Q38890 (GUN25_ARATH) Reviewed, UniProtKB/Swiss-Prot

Names and origin

Protein names

Recommended name:
Endoglucanase 25
EC=3.2.1.4

Alternative name(s):
Cellulase homolog OR16pep
Endo-1,4-beta glucanase 25
Protein KORRIGAN
Protein RADIALLY SWOLLEN 2

Gene names

Name:	KOR
Synonyms:	DEC, KOR1, RSW2
Ordered Locus Names:	At5g49720
ORF Names:	K2I5.8

Organism

Arabidopsis thaliana (Mouse-ear cress) [Reference proteome]

Taxonomic identifier

3702 [NCBI]

Taxonomic lineage

cellular organisms › Eukaryota › Viridiplantae › Streptophyta › Streptophytina › Embryophyta › Tracheophyta › Euphyllophyta › Spermatophyta › Magnoliophyta › eudicotyledons › core eudicotyledons › rosids › malvids › Brassicales › Brassicaceae › Camelineae › Arabidopsis

Protein attributes

Sequence length	621 AA.
Sequence status	Complete.
Protein existence	Evidence at protein level

General annotation (Comments)

Function	Required for cellulose microfibrils formation. Involved in cell wall assembly during cell elongation and cell plate maturation in cytokinesis. Required for secondary cell wall formation in the developing xylem. May cycle through different intracellular compartments, including plasma membrane. Ref.2 Ref.3 Ref.9 Ref.13 Ref.14
Catalytic activity	Endohydrolysis of (1->4)-beta-D-glucosidic linkages in cellulose, lichenin and cereal beta-D-glucans.
Subcellular location	Cell membrane; Single-pass type II membrane protein. Note: Cell plate. Ref.2
Tissue specificity	Highly expressed in roots and stems, at intermediate levels in leaves and flowers, and at lower levels in siliques. Expressed in xylem (at protein level). Ref.2 Ref.3 Ref.13
Developmental stage	Expressed during hypocotyl elongation in the dark. Ref.2
Post-translational modification	Glycosylated. N-glycosylation of KOR in the endoplasmic reticulum followed by N-glycan modifications in the Golgi are essential for catalytic activity. Ref.15
Disruption phenotype	Plants are extremely dwarf and show severe abnormal morphology with incomplete cell walls, aberrant cell plates and multinucleated cells. Ref.2
Sequence similarities	Belongs to the glycosyl hydrolase 9 (cellulase E) family.

Ontologies

Keywords
Biological process	Carbohydrate metabolism Cell wall biogenesis/degradation Cellulose degradation Polysaccharide degradation
Cellular component	Cell membrane Membrane
Domain	Signal-anchor Transmembrane Transmembrane helix
Molecular function	Glycosidase Hydrolase
PTM	Glycoprotein Phosphoprotein
Technical term	Complete proteome Reference proteome
Gene Ontology (GO)
Biological_process	cellulose biosynthetic process Inferred from mutant phenotype PubMed 18583534. Source: TAIR cellulose catabolic process Inferred from electronic annotation. Source: UniProtKB-KW cortical microtubule organization Inferred from mutant phenotype PubMed 18583534. Source: TAIR hyperosmotic salinity response Inferred from mutant phenotype Ref.15. Source: TAIR response to cytokinin stimulus Inferred from mutant phenotype PubMed 12060228. Source: TAIR shoot system development Inferred from mutant phenotype PubMed 12060228. Source: TAIR unidimensional cell growth Inferred from mutant phenotype PubMed 10987560. Source: TAIR
Cellular_component	Golgi apparatus Inferred from direct assay Ref.14PubMed 22430844PubMed 22923678. Source: TAIR cell plate Inferred from direct assay Ref.3. Source: TAIR early endosome Inferred from direct assay Ref.14. Source: TAIR integral to membrane Inferred from electronic annotation. Source: UniProtKB-KW plasma membrane Inferred from direct assay Ref.10PubMed 17317660. Source: TAIR trans-Golgi network Inferred from direct assay PubMed 22923678. Source: TAIR
Molecular_function	cellulase activity Inferred from sequence or structural similarity Ref.2. Source: TAIR
Complete GO annotation...

Sequence annotation (Features)

	Feature key	Position(s)	Length	Description	Graphical view	Feature identifier
Molecule processing
	Chain	1 – 621	621	Endoglucanase 25		PRO_0000249277
Regions
	Topological domain	1 – 70	70	Cytoplasmic Potential
	Transmembrane	71 – 91	21	Helical; Signal-anchor for type II membrane protein; Potential
	Topological domain	92 – 621	530	Extracellular Potential
	Region	48 – 49	2	Polarized targeting signal 1 (PTS1)
	Region	59 – 62	4	Polarized targeting signal 2 (PTS2)
Sites
	Active site	513	1	By similarity
	Active site	561	1	By similarity
	Active site	570	1	By similarity
Amino acid modifications
	Modified residue	18	1	Phosphoserine Ref.10
	Modified residue	20	1	Phosphothreonine Ref.10 Ref.12
	Modified residue	25	1	Phosphoserine Ref.10
	Glycosylation	108	1	N-linked (GlcNAc...) Potential
	Glycosylation	133	1	N-linked (GlcNAc...) Potential
	Glycosylation	216	1	N-linked (GlcNAc...) Potential
	Glycosylation	324	1	N-linked (GlcNAc...) Potential
	Glycosylation	345	1	N-linked (GlcNAc...) Potential
	Glycosylation	408	1	N-linked (GlcNAc...) Potential
	Glycosylation	425	1	N-linked (GlcNAc...) Potential
	Glycosylation	567	1	N-linked (GlcNAc...) Potential
Experimental info
	Mutagenesis	48 – 49	2	LL → AA: Abolishes polarized targeting to cell plate.
	Mutagenesis	59	1	Y → A: Abolishes polarized targeting to cell plate. Ref.3
	Mutagenesis	183	1	S → N in rsw2-3; decrease in cellulose production. Ref.9
	Mutagenesis	250	1	P → L in irx2-1; decrease in cellulose production. Ref.13
	Mutagenesis	344	1	G → R in rsw2-4; decrease in cellulose production. Ref.9
	Mutagenesis	429	1	G → R in rsw2-1 and rsw2-2; decrease in cellulose production. Ref.9
	Mutagenesis	553	1	P → L in irx2-2; decrease in cellulose production. Ref.13
	Sequence conflict	7	1	W → S in AAM63370. Ref.7
	Sequence conflict	59	1	Y → D in AAK59818. Ref.6
	Sequence conflict	338	1	W → R in AAM63370. Ref.7
	Sequence conflict	397	1	Y → C in AAK59818. Ref.6
	Sequence conflict	397	1	Y → C in AAN72232. Ref.6
	Sequence conflict	605	1	V → A in BAD94393. Ref.8

Sequences

	Sequence		Length	Mass (Da)
	Q38890 [UniParc]. Last modified November 1, 1996. Version 1. Checksum: 798D8B8CC5DEBF42	FASTA	621	69,191
	10 20 30 40 50 60 MYGRDPWGGP LEINTADSAT DDDRSRNLND LDRAALSRPL DETQQSWLLG PTEQKKKKYV 70 80 90 100 110 120 DLGCIIVSRK IFVWTVGTLV AAALLAGFIT LIVKTVPRHH PKTPPPDNYT IALHKALKFF 130 140 150 160 170 180 NAQKSGKLPK HNNVSWRGNS GLQDGKGETG SFYKDLVGGY YDAGDAIKFN FPMAYAMTML 190 200 210 220 230 240 SWSVIEYSAK YEAAGELTHV KELIKWGTDY FLKTFNSTAD SIDDLVSQVG SGNTDDGNTD 250 260 270 280 290 300 PNDHYCWMRP EDMDYKRPVT TCNGGCSDLA AEMAAALASA SIVFKDNKEY SKKLVHGAKV 310 320 330 340 350 360 VYQFGRTRRG RYSAGTAESS KFYNSSMYWD EFIWGGAWMY YATGNVTYLN LITQPTMAKH 370 380 390 400 410 420 AGAFWGGPYY GVFSWDNKLA GAQLLLSRLR LFLSPGYPYE EILRTFHNQT SIVMCSYLPI 430 440 450 460 470 480 FNKFNRTNGG LIELNHGAPQ PLQYSVNAAF LATLYSDYLD AADTPGWYCG PNFYSTSVLR 490 500 510 520 530 540 DFARSQIDYI LGKNPRKMSY VVGFGTKYPR HVHHRGASIP KNKVKYNCKG GWKWRDSKKP 550 560 570 580 590 600 NPNTIEGAMV AGPDKRDGYR DVRMNYNYTE PTLAGNAGLV AALVALSGEE EATGKIDKNT 610 620 IFSAVPPLFP TPPPPPAPWK P

References

	« Hide 'large scale' referencesShow 'large scale' references »
[1]	"An Arabidopsis thaliana cDNA homologous to cellulase." Loebler M. DNA Seq. 8:253-256(1998) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA]. Strain: cv. Columbia. Tissue: Leaf.
[2]	"A plasma membrane-bound putative endo-1,4-beta-D-glucanase is required for normal wall assembly and cell elongation in Arabidopsis." Nicol F., His I., Jauneau A., Vernhettes S., Canut H., Hoefte H. EMBO J. 17:5563-5576(1998) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [GENOMIC DNA], FUNCTION, SUBCELLULAR LOCATION, TISSUE SPECIFICITY, DEVELOPMENTAL STAGE, DISRUPTION PHENOTYPE. Strain: cv. Columbia.
[3]	"KORRIGAN, an Arabidopsis endo-1,4-beta-glucanase, localizes to the cell plate by polarized targeting and is essential for cytokinesis." Zuo J., Niu Q.-W., Nishizawa N., Wu Y., Kost B., Chua N.-H. Plant Cell 12:1137-1152(2000) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [GENOMIC DNA / MRNA], FUNCTION, TISSUE SPECIFICITY, MUTAGENESIS OF 48-LYS-LYS-49 AND TYR-59. Strain: cv. Landsberg erecta.
[4]	"Structural analysis of Arabidopsis thaliana chromosome 5. XI." Kaneko T., Katoh T., Asamizu E., Sato S., Nakamura Y., Kotani H., Tabata S. Submitted (APR-1999) to the EMBL/GenBank/DDBJ databases Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]. Strain: cv. Columbia.
[5]	The Arabidopsis Information Resource (TAIR) Submitted (APR-2011) to the EMBL/GenBank/DDBJ databases Cited for: GENOME REANNOTATION. Strain: cv. Columbia.
[6]	"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. , Miranda M., Quach H.L., Tripp M., Chang C.H., Lee J.M., Toriumi M.J., Chan M.M., Tang C.C., Onodera C.S., Deng J.M., Akiyama K., Ansari Y., Arakawa T., Banh J., Banno F., Bowser L., Brooks S.Y., Carninci P., Chao Q., Choy N., Enju A., Goldsmith A.D., Gurjal M., Hansen N.F., Hayashizaki Y., Johnson-Hopson C., Hsuan V.W., Iida K., Karnes M., Khan S., Koesema E., Ishida J., Jiang P.X., Jones T., Kawai J., Kamiya A., Meyers C., Nakajima M., Narusaka M., Seki M., Sakurai T., Satou M., Tamse R., Vaysberg M., Wallender E.K., Wong C., Yamamura Y., Yuan S., Shinozaki K., Davis R.W., Theologis A., Ecker J.R. Science 302:842-846(2003) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA]. Strain: cv. Columbia.
[7]	"Full-length cDNA from Arabidopsis thaliana." Brover V.V., Troukhan M.E., Alexandrov N.A., Lu Y.-P., Flavell R.B., Feldmann K.A. Submitted (MAR-2002) to the EMBL/GenBank/DDBJ databases Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
[8]	"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. , Satou M., Toyoda T., Konagaya A., Carninci P., Kawai J., Hayashizaki Y., Shinozaki K. Submitted (MAR-2005) to the EMBL/GenBank/DDBJ databases Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] OF 366-621. Strain: cv. Columbia.
[9]	"Temperature-sensitive alleles of RSW2 link the KORRIGAN endo-1,4-beta-glucanase to cellulose synthesis and cytokinesis in Arabidopsis." Lane D.R., Wiedemeier A., Peng L., Hoefte H., Vernhettes S., Desprez T., Hocart C.H., Birch R.J., Baskin T.I., Burn J.E., Arioli T., Betzner A.S., Williamson R.E. Plant Physiol. 126:278-288(2001) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION, MUTAGENESIS OF SER-183; GLY-344 AND GLY-429.
[10]	"Large-scale analysis of in vivo phosphorylated membrane proteins by immobilized metal ion affinity chromatography and mass spectrometry." Nuehse T.S., Stensballe A., Jensen O.N., Peck S.C. Mol. Cell. Proteomics 2:1234-1243(2003) [PubMed] [Europe PMC] [Abstract] Cited for: PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT SER-18; THR-20 AND SER-25, MASS SPECTROMETRY. Strain: cv. La-0.
[11]	"Phylogenetic analysis of the plant endo-beta-1,4-glucanase gene family." Libertini E., Li Y., McQueen-Mason S.J. J. Mol. Evol. 58:506-515(2004) [PubMed] [Europe PMC] [Abstract] Cited for: GENE FAMILY.
[12]	"Phosphoproteomics of the Arabidopsis plasma membrane and a new phosphorylation site database." Nuehse T.S., Stensballe A., Jensen O.N., Peck S.C. Plant Cell 16:2394-2405(2004) [PubMed] [Europe PMC] [Abstract] Cited for: PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT THR-20, MASS SPECTROMETRY.
[13]	"The irregular xylem 2 mutant is an allele of korrigan that affects the secondary cell wall of Arabidopsis thaliana." Szyjanowicz P.M., McKinnon I., Taylor N.G., Gardiner J., Jarvis M.C., Turner S.R. Plant J. 37:730-740(2004) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION, TISSUE SPECIFICITY, MUTAGENESIS OF PRO-250 AND PRO-553.
[14]	"An Arabidopsis endo-1,4-beta-D-glucanase involved in cellulose synthesis undergoes regulated intracellular cycling." Robert S., Bichet A., Grandjean O., Kierzkowski D., Satiat-Jeunemaitre B., Pelletier S., Hauser M.-T., Hoefte H., Vernhettes S. Plant Cell 17:3378-3389(2005) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION.
[15]	"Salt tolerance of Arabidopsis thaliana requires maturation of N-glycosylated proteins in the Golgi apparatus." Kang J.S., Frank J., Kang C.H., Kajiura H., Vikram M., Ueda A., Kim S., Bahk J.D., Triplett B., Fujiyama K., Lee S.Y., von Schaewen A., Koiwa H. Proc. Natl. Acad. Sci. U.S.A. 105:5933-5938(2008) [PubMed] [Europe PMC] [Abstract] Cited for: GLYCOSYLATION.
+	Additional computationally mapped references.

Cross-references

Sequence databases
EMBL GenBank DDBJ	U37702 mRNA. Translation: AAB60304.1. AF073875 Genomic DNA. Translation: AAC83240.1. AF074092 Genomic DNA. Translation: AAC33467.1. AF074375 mRNA. Translation: AAC35344.1. AB025613 Genomic DNA. Translation: BAA98160.1. CP002688 Genomic DNA. Translation: AED95850.1. AY037218 mRNA. Translation: AAK59818.1. BT002221 mRNA. Translation: AAN72232.1. AY086165 mRNA. Translation: AAM63370.1. AK221941 mRNA. Translation: BAD94393.1. AK222193 mRNA. Translation: BAD95336.1.
IPI	IPI00523242.
PIR	S71215.
RefSeq	NP_199783.1. NM_124350.2.
UniGene	At.21098. At.23416.
3D structure databases
ProteinModelPortal	Q38890.
SMR	Q38890. Positions 105-618.
ModBase	Search...
Protein family/group databases
CAZy	GH9. Glycoside Hydrolase Family 9.
Proteomic databases
PaxDb	Q38890.
PRIDE	Q38890.
Protocols and materials databases
StructuralBiologyKnowledgebase	Search...
Genome annotation databases
EnsemblPlants	AT5G49720.1; AT5G49720.1; AT5G49720.
GeneID	835035.
KEGG	ath:AT5G49720.
Organism-specific databases
TAIR	At5g49720.
Phylogenomic databases
eggNOG	NOG257354.
HOGENOM	HOG000021033.
InParanoid	Q38890.
OMA	MMIVKLV.
PhylomeDB	Q38890.
ProtClustDB	CLSN2679649.
Enzyme and pathway databases
BioCyc	MetaCyc:MONOMER-2367.
Gene expression databases
ArrayExpress	Q38890.
Genevestigator	Q38890.
GermOnline	AT5G49720. Arabidopsis thaliana.
Family and domain databases
Gene3D	1.50.10.10. 1 hit.
InterPro	IPR008928. 6-hairpin_glycosidase-like. IPR012341. 6hp_glycosidase. IPR001701. Glyco_hydro_9. IPR018221. Glyco_hydro_9_AS. [Graphical view]
Pfam	PF00759. Glyco_hydro_9. 1 hit. [Graphical view]
SUPFAM	SSF48208. Glyco_trans_6hp. 1 hit.
PROSITE	PS00592. GLYCOSYL_HYDROL_F9_1. 1 hit. PS00698. GLYCOSYL_HYDROL_F9_2. 1 hit. [Graphical view]
ProtoNet	Search...

Entry information

Entry name

GUN25_ARATH

Accession

Primary (citable) accession number: Q38890
Secondary accession number(s): Q56W54 , Q56WU0, Q8H0S4, Q8LD74, Q94C24

Entry history

Integrated into UniProtKB/Swiss-Prot:	September 5, 2006
Last sequence update:	November 1, 1996
Last modified:	May 29, 2013
This is version 102 of the entry and version 1 of the sequence. [Complete history]

Entry status

Reviewed (UniProtKB/Swiss-Prot)

Annotation program

Plant Protein Annotation Program