Plant Transcription Factor Database
Previous version: v3.0
Transcription Factor Information
Basic Information | Signature Domain | Sequence | 
Basic Information? help Back to Top
TF ID AT1G70510.1
Common NameATK1, F24J13.8, KNAT2
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 TALE
Protein Properties Length: 310aa    MW: 35638.1 Da    PI: 4.6519
Description KNOTTED-like from Arabidopsis thaliana 2
Gene Model
Gene Model ID Type Source Coding Sequence
AT1G70510.1genomeTAIRView CDS
Signature Domain? help Back to Top
Signature Domain
No. Domain Score E-value Start End HMM Start HMM End
     Homeobox  21 knrypsaeereeLAkklgLterqVkvWFqNrRake 55 
                  k +yp++ ++  LA+++gL+++q+ +WF N+R ++
                  569*****************************885 PP

          ELK   1 ELKhqLlrKYsgyLgsLkqEFs 22 
                  +LK+qLlrK++++++sLk EFs
                  6********************8 PP

Protein Features ? help Back to Top
3D Structure
Database Entry ID E-value Start End InterPro ID Description
PROSITE profilePS512139.935208228IPR005539ELK domain
SMARTSM011884.2E-6208229IPR005539ELK domain
PfamPF037897.2E-9208229IPR005539ELK domain
PROSITE profilePS5007112.795228291IPR001356Homeobox domain
SMARTSM003891.4E-13230295IPR001356Homeobox domain
CDDcd000865.73E-12237292No hitNo description
PfamPF059202.0E-17248287IPR008422Homeobox KN domain
PROSITE patternPS000270266289IPR017970Homeobox, conserved site
Gene Ontology ? help Back to Top
GO Term GO Category GO Description
GO:0006355Biological Processregulation of transcription, DNA-templated
GO:0009723Biological Processresponse to ethylene
GO:0009736Biological Processcytokinin-activated signaling pathway
GO:0010094Biological Processspecification of carpel identity
GO:0005634Cellular Componentnucleus
GO:0003700Molecular Functiontranscription factor activity, sequence-specific DNA binding
GO:0005515Molecular Functionprotein binding
GO:0043565Molecular Functionsequence-specific DNA binding
Plant Ontology ? help Back to Top
PO Term PO Category PO Description
PO:0000013anatomycauline leaf
PO:0000037anatomyshoot apex
PO:0000226anatomyrib zone
PO:0000230anatomyinflorescence meristem
PO:0009009anatomyplant embryo
PO:0009025anatomyvascular leaf
PO:0009052anatomyflower pedicel
PO:0025022anatomycollective leaf structure
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:0007611developmental stagepetal differentiation and expansion stage
PO:0007616developmental stageflowering stage
Sequence ? help Back to Top
Protein Sequence    Length: 310 aa     Download sequence    Send to blast
Expression -- Microarray ? help Back to Top
Source ID E-value
Expression AtlasAT1G70510-
Expression -- Description ? help Back to Top
Source Description
UniprotTISSUE SPECIFICITY: Expressed predominantly in shoot apices of seedlings, in the receptacle and developing pistil of flowers and in axillary buds of inflorescence stems.
Functional Description ? help Back to Top
Source Description
TAIRA member of class I knotted1-like homeobox gene family (together with KNAT1). Similar to the knotted1 (kn1) homeobox gene of maize. KNAT2 acts synergistically with cytokinins and antagonistically with ethylene based on ectopic expression studies in different mutant backgrounds and hormone treatments. In addition, KNAT2 is negatively regulated by AS and YABBY genes. KNAT2 is strongly expressed in the shoot apex of seedlings, while in mature plants the gene is primarily expressed in flowers and inflorescence stems.
UniProtMay play a role in meristem function, and may be involved in maintaining cells in an undifferentiated, meristematic state. Probably binds to the DNA sequence 5'-TGAC-3'.
Function -- GeneRIF ? help Back to Top
  1. KNAT6 and KNAT2 expression is restricted by BP and PNY to promote correct inflorescence development
    [PMID: 18390591]
  2. Results suggested that the late-flowering phenotype of the as1-1 and as2-1 mutants might have been caused by the ectopic expression of KNAT126 triple mutation.
    [PMID: 19891706]
  3. The data indicate that the ATH1-KNAT2 complex acts redundantly with KNAT6, both of which are negatively regulated by BP during pedicel development.
    [PMID: 22140242]
  4. Arabidopsis ASYMMETRIC LEAVES complex physically interacts with PRC2 and recruits this complex to the homeobox genes BREVIPEDICELLUS and KNAT2 to stably silence these stem cell regulators in differentiating leaves.
    [PMID: 23468429]
  5. Arabidopsis plants harboring loss-of-function KNOX2 alleles exhibit impaired differentiation of all aerial organs and have highly complex leaves, phenocopying gain-of-function KNOX1 alleles
    [PMID: 25671434]
  6. our results indicate that BP interacts with the chromatin remodeling factor BRM to regulate the expression of KNAT2 and KNAT6 in control of inflorescence architecture.
    [PMID: 25822547]
Cis-element ? help Back to Top
Regulation -- PlantRegMap ? help Back to Top
Source Upstream Regulator Target Gene
Regulation -- ATRM (Manually Curated Upstream Regulators) ? help Back to Top
Source Upstream Regulator (A: Activate/R: Repress)
ATRM AT1G65620 (R), AT2G01570 (A), AT2G37630 (R), AT5G66870 (R)
Regulation -- ATRM (Manually Curated Target Genes) ? help Back to Top
Source Target Gene (A: Activate/R: Repress)
ATRM AT4G18960(A)
Regulation -- Hormone ? help Back to Top
Source Hormone
Interaction ? help Back to Top
Source Intact With
IntActSearch P46640
Phenotype -- Mutation ? help Back to Top
Source ID
T-DNA ExpressAT1G70510
Annotation -- Nucleotide ? help Back to Top
Source Hit ID E-value Description
GenBankATU141750.0U14175.1 Arabidopsis thaliana clone KNAT2 knotted-like homeobox protein gene, partial cds.
Annotation -- Protein ? help Back to Top
Source Hit ID E-value Description
RefseqNP_177208.20.0homeobox protein knotted-1-like 2
SwissprotP466400.0KNAT2_ARATH; Homeobox protein knotted-1-like 2
TrEMBLD7KY360.0D7KY36_ARALL; Putative uncharacterized protein
STRINGAT1G70510.10.0(Arabidopsis thaliana)
Orthologous Group ? help Back to Top
LineageOrthologous Group IDTaxa NumberGene Number
Representative plantOGRP16717148
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. Byrne ME, et al.
    Asymmetric leaves1 mediates leaf patterning and stem cell function in Arabidopsis.
    Nature, 2000 Dec 21-28. 408(6815): p. 967-71
  3. Tsiantis M
    Control of shoot cell fate: beyond homeoboxes.
    Plant Cell, 2001. 13(4): p. 733-8
  4. Semiarti E, et al.
    The ASYMMETRIC LEAVES2 gene of Arabidopsis thaliana regulates formation of a symmetric lamina, establishment of venation and repression of meristem-related homeobox genes in leaves.
    Development, 2001. 128(10): p. 1771-83
  5. Pautot V, et al.
    KNAT2: evidence for a link between knotted-like genes and carpel development.
    Plant Cell, 2001. 13(8): p. 1719-34
  6. Bellaoui M, et al.
    The Arabidopsis BELL1 and KNOX TALE homeodomain proteins interact through a domain conserved between plants and animals.
    Plant Cell, 2001. 13(11): p. 2455-70
  7. Venglat SP, et al.
    The homeobox gene BREVIPEDICELLUS is a key regulator of inflorescence architecture in Arabidopsis.
    Proc. Natl. Acad. Sci. U.S.A., 2002. 99(7): p. 4730-5
  8. Byrne ME,Simorowski J,Martienssen RA
    ASYMMETRIC LEAVES1 reveals knox gene redundancy in Arabidopsis.
    Development, 2002. 129(8): p. 1957-65
  9. Hamant O, et al.
    The KNAT2 homeodomain protein interacts with ethylene and cytokinin signaling.
    Plant Physiol., 2002. 130(2): p. 657-65
  10. Barley R,Waites R
    Plant meristems: the interplay of KNOX and gibberellins.
    Curr. Biol., 2002. 12(20): p. R696-8
  11. Kumaran MK,Bowman JL,Sundaresan V
    YABBY polarity genes mediate the repression of KNOX homeobox genes in Arabidopsis.
    Plant Cell, 2002. 14(11): p. 2761-70
  12. Ha CM, et al.
    The BLADE-ON-PETIOLE 1 gene controls leaf pattern formation through the modulation of meristematic activity in Arabidopsis.
    Development, 2003. 130(1): p. 161-72
  13. Haberer G,Erschadi S,Torres-Ruiz RA
    The Arabidopsis gene PEPINO/PASTICCINO2 is required for proliferation control of meristematic and non-meristematic cells and encodes a putative anti-phosphatase.
    Dev. Genes Evol., 2002. 212(11): p. 542-50
  14. Micol JL,Hake S
    The development of plant leaves.
    Plant Physiol., 2003. 131(2): p. 389-94
  15. Harrar Y,Bellec Y,Bellini C,Faure JD
    Hormonal control of cell proliferation requires PASTICCINO genes.
    Plant Physiol., 2003. 132(3): p. 1217-27
  16. Lin WC,Shuai B,Springer PS
    The Arabidopsis LATERAL ORGAN BOUNDARIES-domain gene ASYMMETRIC LEAVES2 functions in the repression of KNOX gene expression and in adaxial-abaxial patterning.
    Plant Cell, 2003. 15(10): p. 2241-52
  17. Bae MS,Cho EJ,Choi EY,Park OK
    Analysis of the Arabidopsis nuclear proteome and its response to cold stress.
    Plant J., 2003. 36(5): p. 652-63
  18. Katz A,Oliva M,Mosquna A,Hakim O,Ohad N
    FIE and CURLY LEAF polycomb proteins interact in the regulation of homeobox gene expression during sporophyte development.
    Plant J., 2004. 37(5): p. 707-19
  19. Hackbusch J,Richter K,Müller J,Salamini F,Uhrig JF
    A central role of Arabidopsis thaliana ovate family proteins in networking and subcellular localization of 3-aa loop extension homeodomain proteins.
    Proc. Natl. Acad. Sci. U.S.A., 2005. 102(13): p. 4908-12
  20. Kwon CS,Chen C,Wagner D
    WUSCHEL is a primary target for transcriptional regulation by SPLAYED in dynamic control of stem cell fate in Arabidopsis.
    Genes Dev., 2005. 19(8): p. 992-1003
  21. Yanai O, et al.
    Arabidopsis KNOXI proteins activate cytokinin biosynthesis.
    Curr. Biol., 2005. 15(17): p. 1566-71
  22. Ledbedeva OV,Ezhova TA,Melzer S
    [Gene TAENIATA is a novel negative regulator of the Arabidopsis thaliana homeobox genes KNAT1, KNAT2, KNAT6, and STM].
    Genetika, 2005. 41(8): p. 1068-74
  23. Chua YL,Channeli
    The bromodomain protein GTE6 controls leaf development in Arabidopsis by histone acetylation at ASYMMETRIC LEAVES1.
    Genes Dev., 2005. 19(18): p. 2245-54
  24. Belles-Boix E, et al.
    KNAT6: an Arabidopsis homeobox gene involved in meristem activity and organ separation.
    Plant Cell, 2006. 18(8): p. 1900-7
  25. Okushima Y,Fukaki H,Onoda M,Theologis A,Tasaka M
    ARF7 and ARF19 regulate lateral root formation via direct activation of LBD/ASL genes in Arabidopsis.
    Plant Cell, 2007. 19(1): p. 118-30
  26. Krupkov
    The TUMOROUS SHOOT DEVELOPMENT2 gene of Arabidopsis encoding a putative methyltransferase is required for cell adhesion and co-ordinated plant development.
    Plant J., 2007. 50(4): p. 735-50
  27. Scofield S,Dewitte W,Murray JA
    The KNOX gene SHOOT MERISTEMLESS is required for the development of reproductive meristematic tissues in Arabidopsis.
    Plant J., 2007. 50(5): p. 767-81
  28. Alonso-Cantabrana H, et al.
    Common regulatory networks in leaf and fruit patterning revealed by mutations in the Arabidopsis ASYMMETRIC LEAVES1 gene.
    Development, 2007. 134(14): p. 2663-71
  29. Ha CM,Jun JH,Nam HG,Fletcher JC
    BLADE-ON-PETIOLE 1 and 2 control Arabidopsis lateral organ fate through regulation of LOB domain and adaxial-abaxial polarity genes.
    Plant Cell, 2007. 19(6): p. 1809-25
  30. Kumar R, et al.
    The Arabidopsis BEL1-LIKE HOMEODOMAIN proteins SAW1 and SAW2 act redundantly to regulate KNOX expression spatially in leaf margins.
    Plant Cell, 2007. 19(9): p. 2719-35
  31. Guo M,Thomas J,Collins G,Timmermans MC
    Direct repression of KNOX loci by the ASYMMETRIC LEAVES1 complex of Arabidopsis.
    Plant Cell, 2008. 20(1): p. 48-58
  32. Ragni L,Belles-Boix E,Günl M,Pautot V
    Interaction of KNAT6 and KNAT2 with BREVIPEDICELLUS and PENNYWISE in Arabidopsis inflorescences.
    Plant Cell, 2008. 20(4): p. 888-900
  33. Hou X, et al.
    Global identification of DELLA target genes during Arabidopsis flower development.
    Plant Physiol., 2008. 147(3): p. 1126-42
  34. Yu L,Patibanda V,Smith HM
    A novel role of BELL1-like homeobox genes, PENNYWISE and POUND-FOOLISH, in floral patterning.
    Planta, 2009. 229(3): p. 693-707
  35. Soyano T,Thitamadee S,Machida Y,Chua NH
    ASYMMETRIC LEAVES2-LIKE19/LATERAL ORGAN BOUNDARIES DOMAIN30 and ASL20/LBD18 regulate tracheary element differentiation in Arabidopsis.
    Plant Cell, 2008. 20(12): p. 3359-73
  36. Scofield S,Dewitte W,Murray JA
    A model for Arabidopsis class-1 KNOX gene function.
    Plant Signal Behav, 2008. 3(4): p. 257-9
  37. Ikezaki M, et al.
    Genetic networks regulated by ASYMMETRIC LEAVES1 (AS1) and AS2 in leaf development in Arabidopsis thaliana: KNOX genes control five morphological events.
    Plant J., 2010. 61(1): p. 70-82
  38. Tabata R, et al.
    Arabidopsis auxin response factor6 and 8 regulate jasmonic acid biosynthesis and floral organ development via repression of class 1 KNOX genes.
    Plant Cell Physiol., 2010. 51(1): p. 164-75
  39. Li Y,Pi L,Huang H,Xu L
    ATH1 and KNAT2 proteins act together in regulation of plant inflorescence architecture.
    J. Exp. Bot., 2012. 63(3): p. 1423-33
  40. Butenko MA,Shi CL,Aalen RB
    KNAT1, KNAT2 and KNAT6 act downstream in the IDA-HAE/HSL2 signaling pathway to regulate floral organ abscission.
    Plant Signal Behav, 2012. 7(1): p. 135-8
  41. Luo M, et al.
    Histone deacetylase HDA6 is functionally associated with AS1 in repression of KNOX genes in arabidopsis.
    PLoS Genet., 2012. 8(12): p. e1003114
  42. Lodha M,Marco CF,Timmermans MC
    The ASYMMETRIC LEAVES complex maintains repression of KNOX homeobox genes via direct recruitment of Polycomb-repressive complex2.
    Genes Dev., 2013. 27(6): p. 596-601
  43. Scofield S,Dewitte W,Nieuwland J,Murray JA
    The Arabidopsis homeobox gene SHOOT MERISTEMLESS has cellular and meristem-organisational roles with differential requirements for cytokinin and CYCD3 activity.
    Plant J., 2013. 75(1): p. 53-66
  44. Stammler A, et al.
    Duplicated STM-like KNOX I genes act in floral meristem activity in Eschscholzia californica (Papaveraceae).
    Dev. Genes Evol., 2013. 223(5): p. 289-301
  45. Furumizu C,Alvarez JP,Sakakibara K,Bowman JL
    Antagonistic roles for KNOX1 and KNOX2 genes in patterning the land plant body plan following an ancient gene duplication.
    PLoS Genet., 2015. 11(2): p. e1004980
  46. 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
  47. Zhao M, et al.
    Arabidopsis BREVIPEDICELLUS interacts with the SWI2/SNF2 chromatin remodeling ATPase BRAHMA to regulate KNAT2 and KNAT6 expression in control of inflorescence architecture.
    PLoS Genet., 2015. 11(3): p. e1005125
  48. Dockx J, et al.
    The homeobox gene ATK1 of Arabidopsis thaliana is expressed in the shoot apex of the seedling and in flowers and inflorescence stems of mature plants.
    Plant Mol. Biol., 1995. 28(4): p. 723-37
  49. Lincoln C,Long J,Yamaguchi J,Serikawa K,Hake S
    A knotted1-like homeobox gene in Arabidopsis is expressed in the vegetative meristem and dramatically alters leaf morphology when overexpressed in transgenic plants.
    Plant Cell, 1994. 6(12): p. 1859-76
  50. Granger CL,Callos JD,Medford JI
    Isolation of an Arabidopsis homologue of the maize homeobox Knotted-1 gene.
    Plant Mol. Biol., 1996. 31(2): p. 373-8