMID: 22775442]
  4. DYT1 regulates the expression of AMS, MS188/MYB80, TEK and MS1 for pollen wall formation, primarily via TDF1.
    [PMID: 25284309]
  5. DYT1 forms complexes with bHLH010, bHLH089 and bHLH091 subsequently affecting the correct expression of many DYT1 target genes.
    [PMID: 26216374]
  6. Data said that the normal spatio-temporal subcellular localization of DYT1 is important for DYT1 function and/or that the BIF domains from different bHLH members might be functionally distinct.
    [PMID: 27113773]
  7. Here the authors demonstrate abnormal risk taking in DYT1 dystonia patients, which is correlated with disease severity, thereby supporting striatal plasticity in shaping choice behavior in humans.
    [PMID: 27249418]
  8. DYT1 dystonia is a network disorder, with crucial nodes in sensory-motor integration of posterior parietal structures.
    [PMID: 27453152]
  9. we show the behavioral and molecular consequences of stressing the endoplasmic reticulum in DYT1 mice by increasing the amount of misfolded proteins
    [PMID: 27707963]
  10. The results of this study suggested that motor impairments in DYT1 KI animals only exhibit when the sensory stimuli are strongly altered during performance.
    [PMID: 27769743]
  11. Mice with the DYT1 dystonia mutation exhibit an enhanced response to mu receptor activation, dependent on selective receptor gene upregulation..
    [PMID: 29150865]
  12. Early onset torsion dystonia (DYT1) patient fibroblasts with heterozygous DYT1 mutant alleles showed decreased herpes simplex virus 1 plaque formation as compared to controls.
    [PMID: 29396398]
Binding Motif ? help Back to Top
Motif ID Method Source Motif file
Motif logo
Cis-element ? help Back to Top
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 AT2G16910(A), AT3G23770(A), AT3G28470(A), AT3G51590(A), AT5G22260(A)
Interaction ? help Back to Top
Source Intact With
BioGRIDAT4G21330, AT1G06170
Phenotype -- Mutation ? help Back to Top
Source ID
T-DNA ExpressAT4G21330
Annotation -- Nucleotide ? help Back to Top
Source Hit ID E-value Description
GenBankAL0311871e-136AL031187.1 Arabidopsis thaliana DNA chromosome 4, BAC clone T6K22 (ESSAII project).
GenBankAL1615541e-136AL161554.2 Arabidopsis thaliana DNA chromosome 4, contig fragment No. 54.
GenBankCP0026871e-136CP002687.1 Arabidopsis thaliana chromosome 4 sequence.
GenBankEF6370831e-136EF637083.1 Arabidopsis thaliana ecotype Cvi-0 S-locus region genomic sequence.
Annotation -- Protein ? help Back to Top
Source Hit ID E-value Description
RefseqNP_193864.11e-151basic helix-loop-helix (bHLH) DNA-binding superfamily protein
SwissprotO819001e-152DYT1_ARATH; Transcription factor DYT1
STRINGAT4G21330.11e-150(Arabidopsis thaliana)
Orthologous Group ? help Back to Top
LineageOrthologous Group IDTaxa NumberGene Number
Representative plantOGRP1030948
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. Heim MA, et al.
    The basic helix-loop-helix transcription factor family in plants: a genome-wide study of protein structure and functional diversity.
    Mol. Biol. Evol., 2003. 20(5): p. 735-47
  3. Toledo-Ortiz G,Huq E,Quail PH
    The Arabidopsis basic/helix-loop-helix transcription factor family.
    Plant Cell, 2003. 15(8): p. 1749-70
  4. Bailey PC, et al.
    Update on the basic helix-loop-helix transcription factor gene family in Arabidopsis thaliana.
    Plant Cell, 2003. 15(11): p. 2497-502
  5. Bauer P, et al.
    Analysis of sequence, map position, and gene expression reveals conserved essential genes for iron uptake in Arabidopsis and tomato.
    Plant Physiol., 2004. 136(4): p. 4169-83
  6. Zhang W, et al.
    Regulation of Arabidopsis tapetum development and function by DYSFUNCTIONAL TAPETUM1 (DYT1) encoding a putative bHLH transcription factor.
    Development, 2006. 133(16): p. 3085-95
  7. Li N, et al.
    The rice tapetum degeneration retardation gene is required for tapetum degradation and anther development.
    Plant Cell, 2006. 18(11): p. 2999-3014
  8. Tang C, et al.
    The evolution of selfing in Arabidopsis thaliana.
    Science, 2007. 317(5841): p. 1070-2
  9. Wijeratne AJ, et al.
    Differential gene expression in Arabidopsis wild-type and mutant anthers: insights into anther cell differentiation and regulatory networks.
    Plant J., 2007. 52(1): p. 14-29
  10. Xing S,Zachgo S
    ROXY1 and ROXY2, two Arabidopsis glutaredoxin genes, are required for anther development.
    Plant J., 2008. 53(5): p. 790-801
  11. Zhu J, et al.
    Defective in Tapetal development and function 1 is essential for anther development and tapetal function for microspore maturation in Arabidopsis.
    Plant J., 2008. 55(2): p. 266-77
  12. Xu J, et al.
    The ABORTED MICROSPORES regulatory network is required for postmeiotic male reproductive development in Arabidopsis thaliana.
    Plant Cell, 2010. 22(1): p. 91-107
  13. Skinner MK,Rawls A,Wilson-Rawls J,Roalson EH
    Basic helix-loop-helix transcription factor gene family phylogenetics and nomenclature.
    Differentiation, 2010. 80(1): p. 1-8
  14. Zhu J,Lou Y,Xu X,Yang ZN
    A genetic pathway for tapetum development and function in Arabidopsis.
    J Integr Plant Biol, 2011. 53(11): p. 892-900
  15. Wang XT,Yuan C,Yuan TT,Cui SJ
    The Arabidopsis LFR gene is required for the formation of anther cell layers and normal expression of key regulatory genes.
    Mol Plant, 2012. 5(5): p. 993-1000
  16. Feng B, et al.
    Regulation of the Arabidopsis anther transcriptome by DYT1 for pollen development.
    Plant J., 2012. 72(4): p. 612-24
  17. Jiang J,Zhang Z,Cao J
    Pollen wall development: the associated enzymes and metabolic pathways.
    Plant Biol (Stuttg), 2013. 15(2): p. 249-63
  18. Gu JN, et al.
    DYT1 directly regulates the expression of TDF1 for tapetum development and pollen wall formation in Arabidopsis.
    Plant J., 2014. 80(6): p. 1005-13
  19. Qian H, et al.
    Trace concentrations of imazethapyr (IM) affect floral organs development and reproduction in Arabidopsis thaliana: IM-induced inhibition of key genes regulating anther and pollen biosynthesis.
    Ecotoxicology, 2015. 24(1): p. 163-71
  20. 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
  21. Shumin Z, et al.
    One novel cis-element is essential for correct DYSFUNCTIONAL TAPETUM 1 (DYT1) expression in Arabidopsis thaliana.
    Plant Cell Rep., 2015. 34(10): p. 1773-80
  22. Zhu E, et al.
    The DYT1-interacting proteins bHLH010, bHLH089 and bHLH091 are redundantly required for Arabidopsis anther development and transcriptome.
    Plant J., 2015. 83(6): p. 976-90
  23. Cui J, et al.
    Feedback Regulation of DYT1 by Interactions with Downstream bHLH Factors Promotes DYT1 Nuclear Localization and Anther Development.
    Plant Cell, 2016. 28(5): p. 1078-93
  24. Arkadir D, et al.
    DYT1 dystonia increases risk taking in humans.
    Elife, 2017.
  25. Premi E, et al.
    Functional Connectivity Networks in Asymptomatic and Symptomatic DYT1 Carriers.
    Mov. Disord., 2016. 31(11): p. 1739-1743
  26. Beauvais G, et al.
    Disruption of Protein Processing in the Endoplasmic Reticulum of DYT1 Knock-in Mice Implicates Novel Pathways in Dystonia Pathogenesis.
    J. Neurosci., 2016. 36(40): p. 10245-10256
  27. Richter F,Gerstenberger J,Bauer A,Liang CC,Richter A
    Sensorimotor tests unmask a phenotype in the DYT1 knock-in mouse model of dystonia.
    Behav. Brain Res., 2017. 317: p. 536-541
  28. Zhou S, et al.
    Function Identification of the Nucleotides in Key cis-Element of DYSFUNCTIONAL TAPETUM1 (DYT1) Promoter.
    Front Plant Sci, 2017. 8: p. 153
  29. Li DD,Xue JS,Zhu J,Yang ZN
    Gene Regulatory Network for Tapetum Development in Arabidopsis thaliana.
    Front Plant Sci, 2017. 8: p. 1559
  30. Ponterio G, et al.
    Enhanced mu opioid receptor-dependent opioidergic modulation of striatal cholinergic transmission in DYT1 dystonia.
    Mov. Disord., 2018. 33(2): p. 310-320
  31. Gy├Ârgy B, et al.
    Mutant torsinA in the heterozygous DYT1 state compromises HSV propagation in infected neurons and fibroblasts.
    Sci Rep, 2018. 8(1): p. 2324