WOX9_ARATH
ID WOX9_ARATH Reviewed; 378 AA.
AC Q6X7J4; O23662; Q8GX13;
DT 15-MAR-2005, integrated into UniProtKB/Swiss-Prot.
DT 05-JUL-2004, sequence version 1.
DT 03-AUG-2022, entry version 130.
DE RecName: Full=WUSCHEL-related homeobox 9 {ECO:0000303|PubMed:14711878};
DE AltName: Full=Protein STIMPY {ECO:0000303|PubMed:15753038};
GN Name=WOX9 {ECO:0000303|PubMed:14711878};
GN Synonyms=STIP {ECO:0000303|PubMed:15753038};
GN OrderedLocusNames=At2g33880 {ECO:0000312|Araport:AT2G33880};
GN ORFNames=T1B8.31 {ECO:0000312|EMBL:AAC69146.1};
OS Arabidopsis thaliana (Mouse-ear cress).
OC Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
OC Spermatophyta; Magnoliopsida; eudicotyledons; Gunneridae; Pentapetalae;
OC rosids; malvids; Brassicales; Brassicaceae; Camelineae; Arabidopsis.
OX NCBI_TaxID=3702;
RN [1]
RP NUCLEOTIDE SEQUENCE [MRNA], DEVELOPMENTAL STAGE, AND TISSUE SPECIFICITY.
RC STRAIN=cv. Landsberg erecta;
RX PubMed=14711878; DOI=10.1242/dev.00963;
RA Haecker A., Gross-Hardt R., Geiges B., Sarkar A., Breuninger H.,
RA Herrmann M., Laux T.;
RT "Expression dynamics of WOX genes mark cell fate decisions during early
RT embryonic patterning in Arabidopsis thaliana.";
RL Development 131:657-668(2004).
RN [2]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC STRAIN=cv. Columbia;
RX PubMed=10617197; DOI=10.1038/45471;
RA Lin X., Kaul S., Rounsley S.D., Shea T.P., Benito M.-I., Town C.D.,
RA Fujii C.Y., Mason T.M., Bowman C.L., Barnstead M.E., Feldblyum T.V.,
RA Buell C.R., Ketchum K.A., Lee J.J., Ronning C.M., Koo H.L., Moffat K.S.,
RA Cronin L.A., Shen M., Pai G., Van Aken S., Umayam L., Tallon L.J.,
RA Gill J.E., Adams M.D., Carrera A.J., Creasy T.H., Goodman H.M.,
RA Somerville C.R., Copenhaver G.P., Preuss D., Nierman W.C., White O.,
RA Eisen J.A., Salzberg S.L., Fraser C.M., Venter J.C.;
RT "Sequence and analysis of chromosome 2 of the plant Arabidopsis thaliana.";
RL Nature 402:761-768(1999).
RN [3]
RP GENOME REANNOTATION.
RC STRAIN=cv. Columbia;
RX PubMed=27862469; DOI=10.1111/tpj.13415;
RA Cheng C.Y., Krishnakumar V., Chan A.P., Thibaud-Nissen F., Schobel S.,
RA Town C.D.;
RT "Araport11: a complete reannotation of the Arabidopsis thaliana reference
RT genome.";
RL Plant J. 89:789-804(2017).
RN [4]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
RC STRAIN=cv. Columbia;
RX PubMed=11910074; DOI=10.1126/science.1071006;
RA Seki M., Narusaka M., Kamiya A., Ishida J., Satou M., Sakurai T.,
RA Nakajima M., Enju A., Akiyama K., Oono Y., Muramatsu M., Hayashizaki Y.,
RA Kawai J., Carninci P., Itoh M., Ishii Y., Arakawa T., Shibata K.,
RA Shinagawa A., Shinozaki K.;
RT "Functional annotation of a full-length Arabidopsis cDNA collection.";
RL Science 296:141-145(2002).
RN [5]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] OF 74-378.
RC STRAIN=cv. Columbia;
RX PubMed=14593172; DOI=10.1126/science.1088305;
RA Yamada K., Lim J., Dale J.M., Chen H., Shinn P., Palm C.J., Southwick A.M.,
RA Wu H.C., Kim C.J., Nguyen M., Pham P.K., Cheuk R.F., Karlin-Newmann G.,
RA Liu S.X., Lam B., Sakano H., Wu T., Yu G., Miranda M., Quach H.L.,
RA Tripp M., Chang C.H., Lee J.M., Toriumi M.J., Chan M.M., Tang C.C.,
RA Onodera C.S., Deng J.M., Akiyama K., Ansari Y., Arakawa T., Banh J.,
RA Banno F., Bowser L., Brooks S.Y., Carninci P., Chao Q., Choy N., Enju A.,
RA Goldsmith A.D., Gurjal M., Hansen N.F., Hayashizaki Y., Johnson-Hopson C.,
RA Hsuan V.W., Iida K., Karnes M., Khan S., Koesema E., Ishida J., Jiang P.X.,
RA Jones T., Kawai J., Kamiya A., Meyers C., Nakajima M., Narusaka M.,
RA Seki M., Sakurai T., Satou M., Tamse R., Vaysberg M., Wallender E.K.,
RA Wong C., Yamamura Y., Yuan S., Shinozaki K., Davis R.W., Theologis A.,
RA Ecker J.R.;
RT "Empirical analysis of transcriptional activity in the Arabidopsis
RT genome.";
RL Science 302:842-846(2003).
RN [6]
RP FUNCTION, TISSUE SPECIFICITY, AND DISRUPTION PHENOTYPE.
RX PubMed=15753038; DOI=10.1016/j.cub.2004.12.079;
RA Wu X., Dabi T., Weigel D.;
RT "Requirement of homeobox gene STIMPY/WOX9 for Arabidopsis meristem growth
RT and maintenance.";
RL Curr. Biol. 15:436-440(2005).
RN [7]
RP FUNCTION, DEVELOPMENTAL STAGE, SUBCELLULAR LOCATION, AND DISRUPTION
RP PHENOTYPE.
RX PubMed=17706632; DOI=10.1016/j.ydbio.2007.07.019;
RA Wu X., Chory J., Weigel D.;
RT "Combinations of WOX activities regulate tissue proliferation during
RT Arabidopsis embryonic development.";
RL Dev. Biol. 309:306-316(2007).
RN [8]
RP FUNCTION, AND TISSUE SPECIFICITY.
RX PubMed=20110319; DOI=10.1242/dev.041426;
RA Skylar A., Hong F., Chory J., Weigel D., Wu X.;
RT "STIMPY mediates cytokinin signaling during shoot meristem establishment in
RT Arabidopsis seedlings.";
RL Development 137:541-549(2010).
RN [9]
RP FUNCTION.
RX PubMed=21057190; DOI=10.4161/psb.5.11.13356;
RA Skylar A., Wu X.;
RT "STIMPY mutants have increased cytokinin sensitivity during dark
RT germination.";
RL Plant Signal. Behav. 5:1437-1439(2010).
RN [10]
RP FUNCTION.
RX PubMed=21185286; DOI=10.1016/j.ydbio.2010.12.019;
RA Skylar A., Sung F., Hong F., Chory J., Wu X.;
RT "Metabolic sugar signal promotes Arabidopsis meristematic proliferation via
RT G2.";
RL Dev. Biol. 351:82-89(2011).
RN [11]
RP INDUCTION BY WRKY2.
RX PubMed=21316593; DOI=10.1016/j.devcel.2011.01.009;
RA Ueda M., Zhang Z., Laux T.;
RT "Transcriptional activation of Arabidopsis axis patterning genes WOX8/9
RT links zygote polarity to embryo development.";
RL Dev. Cell 20:264-270(2011).
CC -!- FUNCTION: Homeodomain transcription factor required for meristem growth
CC and early development (PubMed:15753038). Promotes cell proliferation
CC and prevents premature differentiation in meristematic tissues during
CC postembryonic development (PubMed:15753038). Essential for maintaining
CC tissue growth during embryogenesis (PubMed:17706632). May act by
CC repressing TSS to promote meristematic proliferation (PubMed:21185286).
CC Involved in the transcriptional activation of a subset of cytokinin
CC response factors (PubMed:20110319). May act as a negative regulator of
CC cytokinin signaling in the dark (PubMed:21057190).
CC {ECO:0000269|PubMed:15753038, ECO:0000269|PubMed:17706632,
CC ECO:0000269|PubMed:20110319, ECO:0000269|PubMed:21057190,
CC ECO:0000303|PubMed:21185286}.
CC -!- SUBCELLULAR LOCATION: Nucleus {ECO:0000269|PubMed:17706632}. Cytoplasm
CC {ECO:0000269|PubMed:17706632}.
CC -!- TISSUE SPECIFICITY: Expressed in the basal cell and later at the
CC boundary between suspensor and proembryo (PubMed:14711878). Expressed
CC at low levels in proliferating tissues post embryonically
CC (PubMed:15753038). Detected in vegetative shoot apical meristem, leaf
CC primordia, floral meristems, emerging floral organs, epidermal layer of
CC the placenta and in the upper portion of the root meristematic zone
CC (PubMed:15753038, PubMed:20110319). {ECO:0000269|PubMed:14711878,
CC ECO:0000269|PubMed:15753038, ECO:0000269|PubMed:20110319}.
CC -!- DEVELOPMENTAL STAGE: First detected in the basal daughter cell of the
CC zygote (PubMed:14711878). Unlike WOX2 and WOX8, it is not expressed in
CC the egg cell or the zygote (PubMed:14711878). During two subsequent
CC rounds of transverse cell divisions, it is restricted to the hypophysis
CC (PubMed:14711878). At the 8-cell stage, it expands into the central
CC domain of the embryo, in addition to weakening in the hypophysis
CC (PubMed:14711878). After protoderm formation, expression in the central
CC embryo domain is restricted to the protodermal cells and also
CC disappears from the hypophyseal cell (PubMed:14711878). In subsequent
CC stages, a ring of expression remains at the presumptive boundary
CC between root and hypocotyl, at about the same position as WOX2
CC expression in heart stage embryos (PubMed:14711878). In addition to its
CC embryonic expression, it is expressed postembryonically in the
CC epidermal cells of the placenta during gynoecium development, but not
CC in the developing ovules (PubMed:14711878). The placental expression
CC disappears soon after fertilization (PubMed:14711878). Expression
CC dynamics require MP and BDL, but not GN activity (PubMed:14711878).
CC Detected as early as the first zygotic division in both the apical and
CC the basal daughter cells (PubMed:17706632). By late globular to early
CC transition stage, the protein is evenly distributed through out the
CC embryo and the suspensor (PubMed:17706632). The basal half of the
CC embryo, especially cells in the protoderm layer, starts to show
CC slightly higher levels of expression by early heart stage
CC (PubMed:17706632). {ECO:0000269|PubMed:14711878,
CC ECO:0000269|PubMed:17706632}.
CC -!- INDUCTION: Up-regulated in the zygote after fertilization by the
CC transcription factor WRKY2. {ECO:0000269|PubMed:21316593}.
CC -!- DISRUPTION PHENOTYPE: Embryonic lethality when homozygous
CC (PubMed:17706632). Reduction in embryonic growth, failure in axillary
CC shoot apical meristem and leaf-primordia initiation and growth, and
CC failure in primary root growth and lateral root initiation when
CC heterozygous (PubMed:15753038, PubMed:17706632). Wox8 and wox9 double
CC mutants are embryo lethal, with embryos disrupted as early as the first
CC cell division in the embryo proper (PubMed:17706632).
CC {ECO:0000269|PubMed:15753038, ECO:0000269|PubMed:17706632}.
CC -!- MISCELLANEOUS: The seedling growth arrest phenotype of the stip mutants
CC likely resulted from a G2 cell cycle arrest of the meristematic tissue
CC (PubMed:17706632). Exogenous sucrose in the growth medium reactivate
CC the stip meristems by repressing TSS and activating the expression of
CC cell cycle regulators, and thus promoting G2 to M transition in
CC meristematic tissues (PubMed:21185286). {ECO:0000303|PubMed:17706632,
CC ECO:0000303|PubMed:21185286}.
CC -!- SIMILARITY: Belongs to the WUS homeobox family. {ECO:0000305}.
CC -!- SEQUENCE CAUTION:
CC Sequence=AAC69146.1; Type=Erroneous gene model prediction; Evidence={ECO:0000305};
CC Sequence=BAC43105.1; Type=Erroneous termination; Note=Truncated C-terminus.; Evidence={ECO:0000305};
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DR EMBL; AY251401; AAP37139.1; -; mRNA.
DR EMBL; U78721; AAC69146.1; ALT_SEQ; Genomic_DNA.
DR EMBL; CP002685; AEC08902.1; -; Genomic_DNA.
DR EMBL; AK118501; BAC43105.1; ALT_SEQ; mRNA.
DR EMBL; BT005296; AAO63360.1; -; mRNA.
DR PIR; G84750; G84750.
DR RefSeq; NP_180944.2; NM_128948.4.
DR AlphaFoldDB; Q6X7J4; -.
DR SMR; Q6X7J4; -.
DR BioGRID; 3305; 11.
DR IntAct; Q6X7J4; 10.
DR STRING; 3702.AT2G33880.1; -.
DR PaxDb; Q6X7J4; -.
DR PRIDE; Q6X7J4; -.
DR EnsemblPlants; AT2G33880.1; AT2G33880.1; AT2G33880.
DR GeneID; 817958; -.
DR Gramene; AT2G33880.1; AT2G33880.1; AT2G33880.
DR KEGG; ath:AT2G33880; -.
DR Araport; AT2G33880; -.
DR TAIR; locus:2057614; AT2G33880.
DR eggNOG; ENOG502QVSY; Eukaryota.
DR HOGENOM; CLU_030463_1_0_1; -.
DR InParanoid; Q6X7J4; -.
DR PhylomeDB; Q6X7J4; -.
DR PRO; PR:Q6X7J4; -.
DR Proteomes; UP000006548; Chromosome 2.
DR ExpressionAtlas; Q6X7J4; baseline and differential.
DR Genevisible; Q6X7J4; AT.
DR GO; GO:0005737; C:cytoplasm; IDA:TAIR.
DR GO; GO:0005634; C:nucleus; IDA:TAIR.
DR GO; GO:0003700; F:DNA-binding transcription factor activity; ISS:TAIR.
DR GO; GO:0000976; F:transcription cis-regulatory region binding; IPI:TAIR.
DR GO; GO:0009793; P:embryo development ending in seed dormancy; IMP:TAIR.
DR GO; GO:0008284; P:positive regulation of cell population proliferation; IMP:TAIR.
DR GO; GO:0010075; P:regulation of meristem growth; IMP:TAIR.
DR CDD; cd00086; homeodomain; 1.
DR InterPro; IPR009057; Homeobox-like_sf.
DR InterPro; IPR001356; Homeobox_dom.
DR InterPro; IPR044557; WOX8/9-like.
DR PANTHER; PTHR47288; PTHR47288; 1.
DR Pfam; PF00046; Homeodomain; 1.
DR SMART; SM00389; HOX; 1.
DR SUPFAM; SSF46689; SSF46689; 1.
DR PROSITE; PS50071; HOMEOBOX_2; 1.
PE 2: Evidence at transcript level;
KW Cytoplasm; Developmental protein; DNA-binding; Homeobox; Nucleus;
KW Reference proteome; Transcription; Transcription regulation.
FT CHAIN 1..378
FT /note="WUSCHEL-related homeobox 9"
FT /id="PRO_0000049376"
FT DNA_BIND 51..115
FT /note="Homeobox; WUS-type"
FT /evidence="ECO:0000255|PROSITE-ProRule:PRU00108"
FT REGION 1..60
FT /note="Disordered"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT REGION 123..173
FT /note="Disordered"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT COMPBIAS 25..44
FT /note="Polar residues"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT COMPBIAS 123..171
FT /note="Polar residues"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
SQ SEQUENCE 378 AA; 42089 MW; 5F63ECE82341D28A CRC64;
MASSNRHWPS MFKSKPHPHQ WQHDINSPLL PSASHRSSPF SSGCEVERSP EPKPRWNPKP
EQIRILEAIF NSGMVNPPRE EIRRIRAQLQ EYGQVGDANV FYWFQNRKSR SKHKLRLLHN
HSKHSLPQTQ PQPQPQPSAS SSSSSSSSSS KSTKPRKSKN KNNTNLSLGG SQMMGMFPPE
PAFLFPVSTV GGFEGITVSS QLGFLSGDMI EQQKPAPTCT GLLLSEIMNG SVSYGTHHQQ
HLSEKEVEEM RMKMLQQPQT QICYATTNHQ IASYNNNNNN NNIMLHIPPT TSTATTITTS
HSLATVPSTS DQLQVQADAR IRVFINEMEL EVSSGPFNVR DAFGEEVVLI NSAGQPIVTD
EYGVALHPLQ HGASYYLI
