AS2_ARATH
ID AS2_ARATH Reviewed; 199 AA.
AC O04479; B7XG54; Q53YQ2;
DT 28-MAR-2003, integrated into UniProtKB/Swiss-Prot.
DT 01-JUL-1997, sequence version 1.
DT 03-AUG-2022, entry version 123.
DE RecName: Full=Protein ASYMMETRIC LEAVES 2 {ECO:0000303|PubMed:12040093};
DE AltName: Full=LOB domain-containing protein 6 {ECO:0000303|PubMed:12068116};
GN Name=AS2 {ECO:0000303|PubMed:12040093};
GN Synonyms=LBD6 {ECO:0000303|PubMed:12068116};
GN OrderedLocusNames=At1g65620 {ECO:0000312|Araport:AT1G65620};
GN ORFNames=F5I14.15 {ECO:0000312|EMBL:AAB60912.1};
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], TISSUE SPECIFICITY, GENE FAMILY, AND
RP NOMENCLATURE.
RC STRAIN=cv. Columbia;
RX PubMed=12068116; DOI=10.1104/pp.010926;
RA Shuai B., Reynaga-Pena C.G., Springer P.S.;
RT "The LATERAL ORGAN BOUNDARIES gene defines a novel, plant-specific gene
RT family.";
RL Plant Physiol. 129:747-761(2002).
RN [2]
RP NUCLEOTIDE SEQUENCE [MRNA], TISSUE SPECIFICITY, SUBCELLULAR LOCATION, GENE
RP FAMILY, AND NOMENCLATURE.
RC STRAIN=cv. Columbia;
RX PubMed=12040093; DOI=10.1093/pcp/pcf077;
RA Iwakawa H., Ueno Y., Semiarti E., Onouchi H., Kojima S., Tsukaya H.,
RA Hasebe M., Soma T., Ikezaki M., Machida C., Machida Y.;
RT "The ASYMMETRIC LEAVES2 gene of Arabidopsis thaliana, required for
RT formation of a symmetric flat leaf lamina, encodes a member of a novel
RT family of proteins characterized by cysteine repeats and a leucine
RT zipper.";
RL Plant Cell Physiol. 43:467-478(2002).
RN [3]
RP NUCLEOTIDE SEQUENCE [MRNA].
RC STRAIN=cv. Columbia;
RX PubMed=19154202; DOI=10.1111/j.1365-313x.2009.03797.x;
RA Matsumura Y., Iwakawa H., Machida Y., Machida C.;
RT "Characterization of genes in the ASYMMETRIC LEAVES2/LATERAL ORGAN
RT BOUNDARIES (AS2/LOB) family in Arabidopsis thaliana, and functional and
RT molecular comparisons between AS2 and other family members.";
RL Plant J. 58:525-537(2009).
RN [4]
RP NUCLEOTIDE SEQUENCE [GENOMIC DNA].
RC STRAIN=cv. Bla-1;
RA Lebedeva O.V., Ezhova T.A.;
RT "Gene AS2 from ecotype Blanes Arabidopsis thaliana.";
RL Submitted (JAN-2004) to the EMBL/GenBank/DDBJ databases.
RN [5]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC STRAIN=cv. Columbia;
RX PubMed=11130712; DOI=10.1038/35048500;
RA Theologis A., Ecker J.R., Palm C.J., Federspiel N.A., Kaul S., White O.,
RA Alonso J., Altafi H., Araujo R., Bowman C.L., Brooks S.Y., Buehler E.,
RA Chan A., Chao Q., Chen H., Cheuk R.F., Chin C.W., Chung M.K., Conn L.,
RA Conway A.B., Conway A.R., Creasy T.H., Dewar K., Dunn P., Etgu P.,
RA Feldblyum T.V., Feng J.-D., Fong B., Fujii C.Y., Gill J.E., Goldsmith A.D.,
RA Haas B., Hansen N.F., Hughes B., Huizar L., Hunter J.L., Jenkins J.,
RA Johnson-Hopson C., Khan S., Khaykin E., Kim C.J., Koo H.L.,
RA Kremenetskaia I., Kurtz D.B., Kwan A., Lam B., Langin-Hooper S., Lee A.,
RA Lee J.M., Lenz C.A., Li J.H., Li Y.-P., Lin X., Liu S.X., Liu Z.A.,
RA Luros J.S., Maiti R., Marziali A., Militscher J., Miranda M., Nguyen M.,
RA Nierman W.C., Osborne B.I., Pai G., Peterson J., Pham P.K., Rizzo M.,
RA Rooney T., Rowley D., Sakano H., Salzberg S.L., Schwartz J.R., Shinn P.,
RA Southwick A.M., Sun H., Tallon L.J., Tambunga G., Toriumi M.J., Town C.D.,
RA Utterback T., Van Aken S., Vaysberg M., Vysotskaia V.S., Walker M., Wu D.,
RA Yu G., Fraser C.M., Venter J.C., Davis R.W.;
RT "Sequence and analysis of chromosome 1 of the plant Arabidopsis thaliana.";
RL Nature 408:816-820(2000).
RN [6]
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 [7]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
RC STRAIN=cv. Columbia;
RA Shinn P., Chen H., Kim C.J., Ecker J.R.;
RT "Arabidopsis ORF clones.";
RL Submitted (APR-2006) to the EMBL/GenBank/DDBJ databases.
RN [8]
RP FUNCTION.
RX PubMed=11311158; DOI=10.1242/dev.128.10.1771;
RA Semiarti E., Ueno Y., Tsukaya H., Iwakawa H., Machida C., Machida Y.;
RT "The ASYMMETRIC LEAVES2 gene of Arabidopsis thaliana regulates formation of
RT a symmetric lamina, establishment of venation and repression of meristem-
RT related homeobox genes in leaves.";
RL Development 128:1771-1783(2001).
RN [9]
RP INDUCTION.
RX PubMed=11934861; DOI=10.1242/dev.129.8.1957;
RA Byrne M.E., Simorowski J., Martienssen R.A.;
RT "ASYMMETRIC LEAVES1 reveals knox gene redundancy in Arabidopsis.";
RL Development 129:1957-1965(2002).
RN [10]
RP FUNCTION, AND INTERACTION WITH AS1.
RX PubMed=12874130; DOI=10.1242/dev.00622;
RA Xu L., Xu Y., Dong A., Sun Y., Pi L., Xu Y., Huang H.;
RT "Novel as1 and as2 defects in leaf adaxial-abaxial polarity reveal the
RT requirement for ASYMMETRIC LEAVES1 and 2 and ERECTA functions in specifying
RT leaf adaxial identity.";
RL Development 130:4097-4107(2003).
RN [11]
RP FUNCTION.
RX PubMed=12787254; DOI=10.1046/j.1365-313x.2003.01758.x;
RA Nakazawa M., Ichikawa T., Ishikawa A., Kobayashi H., Tsuhara Y.,
RA Kawashima M., Suzuki K., Muto S., Matsui M.;
RT "Activation tagging, a novel tool to dissect the functions of a gene
RT family.";
RL Plant J. 34:741-750(2003).
RN [12]
RP FUNCTION.
RX PubMed=14508003; DOI=10.1105/tpc.014969;
RA Lin W.-C., Shuai B., Springer P.S.;
RT "The Arabidopsis LATERAL ORGAN BOUNDARIES-domain gene ASYMMETRIC LEAVES2
RT functions in the repression of KNOX gene expression and in adaxial-abaxial
RT patterning.";
RL Plant Cell 15:2241-2252(2003).
RN [13]
RP FUNCTION.
RX PubMed=16006579; DOI=10.1105/tpc.105.033449;
RA Li H., Xu L., Wang H., Yuan Z., Cao X., Yang Z., Zhang D., Xu Y., Huang H.;
RT "The Putative RNA-dependent RNA polymerase RDR6 acts synergistically with
RT ASYMMETRIC LEAVES1 and 2 to repress BREVIPEDICELLUS and MicroRNA165/166 in
RT Arabidopsis leaf development.";
RL Plant Cell 17:2157-2171(2005).
RN [14]
RP FUNCTION.
RX PubMed=16699177; DOI=10.1093/pcp/pcj057;
RA Xu L., Yang L., Pi L., Liu Q., Ling Q., Wang H., Poethig R.S., Huang H.;
RT "Genetic interaction between the AS1-AS2 and RDR6-SGS3-AGO7 pathways for
RT leaf morphogenesis.";
RL Plant Cell Physiol. 47:853-863(2006).
RN [15]
RP FUNCTION.
RX PubMed=17395603; DOI=10.1093/pcp/pcm040;
RA Fu Y., Xu L., Xu B., Yang L., Ling Q., Wang H., Huang H.;
RT "Genetic interactions between leaf polarity-controlling genes and
RT ASYMMETRIC LEAVES1 and 2 in Arabidopsis leaf patterning.";
RL Plant Cell Physiol. 48:724-735(2007).
RN [16]
RP FUNCTION, AND TISSUE SPECIFICITY.
RX PubMed=17559509; DOI=10.1111/j.1365-313x.2007.03132.x;
RA Iwakawa H., Iwasaki M., Kojima S., Ueno Y., Soma T., Tanaka H.,
RA Semiarti E., Machida Y., Machida C.;
RT "Expression of the ASYMMETRIC LEAVES2 gene in the adaxial domain of
RT Arabidopsis leaves represses cell proliferation in this domain and is
RT critical for the development of properly expanded leaves.";
RL Plant J. 51:173-184(2007).
RN [17]
RP FUNCTION, INTERACTION WITH AS1, AND SUBCELLULAR LOCATION.
RX PubMed=23271976; DOI=10.1371/journal.pgen.1003114;
RA Luo M., Yu C.W., Chen F.F., Zhao L., Tian G., Liu X., Cui Y., Yang J.Y.,
RA Wu K.;
RT "Histone deacetylase HDA6 is functionally associated with AS1 in repression
RT of KNOX genes in arabidopsis.";
RL PLoS Genet. 8:E1003114-E1003114(2012).
RN [18]
RP FUNCTION, AND DISRUPTION PHENOTYPE.
RX PubMed=27334696; DOI=10.1242/bio.019109;
RA Matsumura Y., Ohbayashi I., Takahashi H., Kojima S., Ishibashi N., Keta S.,
RA Nakagawa A., Hayashi R., Saez-Vasquez J., Echeverria M., Sugiyama M.,
RA Nakamura K., Machida C., Machida Y.;
RT "A genetic link between epigenetic repressor AS1-AS2 and a putative small
RT subunit processome in leaf polarity establishment of Arabidopsis.";
RL Biol. Open 5:942-954(2016).
RN [19]
RP INTERACTION WITH LFR.
RX PubMed=29775508; DOI=10.1111/tpj.13973;
RA Lin X., Gu D., Zhao H., Peng Y., Zhang G., Yuan T., Li M., Wang Z.,
RA Wang X., Cui S.;
RT "LFR is functionally associated with AS2 to mediate leaf development in
RT Arabidopsis.";
RL Plant J. 0:0-0(2018).
CC -!- FUNCTION: Negative regulator of cell proliferation in the adaxial side
CC of leaves. Regulates the formation of a symmetric lamina and the
CC establishment of venation. Positively regulates LATERAL ORGAN
CC BOUNDARIES (LOB) within the shoot apex, and the class III HD-ZIP genes
CC REV, PHB, and PHV. Interacts directly with ASYMMETRIC LEAVES 1 (AS1) to
CC repress the knox homeobox genes KNAT1, KNAT2, and KNAT6 and the abaxial
CC determinants ARF3, KAN2 and YAB5. May act in parallel with the RDR6-
CC SGS3-AGO7 pathway, an endogenous RNA silencing pathway, to regulate the
CC leaf morphogenesis (PubMed:11311158, PubMed:12787254, PubMed:12874130,
CC PubMed:14508003, PubMed:16006579, PubMed:16699177, PubMed:17395603,
CC PubMed:17559509). Required for the binding of AS1 to the KNOX genes
CC (PubMed:23271976). Involved in leaf polarity establishment by
CC functioning cooperatively with RH10 or RID2 to repress abaxial genes
CC ARF3, ARF4, KAN1, KAN2, YAB1 and YAB5, and the knox homeobox genes
CC KNAT1, KNAT2, KNAT6, and STM to promote adaxial development in leaf
CC primordia at shoot apical meristems at high temperatures
CC (PubMed:27334696). {ECO:0000269|PubMed:11311158,
CC ECO:0000269|PubMed:12787254, ECO:0000269|PubMed:12874130,
CC ECO:0000269|PubMed:14508003, ECO:0000269|PubMed:16006579,
CC ECO:0000269|PubMed:16699177, ECO:0000269|PubMed:17395603,
CC ECO:0000269|PubMed:17559509, ECO:0000269|PubMed:23271976,
CC ECO:0000269|PubMed:27334696}.
CC -!- SUBUNIT: Homo- and heterodimer with AS1 (PubMed:23271976). Interacts
CC with AS1 (PubMed:12874130, PubMed:23271976). Part of the AS1 repressor
CC complex composed of AS1, LBD6/AS2 and HDA6 (PubMed:23271976,
CC PubMed:12874130). Interacts with LFR (PubMed:29775508).
CC {ECO:0000269|PubMed:12874130, ECO:0000269|PubMed:23271976,
CC ECO:0000269|PubMed:29775508}.
CC -!- INTERACTION:
CC O04479; O80931: AS1; NbExp=4; IntAct=EBI-4443304, EBI-763232;
CC -!- SUBCELLULAR LOCATION: Nucleus {ECO:0000269|PubMed:12040093,
CC ECO:0000269|PubMed:23271976}.
CC -!- TISSUE SPECIFICITY: Expressed in young shoots, roots, stems, leaves,
CC flowers and adaxial domains of cotyledonary and leaves primordia.
CC {ECO:0000269|PubMed:12040093, ECO:0000269|PubMed:12068116,
CC ECO:0000269|PubMed:17559509}.
CC -!- INDUCTION: Negatively regulated by SHOOT MERISTEMLESS (STM).
CC {ECO:0000269|PubMed:11934861}.
CC -!- DISRUPTION PHENOTYPE: Plants are indistinguishable from that of wild-
CC type at 16 degrees Celsius, however they generate a weak phenotype of
CC pointed leaves at 22 degrees Celsius which become narrower at 26
CC degrees Celsius. In the leaves of plants grown at 26 degrees Celsius,
CC the xylems are located on the adaxial sides and the phloems are on the
CC abaxial sides, similar to those in the wild-type. Plants with double
CC mutations in this protein and in RH10 or AS1 protein have abaxialized
CC filamentous and trumpet-like leaves with loss of the adaxial domain at
CC high temperatures. In double mutants, shapes of epidermal cells of the
CC filamentous leaves are simple and rectangular, similar to those of a
CC petiole, but different from those of flat leaves of wild-type plants.
CC The filamentous leaves of the double mutant at 26 degrees Celsius show
CC primitive or no vascular tissue without apparent xylem cells inside the
CC bundle sheath, suggesting defects in differentiation of xylem cells on
CC the adaxial side. {ECO:0000269|PubMed:27334696}.
CC -!- MISCELLANEOUS: Gain-of-function mutants iso1-D and iso-2D (T-DNA
CC tagging) show completely curled up hyponastic rosette leaves.
CC -!- SIMILARITY: Belongs to the LOB domain-containing protein family.
CC {ECO:0000305}.
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DR EMBL; AF447887; AAL38032.1; -; mRNA.
DR EMBL; AB080802; BAB88693.1; -; mRNA.
DR EMBL; AB473833; BAH10544.1; -; mRNA.
DR EMBL; AY519483; AAS00609.1; -; Genomic_DNA.
DR EMBL; AC001229; AAB60912.1; -; Genomic_DNA.
DR EMBL; CP002684; AEE34401.1; -; Genomic_DNA.
DR EMBL; CP002684; AEE34402.1; -; Genomic_DNA.
DR EMBL; CP002684; AEE34403.1; -; Genomic_DNA.
DR EMBL; CP002684; AEE34404.1; -; Genomic_DNA.
DR EMBL; CP002684; ANM57892.1; -; Genomic_DNA.
DR EMBL; BT025163; ABE77401.1; -; mRNA.
DR PIR; C96681; C96681.
DR RefSeq; NP_001077777.1; NM_001084308.5.
DR RefSeq; NP_001077778.1; NM_001084309.2.
DR RefSeq; NP_001117553.1; NM_001124081.2.
DR RefSeq; NP_001320370.1; NM_001334217.1.
DR RefSeq; NP_176739.1; NM_105235.5.
DR AlphaFoldDB; O04479; -.
DR SMR; O04479; -.
DR BioGRID; 28094; 13.
DR IntAct; O04479; 2.
DR STRING; 3702.AT1G65620.1; -.
DR PaxDb; O04479; -.
DR PRIDE; O04479; -.
DR ProteomicsDB; 246912; -.
DR EnsemblPlants; AT1G65620.1; AT1G65620.1; AT1G65620.
DR EnsemblPlants; AT1G65620.2; AT1G65620.2; AT1G65620.
DR EnsemblPlants; AT1G65620.3; AT1G65620.3; AT1G65620.
DR EnsemblPlants; AT1G65620.4; AT1G65620.4; AT1G65620.
DR EnsemblPlants; AT1G65620.5; AT1G65620.5; AT1G65620.
DR GeneID; 842873; -.
DR Gramene; AT1G65620.1; AT1G65620.1; AT1G65620.
DR Gramene; AT1G65620.2; AT1G65620.2; AT1G65620.
DR Gramene; AT1G65620.3; AT1G65620.3; AT1G65620.
DR Gramene; AT1G65620.4; AT1G65620.4; AT1G65620.
DR Gramene; AT1G65620.5; AT1G65620.5; AT1G65620.
DR KEGG; ath:AT1G65620; -.
DR Araport; AT1G65620; -.
DR TAIR; locus:2034163; AT1G65620.
DR eggNOG; ENOG502QV43; Eukaryota.
DR HOGENOM; CLU_058353_1_2_1; -.
DR InParanoid; O04479; -.
DR OMA; QMMIRSF; -.
DR OrthoDB; 1117037at2759; -.
DR PhylomeDB; O04479; -.
DR PRO; PR:O04479; -.
DR Proteomes; UP000006548; Chromosome 1.
DR ExpressionAtlas; O04479; baseline and differential.
DR Genevisible; O04479; AT.
DR GO; GO:0005654; C:nucleoplasm; IDA:TAIR.
DR GO; GO:0005634; C:nucleus; IDA:TAIR.
DR GO; GO:0009943; P:adaxial/abaxial axis specification; IMP:TAIR.
DR GO; GO:0048441; P:petal development; IGI:TAIR.
DR GO; GO:0009944; P:polarity specification of adaxial/abaxial axis; IMP:TAIR.
DR GO; GO:0009954; P:proximal/distal pattern formation; IGI:TAIR.
DR GO; GO:0009799; P:specification of symmetry; IMP:TAIR.
DR InterPro; IPR004883; LOB.
DR Pfam; PF03195; LOB; 1.
DR PROSITE; PS50891; LOB; 1.
PE 1: Evidence at protein level;
KW Developmental protein; Nucleus; Reference proteome; Repressor;
KW Stress response.
FT CHAIN 1..199
FT /note="Protein ASYMMETRIC LEAVES 2"
FT /id="PRO_0000132257"
FT DOMAIN 8..109
FT /note="LOB"
FT /evidence="ECO:0000255|PROSITE-ProRule:PRU00291"
SQ SEQUENCE 199 AA; 21710 MW; CF75859362F36686 CRC64;
MASSSTNSPC AACKFLRRKC QPECVFAPYF PPDQPQKFAN VHKVFGASNV TKLLNELHPS
QREDAVNSLA YEADMRLRDP VYGCVGVISL LQHQLRQLQI DLSCAKSELS KYQSLGILAA
THQSLGINLL AGAADGTATA VRDHYHHHQF FPREQMFGGL DVPAGNNYDG GILAIGQITQ
FQQPRAAAGD DGRRTVDPS
