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NAC98_ARATH
ID   NAC98_ARATH             Reviewed;         375 AA.
AC   O04017;
DT   04-DEC-2007, integrated into UniProtKB/Swiss-Prot.
DT   01-JUL-1997, sequence version 1.
DT   03-AUG-2022, entry version 123.
DE   RecName: Full=Protein CUP-SHAPED COTYLEDON 2;
DE   AltName: Full=NAC domain-containing protein 98;
DE            Short=ANAC098;
DE   AltName: Full=NAC domain-containing protein CUC2;
GN   Name=NAC098; Synonyms=CUC2; OrderedLocusNames=At5g53950;
GN   ORFNames=K19P17.12;
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], FUNCTION, AND TISSUE SPECIFICITY.
RC   STRAIN=cv. Columbia;
RX   PubMed=9212461; DOI=10.1105/tpc.9.6.841;
RA   Aida M., Ishida T., Fukaki H., Fujisawa H., Tasaka M.;
RT   "Genes involved in organ separation in Arabidopsis: an analysis of the cup-
RT   shaped cotyledon mutant.";
RL   Plant Cell 9:841-857(1997).
RN   [2]
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC   STRAIN=cv. Columbia;
RX   PubMed=9501997; DOI=10.1093/dnares/4.6.401;
RA   Nakamura Y., Sato S., Kaneko T., Kotani H., Asamizu E., Miyajima N.,
RA   Tabata S.;
RT   "Structural analysis of Arabidopsis thaliana chromosome 5. III. Sequence
RT   features of the regions of 1,191,918 bp covered by seventeen physically
RT   assigned P1 clones.";
RL   DNA Res. 4:401-414(1997).
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;
RA   Bautista V.R., Kim C.J., Chen H., Wu S.Y., De Los Reyes C., Ecker J.R.;
RT   "Arabidopsis ORF clones.";
RL   Submitted (JAN-2007) to the EMBL/GenBank/DDBJ databases.
RN   [5]
RP   FUNCTION, AND DEVELOPMENTAL STAGE.
RX   PubMed=10079219; DOI=10.1242/dev.126.8.1563;
RA   Aida M., Ishida T., Tasaka M.;
RT   "Shoot apical meristem and cotyledon formation during Arabidopsis
RT   embryogenesis: interaction among the CUP-SHAPED COTYLEDON and SHOOT
RT   MERISTEMLESS genes.";
RL   Development 126:1563-1570(1999).
RN   [6]
RP   FUNCTION, TISSUE SPECIFICITY, AND DEVELOPMENTAL STAGE.
RX   PubMed=10750709; DOI=10.1093/pcp/41.1.60;
RA   Ishida T., Aida M., Takada S., Tasaka M.;
RT   "Involvement of CUP-SHAPED COTYLEDON genes in gynoecium and ovule
RT   development in Arabidopsis thaliana.";
RL   Plant Cell Physiol. 41:60-67(2000).
RN   [7]
RP   FUNCTION.
RX   PubMed=12163400; DOI=10.1242/dev.129.17.3965;
RA   Aida M., Vernoux T., Furutani M., Traas J., Tasaka M.;
RT   "Roles of PIN-FORMED1 and MONOPTEROS in pattern formation of the apical
RT   region of the Arabidopsis embryo.";
RL   Development 129:3965-3974(2002).
RN   [8]
RP   FUNCTION.
RX   PubMed=12492830; DOI=10.1046/j.1365-313x.2002.01479.x;
RA   Cary A.J., Che P., Howell S.H.;
RT   "Developmental events and shoot apical meristem gene expression patterns
RT   during shoot development in Arabidopsis thaliana.";
RL   Plant J. 32:867-877(2002).
RN   [9]
RP   GENE FAMILY, AND NOMENCLATURE.
RX   PubMed=15029955; DOI=10.1093/dnares/10.6.239;
RA   Ooka H., Satoh K., Doi K., Nagata T., Otomo Y., Murakami K., Matsubara K.,
RA   Osato N., Kawai J., Carninci P., Hayashizaki Y., Suzuki K., Kojima K.,
RA   Takahara Y., Yamamoto K., Kikuchi S.;
RT   "Comprehensive analysis of NAC family genes in Oryza sativa and Arabidopsis
RT   thaliana.";
RL   DNA Res. 10:239-247(2003).
RN   [10]
RP   FUNCTION.
RX   PubMed=12610213; DOI=10.1093/pcp/pcg038;
RA   Daimon Y., Takabe K., Tasaka M.;
RT   "The CUP-SHAPED COTYLEDON genes promote adventitious shoot formation on
RT   calli.";
RL   Plant Cell Physiol. 44:113-121(2003).
RN   [11]
RP   FUNCTION, AND INDUCTION BY MIR164.
RX   PubMed=15202996; DOI=10.1016/j.cub.2004.06.022;
RA   Mallory A.C., Dugas D.V., Bartel D.P., Bartel B.;
RT   "MicroRNA regulation of NAC-domain targets is required for proper formation
RT   and separation of adjacent embryonic, vegetative, and floral organs.";
RL   Curr. Biol. 14:1035-1046(2004).
RN   [12]
RP   FUNCTION, AND INDUCTION BY MIR164.
RX   PubMed=15294871; DOI=10.1242/dev.01320;
RA   Laufs P., Peaucelle A., Morin H., Traas J.;
RT   "MicroRNA regulation of the CUC genes is required for boundary size control
RT   in Arabidopsis meristems.";
RL   Development 131:4311-4322(2004).
RN   [13]
RP   FUNCTION, MUTAGENESIS OF TRP-373 AND TYR-375, DOMAIN, TRANSACTIVATING
RP   REGION, AND SUBCELLULAR LOCATION.
RX   PubMed=15500463; DOI=10.1111/j.1365-313x.2004.02238.x;
RA   Taoka K., Yanagimoto Y., Daimon Y., Hibara K., Aida M., Tasaka M.;
RT   "The NAC domain mediates functional specificity of CUP-SHAPED COTYLEDON
RT   proteins.";
RL   Plant J. 40:462-473(2004).
RN   [14]
RP   TISSUE SPECIFICITY.
RX   PubMed=15053771; DOI=10.1111/j.1365-313x.2004.02026.x;
RA   Breuil-Broyer S., Morel P., de Almeida-Engler J., Coustham V., Negrutiu I.,
RA   Trehin C.;
RT   "High-resolution boundary analysis during Arabidopsis thaliana flower
RT   development.";
RL   Plant J. 38:182-192(2004).
RN   [15]
RP   FUNCTION, AND INDUCTION BY MIR164.
RX   PubMed=15723790; DOI=10.1016/j.cub.2005.02.017;
RA   Baker C.C., Sieber P., Wellmer F., Meyerowitz E.M.;
RT   "The early extra petals1 mutant uncovers a role for microRNA miR164c in
RT   regulating petal number in Arabidopsis.";
RL   Curr. Biol. 15:303-315(2005).
RN   [16]
RP   INDUCTION BY BRM AND SYD.
RX   PubMed=16854978; DOI=10.1242/dev.02508;
RA   Kwon C.S., Hibara K., Pfluger J., Bezhani S., Metha H., Aida M., Tasaka M.,
RA   Wagner D.;
RT   "A role for chromatin remodeling in regulation of CUC gene expression in
RT   the Arabidopsis cotyledon boundary.";
RL   Development 133:3223-3230(2006).
RN   [17]
RP   FUNCTION.
RX   PubMed=16798887; DOI=10.1105/tpc.106.041988;
RA   Belles-Boix E., Hamant O., Witiak S.M., Morin H., Traas J., Pautot V.;
RT   "KNAT6: an Arabidopsis homeobox gene involved in meristem activity and
RT   organ separation.";
RL   Plant Cell 18:1900-1907(2006).
RN   [18]
RP   FUNCTION, AND DEVELOPMENTAL STAGE.
RX   PubMed=17098808; DOI=10.1105/tpc.106.045617;
RA   Nikovics K., Blein T., Peaucelle A., Ishida T., Morin H., Aida M.,
RA   Laufs P.;
RT   "The balance between the MIR164A and CUC2 genes controls leaf margin
RT   serration in Arabidopsis.";
RL   Plant Cell 18:2929-2945(2006).
RN   [19]
RP   FUNCTION.
RX   PubMed=17122068; DOI=10.1105/tpc.106.045716;
RA   Hibara K., Karim M.R., Takada S., Taoka K., Furutani M., Aida M.,
RA   Tasaka M.;
RT   "Arabidopsis CUP-SHAPED COTYLEDON3 regulates postembryonic shoot meristem
RT   and organ boundary formation.";
RL   Plant Cell 18:2946-2957(2006).
RN   [20]
RP   FUNCTION, DEVELOPMENTAL STAGE, AND INDUCTION BY MIR164.
RX   PubMed=17251269; DOI=10.1242/dev.02774;
RA   Peaucelle A., Morin H., Traas J., Laufs P.;
RT   "Plants expressing a miR164-resistant CUC2 gene reveal the importance of
RT   post-meristematic maintenance of phyllotaxy in Arabidopsis.";
RL   Development 134:1045-1050(2007).
RN   [21]
RP   FUNCTION, AND INDUCTION BY MIR164.
RX   PubMed=17287247; DOI=10.1242/dev.02817;
RA   Sieber P., Wellmer F., Gheyselinck J., Riechmann J.L., Meyerowitz E.M.;
RT   "Redundancy and specialization among plant microRNAs: role of the MIR164
RT   family in developmental robustness.";
RL   Development 134:1051-1060(2007).
CC   -!- FUNCTION: Transcription activator of STM and KNAT6. Involved in
CC       molecular mechanisms regulating shoot apical meristem (SAM) formation
CC       during embryogenesis and organ separation. Required for the fusion of
CC       septa of gynoecia along the length of the ovaries. Activates the shoot
CC       formation in callus in a STM-dependent manner. Controls leaf margin
CC       development and required for leaf serration. Involved in axillary
CC       meristem initiation and separation of the meristem from the main stem.
CC       Regulates the phyllotaxy throughout the plant development. Seems to act
CC       as an inhibitor of cell division. {ECO:0000269|PubMed:10079219,
CC       ECO:0000269|PubMed:10750709, ECO:0000269|PubMed:12163400,
CC       ECO:0000269|PubMed:12492830, ECO:0000269|PubMed:12610213,
CC       ECO:0000269|PubMed:15202996, ECO:0000269|PubMed:15294871,
CC       ECO:0000269|PubMed:15500463, ECO:0000269|PubMed:15723790,
CC       ECO:0000269|PubMed:16798887, ECO:0000269|PubMed:17098808,
CC       ECO:0000269|PubMed:17122068, ECO:0000269|PubMed:17251269,
CC       ECO:0000269|PubMed:17287247, ECO:0000269|PubMed:9212461}.
CC   -!- INTERACTION:
CC       O04017; Q9SFV2: FHA2; NbExp=3; IntAct=EBI-1998502, EBI-15191747;
CC       O04017; O82277: TCP10; NbExp=3; IntAct=EBI-1998502, EBI-3133327;
CC   -!- SUBCELLULAR LOCATION: Nucleus {ECO:0000255|PROSITE-ProRule:PRU00353,
CC       ECO:0000269|PubMed:15500463}.
CC   -!- TISSUE SPECIFICITY: Mostly expressed in buds and flowers, and, to a
CC       lower extent, in the aerial parts of seedling, inflorescence and old
CC       silique. In a general manner, present at the boundaries between
CC       mersitems and araising primordia. {ECO:0000269|PubMed:10750709,
CC       ECO:0000269|PubMed:15053771, ECO:0000269|PubMed:9212461}.
CC   -!- DEVELOPMENTAL STAGE: First expressed in early to mid-globular-stage
CC       embryos. In late globular stage, detected as a stripe running medially
CC       across the top of the embryo. In heart stage embryo, expression is
CC       restricted to a stripe between the cotyledon primordia, but confined to
CC       hypodermal cells. In the bending-cotyledon stage, localized in a region
CC       surrounding the SAM, that correspond to the boundary region of
CC       cotyledon margins (BCM) and the boundaries between SAM and cotyledons,
CC       including protoderm cells. Observed in the margins of leaf primordia,
CC       and later restricted to the leaf sinus region, with a diminution in
CC       outgrowing teeth. Restricted to the proximal part of mature leaves.
CC       Expressed at the boundaries between the inflorescence meristem (IM) and
CC       flower primordia. Once the flower starts to grow out and the internode
CC       begin to elongates, restricted to the axils of the floral pedicels
CC       through several nodes. Detected within floral primordia, between sepal
CC       primordia and the floral meristem. Also present at the boundaries of
CC       individual sepal primordia, as well as in the region surrounding each
CC       petal and stamen primordium. Later detected transiently at the
CC       boundaries between locules of each theca in anthers. Expression at the
CC       inner part of presumtive septal regions that raises to include
CC       presumptive placenta, at the tips of septal primordia, as septum grow.
CC       Localized in the fused region of the septum. Found at the boundaries of
CC       ovule primordia, and later at the boundary between the nucellus and the
CC       chalaza. {ECO:0000269|PubMed:10079219, ECO:0000269|PubMed:10750709,
CC       ECO:0000269|PubMed:17098808, ECO:0000269|PubMed:17251269}.
CC   -!- INDUCTION: By BRM and SYD, at the chromatin level, and conferring a
CC       very specific spatial expression pattern. Precise spatial regulation by
CC       post-transcriptional repression directed by the microRNA miR164.
CC       {ECO:0000269|PubMed:15202996, ECO:0000269|PubMed:15294871,
CC       ECO:0000269|PubMed:15723790, ECO:0000269|PubMed:16854978,
CC       ECO:0000269|PubMed:17251269, ECO:0000269|PubMed:17287247}.
CC   -!- DOMAIN: The NAC domain includes a DNA-binding domain and a dimerization
CC       domain, and confers the specificity of the transactivated target genes.
CC       {ECO:0000269|PubMed:15500463}.
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DR   EMBL; AB002560; BAA19529.1; -; Genomic_DNA.
DR   EMBL; AB007644; BAB10725.1; -; Genomic_DNA.
DR   EMBL; CP002688; AED96432.1; -; Genomic_DNA.
DR   EMBL; BT030026; ABN04764.1; -; mRNA.
DR   RefSeq; NP_200206.1; NM_124774.3.
DR   AlphaFoldDB; O04017; -.
DR   SMR; O04017; -.
DR   BioGRID; 20722; 31.
DR   IntAct; O04017; 25.
DR   STRING; 3702.AT5G53950.1; -.
DR   PaxDb; O04017; -.
DR   PRIDE; O04017; -.
DR   ProteomicsDB; 251013; -.
DR   EnsemblPlants; AT5G53950.1; AT5G53950.1; AT5G53950.
DR   GeneID; 835478; -.
DR   Gramene; AT5G53950.1; AT5G53950.1; AT5G53950.
DR   KEGG; ath:AT5G53950; -.
DR   Araport; AT5G53950; -.
DR   TAIR; locus:2154684; AT5G53950.
DR   eggNOG; ENOG502QSC9; Eukaryota.
DR   HOGENOM; CLU_035664_5_2_1; -.
DR   InParanoid; O04017; -.
DR   OMA; DSGYYNQ; -.
DR   OrthoDB; 1264987at2759; -.
DR   PhylomeDB; O04017; -.
DR   PRO; PR:O04017; -.
DR   Proteomes; UP000006548; Chromosome 5.
DR   ExpressionAtlas; O04017; baseline and differential.
DR   Genevisible; O04017; AT.
DR   GO; GO:0005634; C:nucleus; IEA:UniProtKB-SubCell.
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:0090691; P:formation of plant organ boundary; IGI:TAIR.
DR   GO; GO:0048366; P:leaf development; IMP:TAIR.
DR   GO; GO:0010072; P:primary shoot apical meristem specification; IGI:TAIR.
DR   GO; GO:0090709; P:regulation of timing of plant organ formation; IMP:TAIR.
DR   GO; GO:0010223; P:secondary shoot formation; IMP:TAIR.
DR   Gene3D; 2.170.150.80; -; 1.
DR   InterPro; IPR003441; NAC-dom.
DR   InterPro; IPR036093; NAC_dom_sf.
DR   Pfam; PF02365; NAM; 1.
DR   SUPFAM; SSF101941; SSF101941; 1.
DR   PROSITE; PS51005; NAC; 1.
PE   1: Evidence at protein level;
KW   Developmental protein; DNA-binding; Nucleus; Reference proteome;
KW   Transcription; Transcription regulation.
FT   CHAIN           1..375
FT                   /note="Protein CUP-SHAPED COTYLEDON 2"
FT                   /id="PRO_0000312287"
FT   DOMAIN          17..169
FT                   /note="NAC"
FT                   /evidence="ECO:0000255|PROSITE-ProRule:PRU00353"
FT   DNA_BIND        116..175
FT                   /evidence="ECO:0000255|PROSITE-ProRule:PRU00353"
FT   REGION          175..228
FT                   /note="Disordered"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   REGION          210..247
FT                   /note="Involved in transactivation activity"
FT   REGION          373..375
FT                   /note="Involved in transactivation activity"
FT   COMPBIAS        186..228
FT                   /note="Polar residues"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   MUTAGEN         373
FT                   /note="W->A: Loss of transactivation activity. when
FT                   associated with A-373."
FT                   /evidence="ECO:0000269|PubMed:15500463"
FT   MUTAGEN         375
FT                   /note="Y->A: Loss of transactivation activity. when
FT                   associated with A-371."
FT                   /evidence="ECO:0000269|PubMed:15500463"
SQ   SEQUENCE   375 AA;  41434 MW;  DA2E8AD98F83D11B CRC64;
     MDIPYYHYDH GGDSQYLPPG FRFHPTDEEL ITHYLLRKVL DGCFSSRAIA EVDLNKCEPW
     QLPGRAKMGE KEWYFFSLRD RKYPTGLRTN RATEAGYWKA TGKDREIFSS KTCALVGMKK
     TLVFYKGRAP KGEKSNWVMH EYRLEGKFSY HFISRSSKDE WVISRVFQKT TLASTGAVSE
     GGGGGGATVS VSSGTGPSKK TKVPSTISRN YQEQPSSPSS VSLPPLLDPT TTLGYTDSSC
     SYDSRSTNTT VTASAITEHV SCFSTVPTTT TALGLDVNSF SRLPPPLGFD FDPFPRFVSR
     NVSTQSNFRS FQENFNQFPY FGSSSASTMT SAVNLPSFQG GGGVSGMNYW LPATAEENES
     KVGVLHAGLD CIWNY
 
 
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