ID BLH9_ARATH Reviewed; 575 AA.
AC Q9LZM8; Q8H147; Q8LGJ2;
DT 15-JAN-2008, integrated into UniProtKB/Swiss-Prot.
DT 01-OCT-2000, sequence version 1.
DT 27-MAR-2024, entry version 158.
DE RecName: Full=BEL1-like homeodomain protein 9;
DE Short=BEL1-like protein 9;
DE AltName: Full=Protein BELLRINGER;
DE AltName: Full=Protein LARSON;
DE AltName: Full=Protein PENNYWISE;
DE AltName: Full=Protein REPLUMLESS;
DE AltName: Full=Protein VAAMANA;
GN Name=BLH9; Synonyms=BLR, LSN, PNY, RPL, VAN; OrderedLocusNames=At5g02030;
GN ORFNames=T7H20_80;
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, TISSUE SPECIFICITY, DEVELOPMENTAL
RP STAGE, AND INTERACTION WITH STM AND KNAT1/BP.
RX PubMed=12874117; DOI=10.1242/dev.00620;
RA Byrne M.E., Groover A.T., Fontana J.R., Martienssen R.A.;
RT "Phyllotactic pattern and stem cell fate are determined by the Arabidopsis
RT homeobox gene BELLRINGER.";
RL Development 130:3941-3950(2003).
RN [2]
RP NUCLEOTIDE SEQUENCE [MRNA], FUNCTION, TISSUE SPECIFICITY, AND INTERACTION
RP WITH STM; KNAT1/BP AND KNAT6.
RC STRAIN=cv. Landsberg erecta; TISSUE=Flower;
RX PubMed=15019989; DOI=10.1016/j.gene.2003.12.033;
RA Bhatt A.M., Etchells J.P., Canales C., Lagodienko A., Dickinson H.;
RT "VAAMANA -- a BEL1-like homeodomain protein, interacts with KNOX proteins
RT BP and STM and regulates inflorescence stem growth in Arabidopsis.";
RL Gene 328:103-111(2004).
RN [3]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC STRAIN=cv. Columbia;
RX PubMed=11130714; DOI=10.1038/35048507;
RA Tabata S., Kaneko T., Nakamura Y., Kotani H., Kato T., Asamizu E.,
RA Miyajima N., Sasamoto S., Kimura T., Hosouchi T., Kawashima K., Kohara M.,
RA Matsumoto M., Matsuno A., Muraki A., Nakayama S., Nakazaki N., Naruo K.,
RA Okumura S., Shinpo S., Takeuchi C., Wada T., Watanabe A., Yamada M.,
RA Yasuda M., Sato S., de la Bastide M., Huang E., Spiegel L., Gnoj L.,
RA O'Shaughnessy A., Preston R., Habermann K., Murray J., Johnson D.,
RA Rohlfing T., Nelson J., Stoneking T., Pepin K., Spieth J., Sekhon M.,
RA Armstrong J., Becker M., Belter E., Cordum H., Cordes M., Courtney L.,
RA Courtney W., Dante M., Du H., Edwards J., Fryman J., Haakensen B.,
RA Lamar E., Latreille P., Leonard S., Meyer R., Mulvaney E., Ozersky P.,
RA Riley A., Strowmatt C., Wagner-McPherson C., Wollam A., Yoakum M., Bell M.,
RA Dedhia N., Parnell L., Shah R., Rodriguez M., Hoon See L., Vil D.,
RA Baker J., Kirchoff K., Toth K., King L., Bahret A., Miller B., Marra M.A.,
RA Martienssen R., McCombie W.R., Wilson R.K., Murphy G., Bancroft I.,
RA Volckaert G., Wambutt R., Duesterhoeft A., Stiekema W., Pohl T.,
RA Entian K.-D., Terryn N., Hartley N., Bent E., Johnson S., Langham S.-A.,
RA McCullagh B., Robben J., Grymonprez B., Zimmermann W., Ramsperger U.,
RA Wedler H., Balke K., Wedler E., Peters S., van Staveren M., Dirkse W.,
RA Mooijman P., Klein Lankhorst R., Weitzenegger T., Bothe G., Rose M.,
RA Hauf J., Berneiser S., Hempel S., Feldpausch M., Lamberth S.,
RA Villarroel R., Gielen J., Ardiles W., Bents O., Lemcke K., Kolesov G.,
RA Mayer K.F.X., Rudd S., Schoof H., Schueller C., Zaccaria P., Mewes H.-W.,
RA Bevan M., Fransz P.F.;
RT "Sequence and analysis of chromosome 5 of the plant Arabidopsis thaliana.";
RL Nature 408:823-826(2000).
RN [4]
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 [5]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
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 NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
RC STRAIN=cv. Columbia;
RA Cheuk R.F., Chen H., Kim C.J., Shinn P., Ecker J.R.;
RT "Arabidopsis ORF clones.";
RL Submitted (JAN-2005) to the EMBL/GenBank/DDBJ databases.
RN [7]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
RA Brover V.V., Troukhan M.E., Alexandrov N.A., Lu Y.-P., Flavell R.B.,
RA Feldmann K.A.;
RT "Full-length cDNA from Arabidopsis thaliana.";
RL Submitted (MAR-2002) to the EMBL/GenBank/DDBJ databases.
RN [8]
RP FUNCTION, AND TISSUE SPECIFICITY.
RX PubMed=13678595; DOI=10.1016/j.cub.2003.08.027;
RA Roeder A.H.K., Ferrandiz C., Yanofsky M.F.;
RT "The role of the REPLUMLESS homeodomain protein in patterning the
RT Arabidopsis fruit.";
RL Curr. Biol. 13:1630-1635(2003).
RN [9]
RP FUNCTION, TISSUE SPECIFICITY, AND INTERACTION WITH STM AND KNAT1/BP.
RX PubMed=12897247; DOI=10.1105/tpc.012856;
RA Smith H.M.S., Hake S.;
RT "The interaction of two homeobox genes, BREVIPEDICELLUS and PENNYWISE,
RT regulates internode patterning in the Arabidopsis inflorescence.";
RL Plant Cell 15:1717-1727(2003).
RN [10]
RP GENE FAMILY ORGANIZATION, AND FUNCTION.
RX PubMed=15120075; DOI=10.1016/j.cub.2004.04.032;
RA Smith H.M.S., Campbell B.C.C., Hake S.;
RT "Competence to respond to floral inductive signals requires the homeobox
RT genes PENNYWISE and POUND-FOOLISH.";
RL Curr. Biol. 14:812-817(2004).
RN [11]
RP FUNCTION.
RX PubMed=15155890; DOI=10.1105/tpc.021147;
RA Bao X., Franks R.G., Levin J.Z., Liu Z.;
RT "Repression of AGAMOUS by BELLRINGER in floral and inflorescence
RT meristems.";
RL Plant Cell 16:1478-1489(2004).
RN [12]
RP INTERACTION WITH STM.
RX PubMed=16513846; DOI=10.1093/nar/gkl016;
RA Cole M., Nolte C., Werr W.;
RT "Nuclear import of the transcription factor SHOOT MERISTEMLESS depends on
RT heterodimerization with BLH proteins expressed in discrete sub-domains of
RT the shoot apical meristem of Arabidopsis thaliana.";
RL Nucleic Acids Res. 34:1281-1292(2006).
RN [13]
RP FUNCTION, AND INTERACTION WITH STM AND KNAT1/BP.
RX PubMed=16741748; DOI=10.1007/s00425-006-0298-9;
RA Kanrar S., Onguka O., Smith H.M.S.;
RT "Arabidopsis inflorescence architecture requires the activities of KNOX-
RT BELL homeodomain heterodimers.";
RL Planta 224:1163-1173(2006).
RN [14]
RP INTERACTION WITH KNATM.
RX PubMed=18398054; DOI=10.1105/tpc.108.058495;
RA Magnani E., Hake S.;
RT "KNOX lost the OX: the Arabidopsis KNATM gene defines a novel class of KNOX
RT transcriptional regulators missing the homeodomain.";
RL Plant Cell 20:875-887(2008).
CC -!- FUNCTION: Transcription factor that is involved in the preservation of
CC the spiral phyllotactic arrangement leading to a regular pattern of
CC organ initiation. Required for maintenance of stem cell fate in the
CC shoot apical meristem, and is essential for specifying floral primordia
CC and establishing early internode patterning events during inflorescence
CC development. Acts as a transcription repressor of AG expression in
CC floral and inflorescence meristems. Is also responsive of the nuclear
CC import of SHOOT MERISTEMLESS (STM). In the fruit, plays a central role
CC in patterning by negatively regulating SHP expression in order to
CC prevent replum cells from adopting a valve margin cell fate.
CC {ECO:0000269|PubMed:12874117, ECO:0000269|PubMed:12897247,
CC ECO:0000269|PubMed:13678595, ECO:0000269|PubMed:15019989,
CC ECO:0000269|PubMed:15120075, ECO:0000269|PubMed:15155890,
CC ECO:0000269|PubMed:16741748}.
CC -!- SUBUNIT: May form heterodimeric complex with the TALE/KNOX proteins
CC STM, KNAT1/BP and KNAT6 (PubMed:12897247, PubMed:16513846,
CC PubMed:16741748). Interacts with KNATM, isoform KNATM-B
CC (PubMed:18398054). {ECO:0000269|PubMed:12897247,
CC ECO:0000269|PubMed:16513846, ECO:0000269|PubMed:16741748,
CC ECO:0000269|PubMed:18398054}.
CC -!- INTERACTION:
CC Q9LZM8; P46639: KNAT1; NbExp=9; IntAct=EBI-530473, EBI-530486;
CC Q9LZM8; P46640: KNAT2; NbExp=4; IntAct=EBI-530473, EBI-1153809;
CC Q9LZM8; Q84JS6: KNAT6; NbExp=6; IntAct=EBI-530473, EBI-530499;
CC Q9LZM8; Q38874: STM; NbExp=9; IntAct=EBI-530473, EBI-530523;
CC -!- SUBCELLULAR LOCATION: Nucleus {ECO:0000305}.
CC -!- TISSUE SPECIFICITY: Predominantly expressed in stems, inflorescences
CC and flowers. {ECO:0000269|PubMed:12874117, ECO:0000269|PubMed:12897247,
CC ECO:0000269|PubMed:13678595, ECO:0000269|PubMed:15019989}.
CC -!- DEVELOPMENTAL STAGE: Expressed during late heart stage embryo in the
CC SAM region between the two developing cotyledons. Expression is
CC initially down-regulated in incipient floral primordia but a strong
CC expression is subsequently detected in the central region of young
CC floral primordia and the inner whorl of developing flowers. In fruit,
CC is expressed in the replum of developing ovaries.
CC {ECO:0000269|PubMed:12874117}.
CC -!- DOMAIN: The SR/KY and BELL domains are responsive for the interaction
CC between the TALE/BELL proteins and the TALE/KNOX proteins.
CC -!- SIMILARITY: Belongs to the TALE/BELL homeobox family. {ECO:0000305}.
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DR EMBL; AY328314; AAP93641.1; -; mRNA.
DR EMBL; AJ519491; CAD58040.1; -; mRNA.
DR EMBL; AL162508; CAB82976.1; -; Genomic_DNA.
DR EMBL; CP002688; AED90423.1; -; Genomic_DNA.
DR EMBL; BT000779; AAN31918.1; -; mRNA.
DR EMBL; BT020537; AAW70383.1; -; mRNA.
DR EMBL; AY084243; AAM60839.1; -; mRNA.
DR PIR; T48224; T48224.
DR RefSeq; NP_195823.1; NM_120281.3.
DR AlphaFoldDB; Q9LZM8; -.
DR SMR; Q9LZM8; -.
DR BioGRID; 17021; 15.
DR IntAct; Q9LZM8; 12.
DR STRING; 3702.Q9LZM8; -.
DR PaxDb; 3702-AT5G02030-1; -.
DR ProteomicsDB; 240434; -.
DR EnsemblPlants; AT5G02030.1; AT5G02030.1; AT5G02030.
DR GeneID; 831745; -.
DR Gramene; AT5G02030.1; AT5G02030.1; AT5G02030.
DR KEGG; ath:AT5G02030; -.
DR Araport; AT5G02030; -.
DR TAIR; AT5G02030; RPL.
DR eggNOG; KOG0773; Eukaryota.
DR HOGENOM; CLU_011058_3_1_1; -.
DR InParanoid; Q9LZM8; -.
DR OMA; CNVGRGI; -.
DR OrthoDB; 468484at2759; -.
DR PhylomeDB; Q9LZM8; -.
DR PRO; PR:Q9LZM8; -.
DR Proteomes; UP000006548; Chromosome 5.
DR ExpressionAtlas; Q9LZM8; baseline and differential.
DR Genevisible; Q9LZM8; AT.
DR GO; GO:0005829; C:cytosol; 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:0043565; F:sequence-specific DNA binding; IDA:TAIR.
DR GO; GO:0048457; P:floral whorl morphogenesis; IMP:TAIR.
DR GO; GO:0010154; P:fruit development; IMP:TAIR.
DR GO; GO:0080006; P:internode patterning; IMP:TAIR.
DR GO; GO:0010076; P:maintenance of floral meristem identity; IGI:TAIR.
DR GO; GO:0010077; P:maintenance of inflorescence meristem identity; IGI:TAIR.
DR GO; GO:0045892; P:negative regulation of DNA-templated transcription; IMP:TAIR.
DR GO; GO:0007389; P:pattern specification process; IMP:TAIR.
DR GO; GO:1905393; P:plant organ formation; IGI:TAIR.
DR GO; GO:0010223; P:secondary shoot formation; IGI:TAIR.
DR GO; GO:0048367; P:shoot system development; IGI:TAIR.
DR GO; GO:0010228; P:vegetative to reproductive phase transition of meristem; IGI:TAIR.
DR GO; GO:0010051; P:xylem and phloem pattern formation; IMP:TAIR.
DR GO; GO:0010089; P:xylem development; IMP:TAIR.
DR CDD; cd00086; homeodomain; 1.
DR Gene3D; 1.10.10.60; Homeodomain-like; 1.
DR InterPro; IPR009057; Homeobox-like_sf.
DR InterPro; IPR001356; Homeobox_dom.
DR InterPro; IPR008422; Homeobox_KN_domain.
DR InterPro; IPR006563; POX_dom.
DR PANTHER; PTHR11850:SF395; BEL1-LIKE HOMEODOMAIN PROTEIN 9; 1.
DR PANTHER; PTHR11850; HOMEOBOX PROTEIN TRANSCRIPTION FACTORS; 1.
DR Pfam; PF05920; Homeobox_KN; 1.
DR Pfam; PF07526; POX; 1.
DR SMART; SM00389; HOX; 1.
DR SMART; SM00574; POX; 1.
DR SUPFAM; SSF46689; Homeodomain-like; 1.
DR PROSITE; PS00027; HOMEOBOX_1; 1.
DR PROSITE; PS50071; HOMEOBOX_2; 1.
PE 1: Evidence at protein level;
KW DNA-binding; Homeobox; Nucleus; Reference proteome; Transcription;
KW Transcription regulation.
FT CHAIN 1..575
FT /note="BEL1-like homeodomain protein 9"
FT /id="PRO_0000315465"
FT DNA_BIND 348..410
FT /note="Homeobox"
FT /evidence="ECO:0000255|PROSITE-ProRule:PRU00108"
FT REGION 24..43
FT /note="Disordered"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT REGION 174..190
FT /note="SR/KY domain"
FT REGION 223..294
FT /note="BELL domain"
FT REGION 306..337
FT /note="Disordered"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT REGION 418..485
FT /note="Disordered"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT REGION 533..557
FT /note="Disordered"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT MOTIF 229..246
FT /note="Bipartite nuclear localization signal"
FT /evidence="ECO:0000255"
FT COMPBIAS 306..333
FT /note="Polar residues"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT COMPBIAS 420..462
FT /note="Polar residues"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT CONFLICT 14
FT /note="S -> G (in Ref. 5; AAN31918)"
FT /evidence="ECO:0000305"
FT CONFLICT 444
FT /note="S -> N (in Ref. 2; CAD58040 and 7; AAM60839)"
FT /evidence="ECO:0000305"
SQ SEQUENCE 575 AA; 62007 MW; 07663D53C5E85E4B CRC64;
MADAYEPYHV LQQSRRDKLR IPSLDSHFHF HPPPPPSSGG GGGVFPLADS DFLAAGGFHS
NNNNNHISNP SYSNFMGFLG GPSSSSSTAV AVAGDHSFNA GLSSGDVLVF KPEPLSLSLS
SHPRLAYDLV VPGVVNSGFC RSAGEANAAA VTIASRSSGP LGPFTGYASI LKGSRFLKPA
QMLLDEFCNV GRGIYTDKVI DDDDSSLLFD PTVENLCGVS DGGGGDNGKK KSKLISMLDE
VYKRYKQYYE QLQAVMGSFE CVAGLGHAAP YANLALKALS KHFKCLKNAI TDQLQFSHNN
KIQQQQQCGH PMNSENKTDS LRFGGSDSSR GLCSAGQRHG FPDHHAPVWR PHRGLPERAV
TVLRAWLFDH FLHPYPTDTD KLMLAKQTGL SRNQVSNWFI NARVRVWKPM VEEIHMLETR
QSQRSSSSSW RDERTSTTVF PDNSNNNPSS SSAQQRPNNS SPPRRARNDD VHGTNNNNSY
VNSGSGGGSA VGFSYGIGSS NVPVMNSSTN GGVSLTLGLH HQIGLPEPFP MTTAQRFGLD
GGSGDGGGGY EGQNRQFGRD FIGGSNHQFL HDFVG
//
