P16371 (GROU_DROME) Reviewed, UniProtKB/Swiss-Prot

Names and origin

Protein names

Recommended name:
Protein groucho

Alternative name(s):
Enhancer of split m9/10 protein
Short name=E(spl)m9/10

Gene names

Name:	gro
Synonyms:	E(spl)m9/m10
ORF Names:	CG8384

Organism

Drosophila melanogaster (Fruit fly) [Reference proteome]

Taxonomic identifier

7227 [NCBI]

Taxonomic lineage

cellular organisms › Eukaryota › Opisthokonta › Metazoa › Eumetazoa › Bilateria › Protostomia › Ecdysozoa › Panarthropoda › Arthropoda › Mandibulata › Pancrustacea › Hexapoda › Insecta › Dicondylia › Pterygota › Neoptera › Endopterygota › Diptera › Brachycera › Muscomorpha › Eremoneura › Cyclorrhapha › Schizophora › Acalyptratae › Ephydroidea › Drosophilidae › Drosophilinae › Drosophilini › Drosophilina › Drosophiliti › Drosophila › Sophophora › melanogaster group › melanogaster subgroup

Protein attributes

Sequence length	730 AA.
Sequence status	Complete.
Protein existence	Evidence at protein level

General annotation (Comments)

Function	Transcriptional corepressor that regulates transcription when recruited to specific target DNA by hairy-related bHLH proteins. Maternally required for neurogenesis; in the segregation of the neuroectoderm. Directly or indirectly interacts with Notch and Delta. Ref.5
Subunit structure	Forms a complex with the hairy/Enhancer of split/deadpan family of basic helix-loop-helix proteins in order to repress transcription. Its activity in regulating transcription depends on other proteins as it lacks a DNA-binding motif.
Subcellular location	Nucleus.
Developmental stage	Expressed both maternally and zygotically. Ref.5
Post-translational modification	Ubiquitinated by XIAP/BIRC4. Ref.7
Sequence similarities	Belongs to the WD repeat Groucho/TLE family. Contains 7 WD repeats.

Ontologies

Keywords
Biological process	Differentiation Neurogenesis Transcription Transcription regulation Wnt signaling pathway
Cellular component	Nucleus
Coding sequence diversity	Alternative splicing
Domain	Repeat WD repeat
Molecular function	Developmental protein Repressor
PTM	Phosphoprotein Ubl conjugation
Technical term	Complete proteome Reference proteome
Gene Ontology (GO)
Biological_process	Wnt receptor signaling pathway Inferred from electronic annotation. Source: UniProtKB-KW actin filament organization Inferred from mutant phenotype PubMed 14527345. Source: FlyBase dendrite morphogenesis Inferred from mutant phenotype PubMed 16547170. Source: FlyBase lateral inhibition Inferred from mutant phenotype PubMed 19363474. Source: FlyBase negative regulation of canonical Wnt receptor signaling pathway Inferred from direct assay PubMed 19460168. Source: UniProtKB negative regulation of smoothened signaling pathway Inferred from mutant phenotype PubMed 17571073. Source: FlyBase sex determination, primary response to X:A ratio Traceable author statement PubMed 10648226. Source: FlyBase
Cellular_component	nucleus Traceable author statement PubMed 10929403. Source: FlyBase protein complex Inferred from physical interaction PubMed 17898168. Source: FlyBase
Molecular_function	RNA polymerase II transcription corepressor activity Inferred from genetic interaction PubMed 12466194. Source: FlyBase repressing transcription factor binding Inferred from genetic interaction PubMed 12466194. Source: FlyBase
Complete GO annotation...

Binary interactions

With	Entry	#Exp.	IntAct	Notes
E(spl)	P13098	4	EBI-153866,EBI-185388
HLHm3	Q01068	4	EBI-153866,EBI-121622
HLHm5	P13096	3	EBI-153866,EBI-104760
HLHmdelta	Q01071	3	EBI-153866,EBI-118907
pan	P91943	2	EBI-153866,EBI-147301
RUNX1	Q01196-1	4	EBI-153866,EBI-925940	From a different organism.
Six3	Q62233	2	EBI-153866,EBI-2297327	From a different organism.

Alternative products

This entry describes 2 isoforms produced by alternative splicing. [Align] [Select]

Isoform E (identifier: P16371-1)

This isoform has been chosen as the 'canonical' sequence. All positional information in this entry refers to it. This is also the sequence that appears in the downloadable versions of the entry.

Note: No experimental confirmation available.

Isoform A (identifier: P16371-2)

Also known as: B;

The sequence of this isoform differs from the canonical sequence as follows:
     127-137: Missing.

Sequence annotation (Features)

	Feature key	Position(s)	Length	Description	Graphical view	Feature identifier
Molecule processing
	Chain	1 – 730	730	Protein groucho		PRO_0000051010
Regions
	Repeat	442 – 480	39	WD 1
	Repeat	488 – 527	40	WD 2
	Repeat	532 – 571	40	WD 3
	Repeat	574 – 613	40	WD 4
	Repeat	615 – 654	40	WD 5
	Repeat	656 – 695	40	WD 6
	Repeat	697 – 730	34	WD 7
	Region	206 – 267	62	CCN domain
	Region	262 – 425	164	Binding to basic helix-loop-helix domain
	Motif	227 – 230	4	Nuclear localization signal Potential
	Compositional bias	93 – 144	52	Gln-rich
	Compositional bias	317 – 423	107	Pro-rich
Amino acid modifications
	Modified residue	205	1	Phosphoserine Ref.6
	Modified residue	207	1	Phosphoserine Ref.6
	Modified residue	218	1	Phosphoserine Ref.6
	Modified residue	242	1	Phosphoserine; by CK2 Potential
	Modified residue	258	1	Phosphoserine; by CDC2 Potential
	Modified residue	267	1	Phosphoserine Ref.6
	Modified residue	326	1	Phosphothreonine Ref.6
	Modified residue	328	1	Phosphothreonine Ref.6
Natural variations
	Alternative sequence	127 – 137	11	Missing in isoform A.		VSP_022308
Experimental info
	Sequence conflict	41	1	Q → H in AAA28512. Ref.1

Sequences

	Sequence		Length	Mass (Da)
	Isoform E [UniParc]. Last modified January 9, 2007. Version 3. Checksum: 9DA08E1ED343B2F5	FASTA	730	80,231
	10 20 30 40 50 60 MYPSPVRHPA AGGPPPQGPI KFTIADTLER IKEEFNFLQA QYHSIKLECE KLSNEKTEMQ 70 80 90 100 110 120 RHYVMYYEMS YGLNVEMHKQ TEIAKRLNTL INQLLPFLQA DHQQQVLQAV ERAKQVTMQE 130 140 150 160 170 180 LNLIIGHQQQ HGIQQLLQQI HAQQVPGGPP QPMGALNPFG ALGATMGLPH GPQGLLNKPP 190 200 210 220 230 240 EHHRPDIKPT GLEGPAAAEE RLRNSVSPAD REKYRTRSPL DIENDSKRRK DEKLQEDEGE 250 260 270 280 290 300 KSDQDLVVDV ANEMESHSPR PNGEHVSMEV RDRESLNGER LEKPSSSGIK QERPPSRSGS 310 320 330 340 350 360 SSSRSTPSLK TKDMEKPGTP GAKARTPTPN AAAPAPGVNP KQMMPQGPPP AGYPGAPYQR 370 380 390 400 410 420 PADPYQRPPS DPAYGRPPPM PYDPHAHVRT NGIPHPSALT GGKPAYSFHM NGEGSLQPVP 430 440 450 460 470 480 FPPDALVGVG IPRHARQINT LSHGEVVCAV TISNPTKYVY TGGKGCVKVW DISQPGNKNP 490 500 510 520 530 540 VSQLDCLQRD NYIRSVKLLP DGRTLIVGGE ASNLSIWDLA SPTPRIKAEL TSAAPACYAL 550 560 570 580 590 600 AISPDSKVCF SCCSDGNIAV WDLHNEILVR QFQGHTDGAS CIDISPDGSR LWTGGLDNTV 610 620 630 640 650 660 RSWDLREGRQ LQQHDFSSQI FSLGYCPTGD WLAVGMENSH VEVLHASKPD KYQLHLHESC 670 680 690 700 710 720 VLSLRFAACG KWFVSTGKDN LLNAWRTPYG ASIFQSKETS SVLSCDISTD DKYIVTGSGD 730 KKATVYEVIY
	Isoform A (B) [UniParc]. Checksum: D495291FD77026A5	FASTA	719	78,920
	10 20 30 40 50 60 MYPSPVRHPA AGGPPPQGPI KFTIADTLER IKEEFNFLQA QYHSIKLECE KLSNEKTEMQ 70 80 90 100 110 120 RHYVMYYEMS YGLNVEMHKQ TEIAKRLNTL INQLLPFLQA DHQQQVLQAV ERAKQVTMQE 130 140 150 160 170 180 LNLIIGQQIH AQQVPGGPPQ PMGALNPFGA LGATMGLPHG PQGLLNKPPE HHRPDIKPTG 190 200 210 220 230 240 LEGPAAAEER LRNSVSPADR EKYRTRSPLD IENDSKRRKD EKLQEDEGEK SDQDLVVDVA 250 260 270 280 290 300 NEMESHSPRP NGEHVSMEVR DRESLNGERL EKPSSSGIKQ ERPPSRSGSS SSRSTPSLKT 310 320 330 340 350 360 KDMEKPGTPG AKARTPTPNA AAPAPGVNPK QMMPQGPPPA GYPGAPYQRP ADPYQRPPSD 370 380 390 400 410 420 PAYGRPPPMP YDPHAHVRTN GIPHPSALTG GKPAYSFHMN GEGSLQPVPF PPDALVGVGI 430 440 450 460 470 480 PRHARQINTL SHGEVVCAVT ISNPTKYVYT GGKGCVKVWD ISQPGNKNPV SQLDCLQRDN 490 500 510 520 530 540 YIRSVKLLPD GRTLIVGGEA SNLSIWDLAS PTPRIKAELT SAAPACYALA ISPDSKVCFS 550 560 570 580 590 600 CCSDGNIAVW DLHNEILVRQ FQGHTDGASC IDISPDGSRL WTGGLDNTVR SWDLREGRQL 610 620 630 640 650 660 QQHDFSSQIF SLGYCPTGDW LAVGMENSHV EVLHASKPDK YQLHLHESCV LSLRFAACGK 670 680 690 700 710 WFVSTGKDNL LNAWRTPYGA SIFQSKETSS VLSCDISTDD KYIVTGSGDK KATVYEVIY

References

	« Hide 'large scale' referencesShow 'large scale' references »
[1]	"A deduced gene product from the Drosophila neurogenic locus, enhancer of split, shows homology to mammalian G-protein beta subunit." Hartley D., Preiss A., Artavanis-Tsakonas S. Cell 55:785-795(1988) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM A).
[2]	"The genome sequence of Drosophila melanogaster." Adams M.D., Celniker S.E., Holt R.A., Evans C.A., Gocayne J.D., Amanatides P.G., Scherer S.E., Li P.W., Hoskins R.A., Galle R.F., George R.A., Lewis S.E., Richards S., Ashburner M., Henderson S.N., Sutton G.G., Wortman J.R., Yandell M.D. , Zhang Q., Chen L.X., Brandon R.C., Rogers Y.-H.C., Blazej R.G., Champe M., Pfeiffer B.D., Wan K.H., Doyle C., Baxter E.G., Helt G., Nelson C.R., Miklos G.L.G., Abril J.F., Agbayani A., An H.-J., Andrews-Pfannkoch C., Baldwin D., Ballew R.M., Basu A., Baxendale J., Bayraktaroglu L., Beasley E.M., Beeson K.Y., Benos P.V., Berman B.P., Bhandari D., Bolshakov S., Borkova D., Botchan M.R., Bouck J., Brokstein P., Brottier P., Burtis K.C., Busam D.A., Butler H., Cadieu E., Center A., Chandra I., Cherry J.M., Cawley S., Dahlke C., Davenport L.B., Davies P., de Pablos B., Delcher A., Deng Z., Mays A.D., Dew I., Dietz S.M., Dodson K., Doup L.E., Downes M., Dugan-Rocha S., Dunkov B.C., Dunn P., Durbin K.J., Evangelista C.C., Ferraz C., Ferriera S., Fleischmann W., Fosler C., Gabrielian A.E., Garg N.S., Gelbart W.M., Glasser K., Glodek A., Gong F., Gorrell J.H., Gu Z., Guan P., Harris M., Harris N.L., Harvey D.A., Heiman T.J., Hernandez J.R., Houck J., Hostin D., Houston K.A., Howland T.J., Wei M.-H., Ibegwam C., Jalali M., Kalush F., Karpen G.H., Ke Z., Kennison J.A., Ketchum K.A., Kimmel B.E., Kodira C.D., Kraft C.L., Kravitz S., Kulp D., Lai Z., Lasko P., Lei Y., Levitsky A.A., Li J.H., Li Z., Liang Y., Lin X., Liu X., Mattei B., McIntosh T.C., McLeod M.P., McPherson D., Merkulov G., Milshina N.V., Mobarry C., Morris J., Moshrefi A., Mount S.M., Moy M., Murphy B., Murphy L., Muzny D.M., Nelson D.L., Nelson D.R., Nelson K.A., Nixon K., Nusskern D.R., Pacleb J.M., Palazzolo M., Pittman G.S., Pan S., Pollard J., Puri V., Reese M.G., Reinert K., Remington K., Saunders R.D.C., Scheeler F., Shen H., Shue B.C., Siden-Kiamos I., Simpson M., Skupski M.P., Smith T.J., Spier E., Spradling A.C., Stapleton M., Strong R., Sun E., Svirskas R., Tector C., Turner R., Venter E., Wang A.H., Wang X., Wang Z.-Y., Wassarman D.A., Weinstock G.M., Weissenbach J., Williams S.M., Woodage T., Worley K.C., Wu D., Yang S., Yao Q.A., Ye J., Yeh R.-F., Zaveri J.S., Zhan M., Zhang G., Zhao Q., Zheng L., Zheng X.H., Zhong F.N., Zhong W., Zhou X., Zhu S.C., Zhu X., Smith H.O., Gibbs R.A., Myers E.W., Rubin G.M., Venter J.C. Science 287:2185-2195(2000) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]. Strain: Berkeley.
[3]	"Annotation of the Drosophila melanogaster euchromatic genome: a systematic review." Misra S., Crosby M.A., Mungall C.J., Matthews B.B., Campbell K.S., Hradecky P., Huang Y., Kaminker J.S., Millburn G.H., Prochnik S.E., Smith C.D., Tupy J.L., Whitfield E.J., Bayraktaroglu L., Berman B.P., Bettencourt B.R., Celniker S.E., de Grey A.D.N.J. , Drysdale R.A., Harris N.L., Richter J., Russo S., Schroeder A.J., Shu S.Q., Stapleton M., Yamada C., Ashburner M., Gelbart W.M., Rubin G.M., Lewis S.E. Genome Biol. 3:RESEARCH0083.1-RESEARCH0083.22(2002) [PubMed] [Europe PMC] [Abstract] Cited for: GENOME REANNOTATION, ALTERNATIVE SPLICING. Strain: Berkeley.
[4]	"A Drosophila complementary DNA resource." Rubin G.M., Hong L., Brokstein P., Evans-Holm M., Frise E., Stapleton M., Harvey D.A. Science 287:2222-2224(2000) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM A). Strain: Berkeley. Tissue: Embryo.
[5]	"Groucho is required for Drosophila neurogenesis, segmentation, and sex determination and interacts directly with hairy-related bHLH proteins." Paroush Z., Finley R.L. Jr., Kidd T., Wainwright S.M., Ingham P.W., Brent R., Ish-Horowicz D. Cell 79:805-815(1994) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION, BINDING TO HAIRY-RELATED PROTEINS, DEVELOPMENTAL STAGE.
[6]	"Phosphoproteome analysis of Drosophila melanogaster embryos." Zhai B., Villen J., Beausoleil S.A., Mintseris J., Gygi S.P. J. Proteome Res. 7:1675-1682(2008) [PubMed] [Europe PMC] [Abstract] Cited for: PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT SER-205; SER-207; SER-218; SER-242; SER-258; SER-267; THR-326 AND THR-328, MASS SPECTROMETRY. Tissue: Embryo.
[7]	"XIAP monoubiquitylates Groucho/TLE to promote canonical Wnt signaling." Hanson A.J., Wallace H.A., Freeman T.J., Beauchamp R.D., Lee L.A., Lee E. Mol. Cell 45:619-628(2012) [PubMed] [Europe PMC] [Abstract] Cited for: UBIQUITINATION BY XIAP/BIRC4.
+	Additional computationally mapped references.

Cross-references

Sequence databases
EMBL GenBank DDBJ	M20571 mRNA. Translation: AAA28512.1. AE014297 Genomic DNA. Translation: AAF56557.1. AE014297 Genomic DNA. Translation: AAN14068.1. AE014297 Genomic DNA. Translation: AAN14069.1. AE014297 Genomic DNA. Translation: AAS65218.1. AF145695 mRNA. Translation: AAD38670.1.
PIR	A30047.
RefSeq	NP_001189293.1. NM_001202364.1. NP_524514.2. NM_079790.5. NP_733133.1. NM_170254.3. NP_733134.1. NM_170255.4. NP_733135.1. NM_170256.2. NP_996298.1. NM_206575.3.
UniGene	Dm.1340.
3D structure databases
ProteinModelPortal	P16371.
SMR	P16371. Positions 401-730.
ModBase	Search...
Protein-protein interaction databases
DIP	DIP-40N.
IntAct	P16371. 20 interactions.
MINT	MINT-314389.
Proteomic databases
PaxDb	P16371.
Protocols and materials databases
StructuralBiologyKnowledgebase	Search...
Genome annotation databases
EnsemblMetazoa	FBtr0084966; FBpp0089115; FBgn0001139.
GeneID	43162.
KEGG	dme:Dmel_CG8384.
UCSC	CG8384-RA. d. melanogaster.
Organism-specific databases
CTD	43162.
FlyBase	FBgn0001139. gro.
Phylogenomic databases
eggNOG	COG2319.
GeneTree	ENSGT00550000074465.
InParanoid	A4V3F6.
KO	K04497.
OMA	HAHVRTN.
OrthoDB	EOG4JM64N.
PhylomeDB	P16371.
Gene expression databases
Bgee	P16371.
GermOnline	CG8384. Drosophila melanogaster.
Family and domain databases
Gene3D	2.130.10.10. 1 hit.
InterPro	IPR005617. Groucho/TLE_N. IPR009146. Groucho_enhance. IPR015943. WD40/YVTN_repeat-like_dom. IPR001680. WD40_repeat. IPR019775. WD40_repeat_CS. IPR017986. WD40_repeat_dom. [Graphical view]
Pfam	PF03920. TLE_N. 1 hit. PF00400. WD40. 5 hits. [Graphical view]
PRINTS	PR01850. GROUCHOFAMLY.
SMART	SM00320. WD40. 7 hits. [Graphical view]
SUPFAM	SSF50978. WD40_like. 1 hit.
PROSITE	PS00678. WD_REPEATS_1. 2 hits. PS50082. WD_REPEATS_2. 3 hits. PS50294. WD_REPEATS_REGION. 2 hits. [Graphical view]
ProtoNet	Search...
Other
ChiTaRS	gro. drosophila.
GenomeRNAi	43162.
NextBio	832487.

Entry information

Entry name

GROU_DROME

Accession

Primary (citable) accession number: P16371
Secondary accession number(s): A4V3F6 , Q0KI08, Q7KRZ4, Q9V3F7

Entry history

Integrated into UniProtKB/Swiss-Prot:	August 1, 1990
Last sequence update:	January 9, 2007
Last modified:	May 1, 2013
This is version 142 of the entry and version 3 of the sequence. [Complete history]

Entry status

Reviewed (UniProtKB/Swiss-Prot)

Annotation program

Drosophila annotation project