{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"Q0WSN2","secondaryAccessions":["O22291"],"uniProtkbId":"DAR2_ARATH","entryAudit":{"firstPublicDate":"2010-08-10","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2006-09-05","entryVersion":115,"sequenceVersion":1},"annotationScore":5.0,"organism":{"scientificName":"Arabidopsis thaliana","commonName":"Mouse-ear cress","taxonId":3702,"lineage":["Eukaryota","Viridiplantae","Streptophyta","Embryophyta","Tracheophyta","Spermatophyta","Magnoliopsida","eudicotyledons","Gunneridae","Pentapetalae","rosids","malvids","Brassicales","Brassicaceae","Camelineae","Arabidopsis"]},"proteinExistence":"1: Evidence at protein level","proteinDescription":{"recommendedName":{"fullName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"18483219"}],"value":"Protein DA1-related 2"}},"alternativeNames":[{"fullName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"21749503"}],"value":"Protein LATERAL ROOT DEVELOPMENT 3"}}]},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"18483219"}],"value":"DAR2"},"synonyms":[{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"21749503"}],"value":"LRD3"}],"orderedLocusNames":[{"value":"At2g39830"}],"orfNames":[{"value":"T5I7.13"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21749503"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23296689"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25757472"}],"value":"Acts redundantly with DA1 and DAR1 to regulate endoreduplication during leaf development. Together with DA1 and DAR1, modulates the protein stability of the transcription factors TCP14 and TCP15, which repress endoreduplication by directly regulating the expression of cell-cycle genes (PubMed:25757472). Involved in root phloem development. Is an essential component of early phloem development, long-distance delivery of phloem content, and proper maintenance of root system architecture (PubMed:21749503). Involved in the control of root meristem size. Functions genetically downstream of cytokinin and IAA3 to maintain normal auxin distribution by influencing polar auxin transport. Acts through the PLETHORA pathway, upstream of PLT1 and PLT2 to influence root stem cell niche activity and thus control root meristem size (PubMed:23296689)"}],"commentType":"FUNCTION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25757472"}],"value":"Interacts with ubiquitin, TCP14 and TCP15"}],"commentType":"SUBUNIT"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21749503"}],"value":"Expressed in the vasculature of leaves, inflorescence stems, flowers, hypocotyls, and primary and lateral roots. In roots, expressed in phloem companion cells"}],"commentType":"TISSUE SPECIFICITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25757472"}],"value":"Highly expressed during early stages leaf development, and expression decreases at the later stages of leaf development"}],"commentType":"DEVELOPMENTAL STAGE"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000305","source":"PubMed","id":"25757472"}],"value":"The UIM domains bind molecules modified by monoubiquitin or ubiquitin chains and promote coupled monoubiquitination"}],"commentType":"DOMAIN"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"28167503"}],"value":"Polyubiquitinated by DA2"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21749503"}],"value":"Reduced primary root growth, increased lateral root formation, and altered root system architecture, due to altered early phloem development"}],"commentType":"DISRUPTION PHENOTYPE"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21749503"}],"value":"Exogenous treatment with auxin rescues the phloem defects of the lrd3 knockout mutant"}],"commentType":"MISCELLANEOUS"},{"commentType":"SEQUENCE CAUTION","sequenceCautionType":"Erroneous gene model prediction","sequence":"AAB87132.1","evidences":[{"evidenceCode":"ECO:0000305"}]}],"features":[{"type":"Chain","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":528,"modifier":"EXACT"}},"description":"Protein DA1-related 2","featureId":"PRO_0000396937"},{"type":"Domain","location":{"start":{"value":99,"modifier":"EXACT"},"end":{"value":118,"modifier":"EXACT"}},"description":"UIM 1","evidences":[{"evidenceCode":"ECO:0000305"}]},{"type":"Domain","location":{"start":{"value":127,"modifier":"EXACT"},"end":{"value":148,"modifier":"EXACT"}},"description":"UIM 2; degenerate","evidences":[{"evidenceCode":"ECO:0000305"}]},{"type":"Domain","location":{"start":{"value":160,"modifier":"EXACT"},"end":{"value":220,"modifier":"EXACT"}},"description":"LIM zinc-binding","evidences":[{"evidenceCode":"ECO:0000255","source":"PROSITE-ProRule","id":"PRU00125"}]},{"type":"Region","location":{"start":{"value":60,"modifier":"EXACT"},"end":{"value":108,"modifier":"EXACT"}},"description":"Disordered","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"MobiDB-lite"}]},{"type":"Region","location":{"start":{"value":447,"modifier":"EXACT"},"end":{"value":474,"modifier":"EXACT"}},"description":"Disordered","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"MobiDB-lite"}]},{"type":"Compositional bias","location":{"start":{"value":91,"modifier":"EXACT"},"end":{"value":106,"modifier":"EXACT"}},"description":"Basic and acidic residues","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"MobiDB-lite"}]},{"type":"Compositional bias","location":{"start":{"value":455,"modifier":"EXACT"},"end":{"value":470,"modifier":"EXACT"}},"description":"Low complexity","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"MobiDB-lite"}]}],"keywords":[{"id":"KW-0440","category":"Domain","name":"LIM domain"},{"id":"KW-0479","category":"Ligand","name":"Metal-binding"},{"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"id":"KW-0677","category":"Domain","name":"Repeat"},{"id":"KW-0832","category":"PTM","name":"Ubl conjugation"},{"id":"KW-0862","category":"Ligand","name":"Zinc"}],"references":[{"referenceNumber":1,"citation":{"id":"10617197","citationType":"journal article","authors":["Lin X.","Kaul S.","Rounsley S.D.","Shea T.P.","Benito M.-I.","Town C.D.","Fujii C.Y.","Mason T.M.","Bowman C.L.","Barnstead M.E.","Feldblyum T.V.","Buell C.R.","Ketchum K.A.","Lee J.J.","Ronning C.M.","Koo H.L.","Moffat K.S.","Cronin L.A.","Shen M.","Pai G.","Van Aken S.","Umayam L.","Tallon L.J.","Gill J.E.","Adams M.D.","Carrera A.J.","Creasy T.H.","Goodman H.M.","Somerville C.R.","Copenhaver G.P.","Preuss D.","Nierman W.C.","White O.","Eisen J.A.","Salzberg S.L.","Fraser C.M.","Venter J.C."],"citationCrossReferences":[{"database":"PubMed","id":"10617197"},{"database":"DOI","id":"10.1038/45471"}],"title":"Sequence and analysis of chromosome 2 of the plant Arabidopsis thaliana.","publicationDate":"1999","journal":"Nature","firstPage":"761","lastPage":"768","volume":"402"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"referenceComments":[{"value":"cv. Columbia","type":"STRAIN"}]},{"referenceNumber":2,"citation":{"id":"27862469","citationType":"journal article","authors":["Cheng C.Y.","Krishnakumar V.","Chan A.P.","Thibaud-Nissen F.","Schobel S.","Town C.D."],"citationCrossReferences":[{"database":"PubMed","id":"27862469"},{"database":"DOI","id":"10.1111/tpj.13415"}],"title":"Araport11: a complete reannotation of the Arabidopsis thaliana reference genome.","publicationDate":"2017","journal":"Plant J.","firstPage":"789","lastPage":"804","volume":"89"},"referencePositions":["GENOME REANNOTATION"],"referenceComments":[{"value":"cv. Columbia","type":"STRAIN"}]},{"referenceNumber":3,"citation":{"id":"CI-CI4AM1G71UC0S","citationType":"submission","authoringGroup":["The Arabidopsis Information Resource (TAIR)"],"title":"Community-consensus reference genome of Arabidopsis thaliana Col-0.","publicationDate":"FEB-2026","submissionDatabase":"EMBL/GenBank/DDBJ databases"},"referencePositions":["GENOME REANNOTATION"],"referenceComments":[{"value":"cv. Columbia","type":"STRAIN"}]},{"referenceNumber":4,"citation":{"id":"CI-1ND7IVR87IF4I","citationType":"submission","authors":["Totoki Y.","Seki M.","Ishida J.","Nakajima M.","Enju A.","Kamiya A.","Narusaka M.","Shin-i T.","Nakagawa M.","Sakamoto N.","Oishi K.","Kohara Y.","Kobayashi M.","Toyoda A.","Sakaki Y.","Sakurai T.","Iida K.","Akiyama K.","Satou M.","Toyoda T.","Konagaya A.","Carninci P.","Kawai J.","Hayashizaki Y.","Shinozaki K."],"title":"Large-scale analysis of RIKEN Arabidopsis full-length (RAFL) cDNAs.","publicationDate":"JUL-2006","submissionDatabase":"EMBL/GenBank/DDBJ databases"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA]"],"referenceComments":[{"value":"cv. Columbia","type":"STRAIN"}]},{"referenceNumber":5,"citation":{"id":"18483219","citationType":"journal article","authors":["Li Y.","Zheng L.","Corke F.","Smith C.","Bevan M.W."],"citationCrossReferences":[{"database":"PubMed","id":"18483219"},{"database":"DOI","id":"10.1101/gad.463608"}],"title":"Control of final seed and organ size by the DA1 gene family in Arabidopsis thaliana.","publicationDate":"2008","journal":"Genes Dev.","firstPage":"1331","lastPage":"1336","volume":"22"},"referencePositions":["GENE FAMILY","NOMENCLATURE"]},{"referenceNumber":6,"citation":{"id":"21749503","citationType":"journal article","authors":["Ingram P.","Dettmer J.","Helariutta Y.","Malamy J.E."],"citationCrossReferences":[{"database":"PubMed","id":"21749503"},{"database":"DOI","id":"10.1111/j.1365-313x.2011.04700.x"}],"title":"Arabidopsis Lateral Root Development 3 is essential for early phloem development and function, and hence for normal root system development.","publicationDate":"2011","journal":"Plant J.","firstPage":"455","lastPage":"467","volume":"68"},"referencePositions":["FUNCTION","TISSUE SPECIFICITY","DISRUPTION PHENOTYPE"]},{"referenceNumber":7,"citation":{"id":"23296689","citationType":"journal article","authors":["Peng Y.","Ma W.","Chen L.","Yang L.","Li S.","Zhao H.","Zhao Y.","Jin W.","Li N.","Bevan M.W.","Li X.","Tong Y.","Li Y."],"citationCrossReferences":[{"database":"PubMed","id":"23296689"},{"database":"DOI","id":"10.1104/pp.112.210237"}],"title":"Control of root meristem size by DA1-RELATED PROTEIN2 in Arabidopsis.","publicationDate":"2013","journal":"Plant Physiol.","firstPage":"1542","lastPage":"1556","volume":"161"},"referencePositions":["FUNCTION"]},{"referenceNumber":8,"citation":{"id":"25757472","citationType":"journal article","authors":["Peng Y.","Chen L.","Lu Y.","Wu Y.","Dumenil J.","Zhu Z.","Bevan M.W.","Li Y."],"citationCrossReferences":[{"database":"PubMed","id":"25757472"},{"database":"DOI","id":"10.1105/tpc.114.132274"}],"title":"The ubiquitin receptors DA1, DAR1, and DAR2 redundantly regulate endoreduplication by modulating the stability of TCP14/15 in Arabidopsis.","publicationDate":"2015","journal":"Plant Cell","firstPage":"649","lastPage":"662","volume":"27"},"referencePositions":["FUNCTION","INTERACTION WITH UBIQUITIN; TCP14 AND TCP15","DEVELOPMENTAL STAGE","DOMAIN"]},{"referenceNumber":9,"citation":{"id":"28167503","citationType":"journal article","authors":["Dong H.","Dumenil J.","Lu F.H.","Na L.","Vanhaeren H.","Naumann C.","Klecker M.","Prior R.","Smith C.","McKenzie N.","Saalbach G.","Chen L.","Xia T.","Gonzalez N.","Seguela M.","Inze D.","Dissmeyer N.","Li Y.","Bevan M.W."],"citationCrossReferences":[{"database":"PubMed","id":"28167503"},{"database":"DOI","id":"10.1101/gad.292235.116"}],"title":"Ubiquitylation activates a peptidase that promotes cleavage and destabilization of its activating E3 ligases and diverse growth regulatory proteins to limit cell proliferation in Arabidopsis.","publicationDate":"2017","journal":"Genes Dev.","firstPage":"197","lastPage":"208","volume":"31"},"referencePositions":["UBIQUITINATION"]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC003000","properties":[{"key":"ProteinId","value":"AAB87132.1"},{"key":"Status","value":"ALT_SEQ"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"EMBL","id":"CP002685","properties":[{"key":"ProteinId","value":"AEC09736.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"EMBL","id":"OZ408684","properties":[{"key":"ProteinId","value":"CAO7038109.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"EMBL","id":"AK227895","properties":[{"key":"ProteinId","value":"BAE99866.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"mRNA"}]},{"database":"PIR","id":"T01013","properties":[{"key":"EntryName","value":"T01013"}]},{"database":"RefSeq","id":"NP_181513.3","properties":[{"key":"NucleotideSequenceId","value":"NM_129542.5"}]},{"database":"AlphaFoldDB","id":"Q0WSN2","properties":[{"key":"Description","value":"-"}]},{"database":"BioGRID","id":"3908","properties":[{"key":"Interactions","value":"5"}]},{"database":"FunCoup","id":"Q0WSN2","properties":[{"key":"Number of interactors","value":"131"}]},{"database":"IntAct","id":"Q0WSN2","properties":[{"key":"Interactions","value":"2"}]},{"database":"STRING","id":"3702.Q0WSN2","properties":[{"key":"Description","value":"-"}]},{"database":"iPTMnet","id":"Q0WSN2","properties":[{"key":"Description","value":"-"}]},{"database":"PaxDb","id":"3702-Q0WSN2","properties":[{"key":"Description","value":"-"}]},{"database":"ProteomicsDB","id":"224676","properties":[{"key":"Description","value":"-"}]},{"database":"GeneID","id":"818570","properties":[{"key":"Description","value":"-"}]},{"database":"KEGG","id":"ath:AT2G39830","properties":[{"key":"Description","value":"-"}]},{"database":"Araport","id":"AT2G39830","properties":[{"key":"Description","value":"-"}]},{"database":"TAIR","id":"AT2G39830","properties":[{"key":"GeneName","value":"DAR2"}]},{"database":"eggNOG","id":"KOG1703","properties":[{"key":"ToxonomicScope","value":"Eukaryota"}]},{"database":"HOGENOM","id":"CLU_015906_4_0_1","properties":[{"key":"Description","value":"-"}]},{"database":"InParanoid","id":"Q0WSN2","properties":[{"key":"Description","value":"-"}]},{"database":"OrthoDB","id":"25414at2759","properties":[{"key":"Description","value":"-"}]},{"database":"PhylomeDB","id":"Q0WSN2","properties":[{"key":"Description","value":"-"}]},{"database":"PRO","id":"PR:Q0WSN2","properties":[{"key":"Description","value":"-"}]},{"database":"Proteomes","id":"UP000006548","properties":[{"key":"Component","value":"Chromosome 2"}]},{"database":"ExpressionAtlas","id":"Q0WSN2","properties":[{"key":"ExpressionPatterns","value":"baseline and differential"}]},{"database":"GO","id":"GO:0009507","properties":[{"key":"GoTerm","value":"C:chloroplast"},{"key":"GoEvidenceType","value":"ISM:TAIR"}]},{"database":"GO","id":"GO:0008233","properties":[{"key":"GoTerm","value":"F:peptidase activity"},{"key":"GoEvidenceType","value":"IDA:TAIR"}],"evidences":[{"evidenceCode":"ECO:0000314","source":"PubMed","id":"32209225"}]},{"database":"GO","id":"GO:0043130","properties":[{"key":"GoTerm","value":"F:ubiquitin binding"},{"key":"GoEvidenceType","value":"ISS:UniProtKB"}]},{"database":"GO","id":"GO:0010088","properties":[{"key":"GoTerm","value":"P:phloem development"},{"key":"GoEvidenceType","value":"IMP:TAIR"}],"evidences":[{"evidenceCode":"ECO:0000315","source":"PubMed","id":"21749503"}]},{"database":"GO","id":"GO:0032875","properties":[{"key":"GoTerm","value":"P:regulation of DNA endoreduplication"},{"key":"GoEvidenceType","value":"IMP:UniProtKB"}],"evidences":[{"evidenceCode":"ECO:0000315","source":"PubMed","id":"25757472"}]},{"database":"GO","id":"GO:0048364","properties":[{"key":"GoTerm","value":"P:root development"},{"key":"GoEvidenceType","value":"IMP:TAIR"}],"evidences":[{"evidenceCode":"ECO:0000315","source":"PubMed","id":"21749503"}]},{"database":"CDD","id":"cd09396","properties":[{"key":"EntryName","value":"LIM_DA1"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.10.110.10:FF:000107","properties":[{"key":"EntryName","value":"Protein DA1-related 2"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.10.110.10","properties":[{"key":"EntryName","value":"Cysteine Rich Protein"},{"key":"MatchStatus","value":"1"}]},{"database":"InterPro","id":"IPR045218","properties":[{"key":"EntryName","value":"DA1-like"}]},{"database":"InterPro","id":"IPR022087","properties":[{"key":"EntryName","value":"DA1-like_dom"}]},{"database":"InterPro","id":"IPR001781","properties":[{"key":"EntryName","value":"Znf_LIM"}]},{"database":"PANTHER","id":"PTHR24209","properties":[{"key":"EntryName","value":"PROTEIN DA1-RELATED 2"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR24209:SF7","properties":[{"key":"EntryName","value":"PROTEIN DA1-RELATED 2"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF12315","properties":[{"key":"EntryName","value":"DA1-like"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00412","properties":[{"key":"EntryName","value":"LIM"},{"key":"MatchStatus","value":"1"}]},{"database":"SMART","id":"SM00132","properties":[{"key":"EntryName","value":"LIM"},{"key":"MatchStatus","value":"1"}]},{"database":"SUPFAM","id":"SSF57716","properties":[{"key":"EntryName","value":"Glucocorticoid receptor-like (DNA-binding domain)"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00478","properties":[{"key":"EntryName","value":"LIM_DOMAIN_1"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS50023","properties":[{"key":"EntryName","value":"LIM_DOMAIN_2"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00142","properties":[{"key":"EntryName","value":"ZINC_PROTEASE"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MDSSSSSSSSSPSSSYGVARVSHISNPCIFGEVGSSSSSTYRDKKWKLMKWVSKLFKSGSNGGGSGAHTNHHPPQFQEDENMVFPLPPSSLDDRSRGARDKEELDRSISLSLADNTKRPHGYGWSMDNNRDFPRPFHGGLNPSSFIPPYEPSYQYRRRQRICGGCNSDIGSGNYLGCMGTFFHPECFRCHSCGYAITEHEFSLSGTKPYHKLCFKELTHPKCEVCHHFIPTNDAGLIEYRCHPFWNQKYCPSHEYDKTARCCSCERLESWDVRYYTLEDGRSLCLECMETAITDTGECQPLYHAIRDYYEGMYMKLDQQIPMLLVQREALNDAIVGEKNGYHHMPETRGLCLSEEQTVTSVLRRPRLGAHRLVGMRTQPQRLTRKCEVTAILVLYGLPRLLTGAILAHELMHGWLRLNGFRNLNPEVEEGICQVLSYMWLESEVLSDPSTRNLPSTSSVATSSSSSFSNKKGGKSNVEKKLGEFFKHQIAHDASPAYGGGFRAANAAACKYGLRRTLDHIRLTGTFPL","length":528,"molWeight":59426,"crc64":"A17599D7815012D6","md5":"A36684886D90698E5E10B5CC427FD889"},"extraAttributes":{"countByCommentType":{"FUNCTION":1,"SUBUNIT":1,"TISSUE SPECIFICITY":1,"DEVELOPMENTAL STAGE":1,"DOMAIN":1,"PTM":1,"DISRUPTION PHENOTYPE":1,"MISCELLANEOUS":1,"SEQUENCE CAUTION":1},"countByFeatureType":{"Chain":1,"Domain":3,"Region":2,"Compositional bias":2},"uniParcId":"UPI0000DB51F5"}}