Q42371 (ERECT_ARATH) Reviewed, UniProtKB/Swiss-Prot
Last modified
May 1, 2013.
Version 106.
History...
Names·Attributes·General annotation·Ontologies·Sequence annotation·Sequences·References·Web links·Cross-refs·Entry info·DocumentsCustomize order
Names·Attributes·General annotation·Ontologies·Sequence annotation·Sequences·References·Web links·Cross-refs·Entry info·DocumentsCustomize orderNames and origin
| Protein names | Recommended name: LRR receptor-like serine/threonine-protein kinase ERECTA EC=2.7.11.1 Alternative name(s): Protein QUANTITATIVE RESISTANCE TO PLECTOSPHAERELLA 1 Protein QUANTITATIVE RESISTANCE TO RALSTONIA SOLANACEARUM 1 Protein TRANSPIRATION EFFICIENCY 1 | ||||||||
| Gene names |
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| Organism | Arabidopsis thaliana (Mouse-ear cress) [Reference proteome] | ||||||||
| Taxonomic identifier | 3702 [NCBI] | ||||||||
| Taxonomic lineage | Eukaryota › Viridiplantae › Streptophyta › Embryophyta › Tracheophyta › Spermatophyta › Magnoliophyta › eudicotyledons › core eudicotyledons › rosids › malvids › Brassicales › Brassicaceae › Camelineae › Arabidopsis![]() |
Protein attributes
| Sequence length | 976 AA. |
| Sequence status | Complete. |
| Sequence processing | The displayed sequence is further processed into a mature form. |
| Protein existence | Evidence at protein level |
General annotation (Comments)
| Function | Receptor kinase that, together with ERL1 and ERL2, regulates aerial architecture, including inflorescence (e.g. shoot apical meristem-originating organ shape, elongation of the internode and pedicels, and adaxial-abaxial polarity), and stomatal patterning (e.g. density and clustering), probably by tuning cell division and expansion. Modulates plant transpiration efficiency by controlling stomatal density, leaf photosynthetic capacity, epidermal cell expansion, mesophyll cell proliferation and cell-cell contact. Probable major trait regulating canalization (maintenance of phenotype despite varying environment) in many aspect of the plant physiology (e.g. plant morphology, light-dependent leaves number, branch number, flowering time, phytate and mineral concentrations) by transducing microenvironmental variation into phenotypic differentiation (ecological amplifier). May maintain development integrity in heat stress conditions. Regulates cell wall composition and structure. Confers resistance to the pathogenic bacteria Ralstonia solanacearum and to the necrotrophic fungi Plectosphaerella cucumerina and Pythium irregulare, and required for callose deposition upon infection. Resistance to P.cucumerina seems cell wall-mediated. Ref.1 Ref.9 Ref.10 Ref.11 Ref.12 Ref.13 Ref.14 Ref.15 Ref.16 Ref.17 Ref.18 Ref.19 Ref.20 Ref.21 Ref.22 Ref.23 Ref.24 Ref.25 |
| Catalytic activity | ATP + a protein = ADP + a phosphoprotein. |
| Subcellular location | Membrane; Single-pass type I membrane protein Potential. |
| Tissue specificity | Mostly expressed in shoot apical meristems (SAM), organ primordia, flowers, siliques and young rosette leaves, and, to a lower extent, in stems and cauline leaves. Ref.1 Ref.8 |
| Developmental stage | Strongly expressed in organ primordia and immature organs but weakly in mature organs. Observed in SAM at low levels during the vegetative growth with an increase at the transition to the reproductive growth phase. At the reproductive stage, localized in the young developing flowers. Expressed in inflorescence meristem and is up-regulated during flower initiation and formation of flower organs. Also found in cells that differentiate into pedicels. Ref.8 Ref.13 |
| Polymorphism | The cultivar Landsberg erecta (cv. Ler) derives from cv. Landsberg (cv. La-0) in which ERECTA is mutated at Ile-750 (variant er). |
| Disruption phenotype | In er-104 and er-105, small curly leaves and compact inflorescence with short thick siliques, increased canalization of rosette leaf number during long days. Ref.1 Ref.22 |
| Sequence similarities | Belongs to the protein kinase superfamily. Ser/Thr protein kinase family. Contains 20 LRR (leucine-rich) repeats. Contains 1 protein kinase domain. |
Ontologies
Sequence annotation (Features)
| Feature key | Position(s) | Length | Description | Graphical view | Feature identifier | ||||
Molecule processing | |||||||||
|---|---|---|---|---|---|---|---|---|---|
| Signal peptide | 1 – 24 | 24 | Potential | ||||||
| Chain | 25 – 976 | 952 | LRR receptor-like serine/threonine-protein kinase ERECTA | PRO_0000389000 | |||||
Regions | |||||||||
| Topological domain | 25 – 580 | 556 | Extracellular Potential | ||||||
| Transmembrane | 581 – 601 | 21 | Helical; Potential | ||||||
| Topological domain | 602 – 976 | 375 | Cytoplasmic Potential | ||||||
| Repeat | 69 – 92 | 24 | LRR 1 | ||||||
| Repeat | 93 – 115 | 23 | LRR 2 | ||||||
| Repeat | 117 – 140 | 24 | LRR 3 | ||||||
| Repeat | 141 – 163 | 23 | LRR 4 | ||||||
| Repeat | 165 – 187 | 23 | LRR 5 | ||||||
| Repeat | 189 – 212 | 24 | LRR 6 | ||||||
| Repeat | 213 – 235 | 23 | LRR 7 | ||||||
| Repeat | 237 – 259 | 23 | LRR 8 | ||||||
| Repeat | 260 – 282 | 23 | LRR 9 | ||||||
| Repeat | 284 – 306 | 23 | LRR 10 | ||||||
| Repeat | 308 – 330 | 23 | LRR 11 | ||||||
| Repeat | 332 – 355 | 24 | LRR 12 | ||||||
| Repeat | 356 – 379 | 24 | LRR 13 | ||||||
| Repeat | 380 – 401 | 22 | LRR 14 | ||||||
| Repeat | 404 – 425 | 22 | LRR 15 | ||||||
| Repeat | 428 – 449 | 22 | LRR 16 | ||||||
| Repeat | 452 – 473 | 22 | LRR 17 | ||||||
| Repeat | 476 – 498 | 23 | LRR 18 | ||||||
| Repeat | 500 – 522 | 23 | LRR 19 | ||||||
| Repeat | 523 – 545 | 23 | LRR 20 | ||||||
| Domain | 648 – 918 | 271 | Protein kinase | ||||||
| Nucleotide binding | 654 – 662 | 9 | ATP By similarity | ||||||
Sites | |||||||||
| Active site | 773 | 1 | Proton acceptor By similarity | ||||||
| Binding site | 676 | 1 | ATP By similarity | ||||||
Amino acid modifications | |||||||||
| Glycosylation | 65 | 1 | N-linked (GlcNAc...) Potential | ||||||
| Glycosylation | 74 | 1 | N-linked (GlcNAc...) Potential | ||||||
| Glycosylation | 221 | 1 | N-linked (GlcNAc...) Potential | ||||||
| Glycosylation | 234 | 1 | N-linked (GlcNAc...) Potential | ||||||
| Glycosylation | 305 | 1 | N-linked (GlcNAc...) Potential | ||||||
| Glycosylation | 329 | 1 | N-linked (GlcNAc...) Potential | ||||||
| Glycosylation | 409 | 1 | N-linked (GlcNAc...) Potential | ||||||
| Glycosylation | 457 | 1 | N-linked (GlcNAc...) Potential | ||||||
| Glycosylation | 510 | 1 | N-linked (GlcNAc...) Potential | ||||||
| Glycosylation | 528 | 1 | N-linked (GlcNAc...) Potential | ||||||
| Glycosylation | 543 | 1 | N-linked (GlcNAc...) Potential | ||||||
Natural variations | |||||||||
| Natural variant | 750 | 1 | I → K Er-1 in strain: cv. Landsberg erecta; round leaves, compact inflorescence, blunt fruits, short and thick siliques and petioles, susceptibility to pathogens such as R.solanacearum, P.irregulare and P.cucumerina, abnormal cell-wall composition and increased canalization of rosette leaf number during long days. In er-101 and er-102, compact inflorescence with short siliques and pedicels. Ref.1 Ref.9 Ref.12 Ref.17 Ref.21 Ref.25 | ||||||
| Natural variant | 886 | 1 | V → M in strain: cv. Mt-0. Ref.7 | ||||||
Experimental info | |||||||||
| Mutagenesis | 282 | 1 | M → I in er-103; compact inflorescence with short siliques, but normal leaves. Ref.1 Ref.18 | ||||||
| Mutagenesis | 489 | 1 | G → D in er-117; compact inflorescence with short siliques and pedicels, and susceptibility to P.cucumerina. Ref.9 Ref.17 | ||||||
| Mutagenesis | 831 | 1 | D → N in er-114; compact inflorescence with short siliques and pedicels, and susceptibility to P.cucumerina. Ref.9 Ref.17 | ||||||
Sequences
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References
| « Hide 'large scale' references | |
| [1] | "The Arabidopsis ERECTA gene encodes a putative receptor protein kinase with extracellular leucine-rich repeats." Torii K.U., Mitsukawa N., Oosumi T., Matsuura Y., Yokoyama R., Whittier R.F., Komeda Y. Plant Cell 8:735-746(1996) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [GENOMIC DNA / MRNA], VARIANT ER-1 LYS-750, FUNCTION, DISRUPTION PHENOTYPE, TISSUE SPECIFICITY, MUTAGENESIS OF MET-282. Strain: cv. Columbia and cv. Landsberg erecta. |
| [2] | "Sequence and analysis of chromosome 2 of the plant Arabidopsis thaliana." 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. Venter J.C.Nature 402:761-768(1999) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]. Strain: cv. Columbia. |
| [3] | The Arabidopsis Information Resource (TAIR) Submitted (APR-2011) to the EMBL/GenBank/DDBJ databases Cited for: GENOME REANNOTATION. Strain: cv. Columbia. |
| [4] | "Empirical analysis of transcriptional activity in the Arabidopsis genome." Yamada K., Lim J., Dale J.M., Chen H., Shinn P., Palm C.J., Southwick A.M., Wu H.C., Kim C.J., Nguyen M., Pham P.K., Cheuk R.F., Karlin-Newmann G., Liu S.X., Lam B., Sakano H., Wu T., Yu G. Ecker J.R.Science 302:842-846(2003) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA]. Strain: cv. Columbia. |
| [5] | "Genome-wide cloning and sequence analysis of leucine-rich repeat receptor-like protein kinase genes in Arabidopsis thaliana." Gou X., He K., Yang H., Yuan T., Lin H., Clouse S.D., Li J. BMC Genomics 11:19-19(2010) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA]. Strain: cv. Columbia. |
| [6] | "Large-scale analysis of RIKEN Arabidopsis full-length (RAFL) cDNAs." 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. Shinozaki K.Submitted (MAR-2005) to the EMBL/GenBank/DDBJ databases Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] OF 536-976. Strain: cv. Columbia. |
| [7] | "The genetic architecture of shoot branching in Arabidopsis thaliana: a comparative assessment of candidate gene associations vs. quantitative trait locus mapping." Ehrenreich I.M., Stafford P.A., Purugganan M.D. Genetics 176:1223-1236(2007) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [GENOMIC DNA] OF 791-935, VARIANT MET-886. Strain: cv. Ag-0, cv. An-1, cv. Br-0, cv. C24, cv. Ct-1, cv. Cvi-1, cv. Edi-0, cv. Ga-0, cv. Kas-2, cv. Kin-0, cv. Landsberg erecta, cv. Ll-0, cv. Lz-0, cv. Mt-0, cv. Nd-1, cv. Nok-3, cv. Oy-0, cv. Se-0, cv. Sorbo, cv. Tsu-1, cv. Van-0, cv. Wa-1 and cv. Wassilewskija. |
| [8] | "The Arabidopsis ERECTA gene is expressed in the shoot apical meristem and organ primordia." Yokoyama R., Takahashi T., Kato A., Torii K.U., Komeda Y. Plant J. 15:301-310(1998) [PubMed] [Europe PMC] [Abstract] Cited for: TISSUE SPECIFICITY, DEVELOPMENTAL STAGE. |
| [9] | "Receptor serine/threonine protein kinases in signalling: analysis of the erecta receptor-like kinase of Arabidopsis thaliana." Lease K.A., Lau N.Y., Schuster R.A., Torii K.U., Walker J.C. New Phytol. 151:133-144(2001) Cited for: FUNCTION, CHARACTERIZATION OF VARIANT ER-1 LYS-750, MUTAGENESIS OF GLY-489 AND ASP-831. |
| [10] | "KNAT1 and ERECTA regulate inflorescence architecture in Arabidopsis." Douglas S.J., Chuck G., Dengler R.E., Pelecanda L., Riggs C.D. Plant Cell 14:547-558(2002) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION. |
| [11] | "Novel as1 and as2 defects in leaf adaxial-abaxial polarity reveal the requirement for ASYMMETRIC LEAVES1 and 2 and ERECTA functions in specifying leaf adaxial identity." Xu L., Xu Y., Dong A., Sun Y., Pi L., Xu Y., Huang H. Development 130:4097-4107(2003) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION. |
| [12] | "ERECTA, an LRR receptor-like kinase protein controlling development pleiotropically affects resistance to bacterial wilt." Godiard L., Sauviac L., Torii K.U., Grenon O., Mangin B., Grimsley N.H., Marco Y. Plant J. 36:353-365(2003) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION, VARIANT ER-1 LYS-750. |
| [13] | "Synergistic interaction of three ERECTA-family receptor-like kinases controls Arabidopsis organ growth and flower development by promoting cell proliferation." Shpak E.D., Berthiaume C.T., Hill E.J., Torii K.U. Development 131:1491-1501(2004) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION, DEVELOPMENTAL STAGE. |
| [14] | "ERECTA is required for protection against heat-stress in the AS1/ AS2 pathway to regulate adaxial-abaxial leaf polarity in Arabidopsis." Qi Y., Sun Y., Xu L., Xu Y., Huang H. Planta 219:270-276(2004) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION. |
| [15] | "Pedicel development in Arabidopsis thaliana: contribution of vascular positioning and the role of the BREVIPEDICELLUS and ERECTA genes." Douglas S.J., Riggs C.D. Dev. Biol. 284:451-463(2005) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION. |
| [16] | "The ERECTA gene regulates plant transpiration efficiency in Arabidopsis." Masle J., Gilmore S.R., Farquhar G.D. Nature 436:866-870(2005) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION. |
| [17] | "ERECTA receptor-like kinase and heterotrimeric G protein from Arabidopsis are required for resistance to the necrotrophic fungus Plectosphaerella cucumerina." Llorente F., Alonso-Blanco C., Sanchez-Rodriguez C., Jorda L., Molina A. Plant J. 43:165-180(2005) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION, MUTAGENESIS OF GLY-489 AND ASP-831, VARIANT ER-1 LYS-750. |
| [18] | "Interaction of auxin and ERECTA in elaborating Arabidopsis inflorescence architecture revealed by the activation tagging of a new member of the YUCCA family putative flavin monooxygenases." Woodward C., Bemis S.M., Hill E.J., Sawa S., Koshiba T., Torii K.U. Plant Physiol. 139:192-203(2005) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION, MUTAGENESIS OF MET-282. |
| [19] | "Stomatal patterning and differentiation by synergistic interactions of receptor kinases." Shpak E.D., McAbee J.M., Pillitteri L.J., Torii K.U. Science 309:290-293(2005) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION. |
| [20] | "Haploinsufficiency after successive loss of signaling reveals a role for ERECTA-family genes in Arabidopsis ovule development." Pillitteri L.J., Bemis S.M., Shpak E.D., Torii K.U. Development 134:3099-3109(2007) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION. |
| [21] | "ABA is an essential signal for plant resistance to pathogens affecting JA biosynthesis and the activation of defenses in Arabidopsis." Adie B.A.T., Perez-Perez J., Perez-Perez M.M., Godoy M., Sanchez-Serrano J.-J., Schmelz E.A., Solano R. Plant Cell 19:1665-1681(2007) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION, VARIANT ER-1 LYS-750. |
| [22] | "Genetics of microenvironmental canalization in Arabidopsis thaliana." Hall M.C., Dworkin I., Ungerer M.C., Purugganan M. Proc. Natl. Acad. Sci. U.S.A. 104:13717-13722(2007) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION, DISRUPTION PHENOTYPE. |
| [23] | "Polymorphic genes of major effect: consequences for variation, selection and evolution in Arabidopsis thaliana." Stinchcombe J.R., Weinig C., Heath K.D., Brock M.T., Schmitt J. Genetics 182:911-922(2009) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION. |
| [24] | "A strong effect of growth medium and organ type on the identification of QTLs for phytate and mineral concentrations in three Arabidopsis thaliana RIL populations." Ghandilyan A., Ilk N., Hanhart C., Mbengue M., Barboza L., Schat H., Koornneef M., El-Lithy M., Vreugdenhil D., Reymond M., Aarts M.G.M. J. Exp. Bot. 60:1409-1425(2009) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION. |
| [25] | "The ERECTA receptor-like kinase regulates cell wall-mediated resistance to pathogens in Arabidopsis thaliana." Sanchez-Rodriguez C., Estevez J.M., Llorente F., Hernandez-Blanco C., Jorda L., Pagan I., Berrocal M., Marco Y., Somerville S., Molina A. Mol. Plant Microbe Interact. 22:953-963(2009) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION, VARIANT ER-1 LYS-750. |
| [26] | "The many functions of ERECTA." van Zanten M., Snoek L.B., Proveniers M.C.G., Peeters A.J.M. Trends Plant Sci. 14:214-218(2009) [PubMed] [Europe PMC] [Abstract] Cited for: REVIEW. |
| + | Additional computationally mapped references. |
Web resources
| PlantP kinase Classification PPC |
| Protein Spotlight A complicated affair - Issue 116 of April 2010 |
Cross-references
Sequence databases | |
|---|---|
| EMBL GenBank DDBJ | D83257 Genomic DNA. Translation: BAA11869.1. U47029 mRNA. Translation: AAC49302.1. AC004484 Genomic DNA. Translation: AAC14518.1. CP002685 Genomic DNA. Translation: AEC07825.1. AY035110 mRNA. Translation: AAK59615.1. FJ708701 mRNA. Translation: ACN59296.1. AK221886 mRNA. Translation: BAD94220.1. EF598332 Genomic DNA. Translation: ABR08864.1. EF598333 Genomic DNA. Translation: ABR08865.1. EF598334 Genomic DNA. Translation: ABR08866.1. EF598335 Genomic DNA. Translation: ABR08867.1. EF598336 Genomic DNA. Translation: ABR08868.1. EF598337 Genomic DNA. Translation: ABR08869.1. EF598338 Genomic DNA. Translation: ABR08870.1. EF598339 Genomic DNA. Translation: ABR08871.1. EF598340 Genomic DNA. Translation: ABR08872.1. EF598341 Genomic DNA. Translation: ABR08873.1. EF598342 Genomic DNA. Translation: ABR08874.1. EF598343 Genomic DNA. Translation: ABR08875.1. EF598344 Genomic DNA. Translation: ABR08876.1. EF598345 Genomic DNA. Translation: ABR08877.1. EF598346 Genomic DNA. Translation: ABR08878.1. EF598347 Genomic DNA. Translation: ABR08879.1. EF598348 Genomic DNA. Translation: ABR08880.1. EF598349 Genomic DNA. Translation: ABR08881.1. EF598350 Genomic DNA. Translation: ABR08882.1. EF598351 Genomic DNA. Translation: ABR08883.1. EF598352 Genomic DNA. Translation: ABR08884.1. EF598353 Genomic DNA. Translation: ABR08885.1. EF598354 Genomic DNA. Translation: ABR08886.1. |
| IPI | IPI00532140. |
| PIR | B84659. |
| RefSeq | NP_180201.1. NM_128190.2. |
| UniGene | At.10804. |
3D structure databases | |
| HSSP | HSSP built from PDB template 1OGQ based on UniProtKB P58822. |
| ProteinModelPortal | Q42371. |
| SMR | Q42371. Positions 36-560, 645-947. |
| ModBase | Search... |
Protein-protein interaction databases | |
| STRING | 3702.AT2G26330.1-P. |
Proteomic databases | |
| PaxDb | Q42371. |
| PRIDE | Q42371. |
Protocols and materials databases | |
| StructuralBiologyKnowledgebase | Search... |
Genome annotation databases | |
| EnsemblPlants | AT2G26330.1; AT2G26330.1; AT2G26330. |
| GeneID | 817173. |
| KEGG | ath:AT2G26330. |
Organism-specific databases | |
| GeneFarm | 2563. 221. |
| TAIR | At2g26330. |
Phylogenomic databases | |
| eggNOG | COG0515. |
| HOGENOM | HOG000116551. |
| InParanoid | Q42371. |
| OMA | PSSDYCV. |
| PhylomeDB | Q42371. |
| ProtClustDB | CLSN2687327. |
Family and domain databases | |
| InterPro | IPR011009. Kinase-like_dom. IPR001611. Leu-rich_rpt. IPR013210. LRR-contain_N2. IPR000719. Prot_kinase_cat_dom. IPR017441. Protein_kinase_ATP_BS. IPR008271. Ser/Thr_kinase_AS. [Graphical view] |
| Pfam | PF00560. LRR_1. 5 hits. PF08263. LRRNT_2. 1 hit. PF00069. Pkinase. 1 hit. [Graphical view] |
| SUPFAM | SSF56112. Kinase_like. 1 hit. |
| PROSITE | PS51450. LRR. 17 hits. PS00107. PROTEIN_KINASE_ATP. 1 hit. PS50011. PROTEIN_KINASE_DOM. 1 hit. PS00108. PROTEIN_KINASE_ST. 1 hit. [Graphical view] |
| ProtoNet | Search... |
Entry information
| Entry name | ERECT_ARATH | ||||||||
| Accession | Primary (citable) accession number: Q42371 Secondary accession number(s): A5YYA0, A5YYB1, Q56WZ3 | ||||||||
| Entry history |
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| Entry status | Reviewed (UniProtKB/Swiss-Prot) | ||||||||
| Annotation program | Plant Protein Annotation Program | ||||||||
Relevant documents
| Arabidopsis thaliana Arabidopsis thaliana: entries and gene names |
| SIMILARITY comments Index of protein domains and families |
| Protein Spotlight Protein Spotlight articles and cited UniProtKB/Swiss-Prot entries |

Clusters with
