Plant Transcription Factor Database
Previous version: v3.0
Transcription Factor Information
Basic Information | Signature Domain | Sequence | 
Basic Information? help Back to Top
TF ID AT4G27330.1
Common NameF27G19.11, NZZ, SPL
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 NZZ/SPL
Protein Properties Length: 314aa    MW: 34078.1 Da    PI: 9.4743
Description sporocyteless (SPL)
Gene Model
Gene Model ID Type Source Coding Sequence
AT4G27330.1genomeTAIRView CDS
Signature Domain? help Back to Top
Signature Domain
No. Domain Score E-value Start End HMM Start HMM End
       NOZZLE   1 matslffmstdqnsvrnpnellrntrlvvnssgeirtetkksrgrkpgsktaqqkqkkptlrgmgvaklerfiieeekkklvvatvgdtssvaaisnt 98 
                  9************************************************************************************************* PP

       NOZZLE  99 atrlpvpvdrgvvlqgfpsslgssrilcggvgsgqvmidpvispwgfvetsatthelssisnpqmynassnnrcdtcfkkkrldgdqnnvvrsngggf 196
                  ************************************************************************************************** PP

       NOZZLE 197 skytmipppmngydeyllqsdhhqrsqgflydqriaraasvsaasasinpyfneatnltgsreefgsvlegnprngsrgvkeyeffpgkydervskva 294
                  ************************************************************************************************** PP

       NOZZLE 295 kvaslvgdcspntidlslkl 314
                  *******************8 PP

Protein Features ? help Back to Top
3D Structure
Database Entry ID E-value Start End InterPro ID Description
PfamPF087441.1E-2023314IPR014855Plant transcription factor NOZZLE
Gene Ontology ? help Back to Top
GO Term GO Category GO Description
GO:0006355Biological Processregulation of transcription, DNA-templated
GO:0048653Biological Processanther development
GO:0005634Cellular Componentnucleus
GO:0003700Molecular Functiontranscription factor activity, sequence-specific DNA binding
GO:0005515Molecular Functionprotein binding
Plant Ontology ? help Back to Top
PO Term PO Category PO Description
PO:0009009anatomyplant embryo
PO:0025022anatomycollective leaf structure
PO:0001078developmental stageplant embryo cotyledonary stage
PO:0007611developmental stagepetal differentiation and expansion stage
PO:0007616developmental stageflowering stage
Sequence ? help Back to Top
Protein Sequence    Length: 314 aa     Download sequence    Send to blast
Expression -- Microarray ? help Back to Top
Source ID E-value
Expression AtlasAT4G27330-
Expression -- Description ? help Back to Top
Source Description
UniprotDEVELOPMENTAL STAGE: Expressed during microsporogenesis and megasoprogenesis (PubMed:10465788). Expressed during ovule development (PubMed:25378179). {ECO:0000269|PubMed:10465788, ECO:0000269|PubMed:25378179}.
UniprotTISSUE SPECIFICITY: Expressed in flower buds (PubMed:10465788, PubMed:10500234). Not found in leaves, siliques and stems (PubMed:10465788). Detected in rosette leaves, stem tissue and seedlings (PubMed:10500234). {ECO:0000269|PubMed:10465788, ECO:0000269|PubMed:10500234}.
Functional Description ? help Back to Top
Source Description
TAIREncodes a putative transcription factor that is required for the initiation of both micro- and megagametogenesis and is expressed in the sporogenous tissue of the anther and the ovule. SPL is a chalaza identity gene that share overlapping functions in establishing the prospective chalaza of the ovule. It also plays a central role in patterning both the proximal-distal and the adaxial-abaxial axes in the ovule and generally interacts with YABBY proteins in vitro. Mutant is defective in the differentiation of primary sporogenous cells into microsporocytes, and does not properly form the anther wall. Regulator of anther cell differenctiation
UniProtTranscriptional regulator of sporocyte development (PubMed:10465788). Acts as an adapter-like transcriptional repressor recruiting TPL/TPR corepressors to inhibit TCP transcription factors (PubMed:25378179). Required for nucellus and embryo sac development (PubMed:10500234). Plays a central role in patterning both the proximal-distal and the adaxial-abaxial axes during ovule development (PubMed:12183381). Involved in establishing the prospective chalaza of the ovule and in controlling the cell number and the length of the funiculus, and is r equired for the development of the integuments (PubMed:10976054). Required, with BEL1, for cytokinin-induced PIN1 expression in ovules (PubMed:22786869). Involved in controlling stamen identity (PubMed:19726570). May also regulate the morphology of lateral organs by repressing auxin production (PubMed:18557819). {ECO:0000269|PubMed:10465788, ECO:0000269|PubMed:10500234, ECO:0000269|PubMed:10976054, ECO:0000269|PubMed:12183381, ECO:0000269|PubMed:18557819, ECO:0000269|PubMed:19726570, ECO:0000269|PubMed:22786869, ECO:0000269|PubMed:25378179}.
Function -- GeneRIF ? help Back to Top
  1. Nozzle protein may coordinate proximal-distal and adaxial-abaxial polarity establishment in the developing ovule by interacting with YABBY proteins in vitro in Arabidopsis.
    [PMID: 15299139]
  2. The cell-free method was used to synthesize SPL as a correctly folded and functional zinc-binding protein.
    [PMID: 17146616]
  3. SPL regulates auxin homeostasis by repressing the transcription of YUC2 and YUC6 and participates in lateral organ morphogenesis.
    [PMID: 18557819]
  4. SPL/NZZ is not only essential for early anther cell differentiation but also is engaged in controlling stamen identity in Arabidopsis
    [PMID: 19726570]
  5. results demonstrate that the miR156-regulated SPLs establish a direct link between developmental programming and trichome distribution
    [PMID: 20622149]
  6. The transcription factors BEL1 and SPL/NZZ, previously described as key regulators of ovule development, are needed for the auxin and cytokinin signaling pathways for the correct patterning of the ovule.
    [PMID: 22786869]
  7. SPL recruits topless/topless-related (TPL/TPR) co-repressors to inhibit the Cincinnata (CIN)-like Teosinte branched1/cycloidea/PCF (TCP) transcription factors.
    [PMID: 25378179]
  8. Data showed that SPOROCYTELESS (SPL) most likely functions as a transcription repressor by interacting with TCP family transcription factors and the TOPLESS (TPL) co-repressors.
    [PMID: 25527103]
Cis-element ? help Back to Top
Regulation -- Description ? help Back to Top
Source Description
UniProtINDUCTION: Up-regulated by the transcription factor AGAMOUS (PubMed:15254538). Regulated by ATXR3/SDG2 via chromatin methylation (PubMed:21037105). {ECO:0000269|PubMed:15254538, ECO:0000269|PubMed:21037105}.
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 AT1G23420 (A), AT4G18960 (A), AT4G37750 (A)
Regulation -- ATRM (Manually Curated Target Genes) ? help Back to Top
Source Target Gene (A: Activate/R: Repress)
ATRM AT1G23420(R), AT1G24260(A), AT2G17950(A), AT4G13260(R), AT4G18960(A), AT4G21750(R), AT4G32540(R), AT4G36920(A), AT4G39400(R), AT5G07280(A), AT5G25620(R), AT5G65700(A)
Regulation -- Hormone ? help Back to Top
Source Hormone
Interaction -- BIND ? help Back to Top
Source Intact With Description
BINDAT1G23420NZZ interacts with INO.
Interaction ? help Back to Top
Source Intact With
BioGRIDAT1G67260, AT4G27330, AT5G08070, AT5G08330, AT5G23280, AT5G41030, AT5G60970, AT1G23420, AT1G30210, AT1G35560, AT1G53230, AT1G58100, AT1G69690
Phenotype -- Disruption Phenotype ? help Back to Top
Source Description
UniProtDISRUPTION PHENOTYPE: Complete male and female sterility. {ECO:0000269|PubMed:10465788}.
Phenotype -- Mutation ? help Back to Top
Source ID
T-DNA ExpressAT4G27330
Annotation -- Nucleotide ? help Back to Top
Source Hit ID E-value Description
GenBankDQ4468730.0DQ446873.1 Arabidopsis thaliana clone pENTR221-At4g27330 sporocyteless (At4g27330) mRNA, complete cds.
Annotation -- Protein ? help Back to Top
Source Hit ID E-value Description
RefseqNP_194463.10.0putative transcription factor SPL
SwissprotO818360.0SPL_ARATH; Protein SPOROCYTELESS
TrEMBLD7MEG80.0D7MEG8_ARALL; Putative uncharacterized protein
STRINGAT4G27330.10.0(Arabidopsis thaliana)
Orthologous Group ? help Back to Top
LineageOrthologous Group IDTaxa NumberGene Number
Representative plantOGRP1001058
Publications ? help Back to Top
  1. Yang WC,Ye D,Xu J,Sundaresan V
    The SPOROCYTELESS gene of Arabidopsis is required for initiation of sporogenesis and encodes a novel nuclear protein.
    Genes Dev., 1999. 13(16): p. 2108-17
  2. Schiefthaler U, et al.
    Molecular analysis of NOZZLE, a gene involved in pattern formation and early sporogenesis during sex organ development in Arabidopsis thaliana.
    Proc. Natl. Acad. Sci. U.S.A., 1999. 96(20): p. 11664-9
  3. Villanueva JM, et al.
    INNER NO OUTER regulates abaxial- adaxial patterning in Arabidopsis ovules.
    Genes Dev., 1999. 13(23): p. 3160-9
  4. Balasubramanian S,Schneitz K
    NOZZLE regulates proximal-distal pattern formation, cell proliferation and early sporogenesis during ovule development in Arabidopsis thaliana.
    Development, 2000. 127(19): p. 4227-38
  5. Balasubramanian S,Schneitz K
    NOZZLE links proximal-distal and adaxial-abaxial pattern formation during ovule development in Arabidopsis thaliana.
    Development, 2002. 129(18): p. 4291-300
  6. Meister RJ, et al.
    Definition and interactions of a positive regulatory element of the Arabidopsis INNER NO OUTER promoter.
    Plant J., 2004. 37(3): p. 426-38
  7. Skinner DJ,Hill TA,Gasser CS
    Regulation of ovule development.
    Plant Cell, 2004. 16 Suppl: p. S32-45
  8. Ito T, et al.
    The homeotic protein AGAMOUS controls microsporogenesis by regulation of SPOROCYTELESS.
    Nature, 2004. 430(6997): p. 356-60
  9. Sieber P,Petrascheck M,Barberis A,Schneitz K
    Organ polarity in Arabidopsis. NOZZLE physically interacts with members of the YABBY family.
    Plant Physiol., 2004. 135(4): p. 2172-85
  10. Sieber P, et al.
    Pattern formation during early ovule development in Arabidopsis thaliana.
    Dev. Biol., 2004. 273(2): p. 321-34
  11. Dinneny JR,Yanofsky MF
    Floral development: an ABC gene chips in downstream.
    Curr. Biol., 2004. 14(19): p. R840-1
  12. Nakayama N, et al.
    Gene trap lines define domains of gene regulation in Arabidopsis petals and stamens.
    Plant Cell, 2005. 17(9): p. 2486-506
  13. Albrecht C,Russinova E,Hecht V,Baaijens E,de Vries S
    The Arabidopsis thaliana SOMATIC EMBRYOGENESIS RECEPTOR-LIKE KINASES1 and 2 control male sporogenesis.
    Plant Cell, 2005. 17(12): p. 3337-49
  14. Yu HJ,Hogan P,Sundaresan V
    Analysis of the female gametophyte transcriptome of Arabidopsis by comparative expression profiling.
    Plant Physiol., 2005. 139(4): p. 1853-69
  15. Hord CL,Chen C,Deyoung BJ,Clark SE,Ma H
    The BAM1/BAM2 receptor-like kinases are important regulators of Arabidopsis early anther development.
    Plant Cell, 2006. 18(7): p. 1667-80
  16. 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
  17. Matsuda T, et al.
    Cell-free synthesis of zinc-binding proteins.
    J. Struct. Funct. Genomics, 2006. 7(2): p. 93-100
  18. Underwood BA,Vanderhaeghen R,Whitford R,Town CD,Hilson P
    Simultaneous high-throughput recombinational cloning of open reading frames in closed and open configurations.
    Plant Biotechnol. J., 2006. 4(3): p. 317-24
  19. 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
  20. Brambilla V, et al.
    Genetic and molecular interactions between BELL1 and MADS box factors support ovule development in Arabidopsis.
    Plant Cell, 2007. 19(8): p. 2544-56
  21. Alves-Ferreira M, et al.
    Global expression profiling applied to the analysis of Arabidopsis stamen development.
    Plant Physiol., 2007. 145(3): p. 747-62
  22. Xing S,Zachgo S
    ROXY1 and ROXY2, two Arabidopsis glutaredoxin genes, are required for anther development.
    Plant J., 2008. 53(5): p. 790-801
  23. Li LC, et al.
    SPOROCYTELESS modulates YUCCA expression to regulate the development of lateral organs in Arabidopsis.
    New Phytol., 2008. 179(3): p. 751-64
  24. Liu X, et al.
    The SPOROCYTELESS/NOZZLE gene is involved in controlling stamen identity in Arabidopsis.
    Plant Physiol., 2009. 151(3): p. 1401-11
  25. Ye Q, et al.
    Brassinosteroids control male fertility by regulating the expression of key genes involved in Arabidopsis anther and pollen development.
    Proc. Natl. Acad. Sci. U.S.A., 2010. 107(13): p. 6100-5
  26. Irish VF
    The flowering of Arabidopsis flower development.
    Plant J., 2010. 61(6): p. 1014-28
  27. Yu N, et al.
    Temporal control of trichome distribution by microRNA156-targeted SPL genes in Arabidopsis thaliana.
    Plant Cell, 2010. 22(7): p. 2322-35
  28. Murmu J, et al.
    Arabidopsis basic leucine-zipper transcription factors TGA9 and TGA10 interact with floral glutaredoxins ROXY1 and ROXY2 and are redundantly required for anther development.
    Plant Physiol., 2010. 154(3): p. 1492-504
  29. Berr A, et al.
    Arabidopsis SET DOMAIN GROUP2 is required for H3K4 trimethylation and is crucial for both sporophyte and gametophyte development.
    Plant Cell, 2010. 22(10): p. 3232-48
  30. Xing S,Salinas M,H
    miR156-targeted and nontargeted SBP-box transcription factors act in concert to secure male fertility in Arabidopsis.
    Plant Cell, 2010. 22(12): p. 3935-50
  31. P
    CYP85A1 is required for the initiation of female gametogenesis in Arabidopsis thaliana.
    Plant Signal Behav, 2011. 6(3): p. 321-6
  32. Ma X,Feng B,Ma H
    AMS-dependent and independent regulation of anther transcriptome and comparison with those affected by other Arabidopsis anther genes.
    BMC Plant Biol., 2012. 12: p. 23
  33. S
    Transcriptional analysis of the Arabidopsis ovule by massively parallel signature sequencing.
    J. Exp. Bot., 2012. 63(10): p. 3829-42
  34. 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
  35. Bencivenga S,Simonini S,Benkov
    The transcription factors BEL1 and SPL are required for cytokinin and auxin signaling during ovule development in Arabidopsis.
    Plant Cell, 2012. 24(7): p. 2886-97
  36. Meinke DW
    A survey of dominant mutations in Arabidopsis thaliana.
    Trends Plant Sci., 2013. 18(2): p. 84-91
  37. Wei B, et al.
    The molecular mechanism of sporocyteless/nozzle in controlling Arabidopsis ovule development.
    Cell Res., 2015. 25(1): p. 121-34
  38. Chen GH,Sun JY,Liu M,Liu J,Yang WC
    SPOROCYTELESS is a novel embryophyte-specific transcription repressor that interacts with TPL and TCP proteins in Arabidopsis.
    J Genet Genomics, 2014. 41(12): p. 617-25
  39. 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