P15330 (DORS_DROME) Reviewed, UniProtKB/Swiss-Prot

Names and origin

Protein names

Recommended name:
Embryonic polarity protein dorsal

Gene names

Name:	dl
ORF Names:	CG6667

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	999 AA.
Sequence status	Complete.
Protein existence	Evidence at protein level

General annotation (Comments)

Function	Embryonic developmental protein. The lateral or ventral identity of a cell depends upon the concentration of dorsal protein in its nucleus during the blastoderm stage. Dorsal is a morphogenetic protein that specifically binds to the kappa B-related consensus sequence 5'-GRGAAAANCC-3', located in the enhancer region of zygotic genes such as Zen, Twist, Snail and Decapentaplegic. Mediates an immune response in Drosophila larvae. Ref.3
Subunit structure	Interacts with tamo via the nuclear localization signal. Interacts with emb, a component of the nuclear pore complex. Ref.7 Ref.8
Subcellular location	Cytoplasm. Nucleus. Note: In ventral regions it is first cytoplasmic, then the protein is relocalized in the nucleus. Its nuclear localization is essential to its function as a morphogen. In dorsal regions it remains cytoplasmic. Tamo negatively regulates nuclear import of dl. Emb in the nuclear pore complex is responsible for export of dl from the nucleus. Ref.2 Ref.7 Ref.8 Isoform A: Nucleus Probable Ref.2 Ref.7 Ref.8.
Tissue specificity	In unchallenged larvae, expression of both isoforms is seen in fat body and gut (isoform A is more abundant). After immune challenge levels of both isoforms are enhanced. Ref.3
Developmental stage	Isoform A is expressed maternally and both isoforms are expressed zygotically from 6-9 hours embryos through to adulthood. Ref.3
Sequence similarities	Contains 1 RHD (Rel-like) domain.

Ontologies

Keywords
Biological process	Transcription Transcription regulation
Cellular component	Cytoplasm Nucleus
Coding sequence diversity	Alternative splicing
Ligand	DNA-binding
Molecular function	Activator Developmental protein
PTM	Phosphoprotein
Technical term	Complete proteome Reference proteome
Gene Ontology (GO)
Biological_process	Toll signaling pathway Traceable author statement PubMed 11283698PubMed 12530956PubMed 15199954. Source: FlyBase anterior/posterior pattern specification Traceable author statement PubMed 9988212. Source: FlyBase dorsal/ventral axis specification Traceable author statement PubMed 10793472PubMed 11700289PubMed 12217523. Source: FlyBase ectodermal cell fate specification Traceable author statement PubMed 10878576. Source: FlyBase gastrulation involving germ band extension Traceable author statement PubMed 15380237. Source: FlyBase germ cell migration Traceable author statement PubMed 9988212. Source: FlyBase heart development Non-traceable author statement PubMed 12027431. Source: FlyBase melanization defense response Inferred from mutant phenotype PubMed 15211580. Source: FlyBase mesodermal cell fate specification Traceable author statement PubMed 10878576. Source: FlyBase negative regulation of transcription from RNA polymerase II promoter Traceable author statement PubMed 11700289. Source: FlyBase peripheral nervous system neuron development Inferred from mutant phenotype PubMed 18000549. Source: FlyBase plasmatocyte differentiation Inferred from mutant phenotype PubMed 15882582. Source: FlyBase regulation of alternative mRNA splicing, via spliceosome Inferred from mutant phenotype PubMed 15492211. Source: FlyBase regulation of hemocyte proliferation Inferred from direct assay PubMed 17060622. Source: FlyBase ventral cord development Non-traceable author statement PubMed 12027434. Source: FlyBase
Cellular_component	cytoplasm Inferred from direct assay PubMed 11943764PubMed 9510254. Source: FlyBase neuromuscular junction Inferred from direct assay PubMed 12532402. Source: FlyBase nucleus Inferred from direct assay PubMed 11943764PubMed 12556495PubMed 22939625PubMed 9510254. Source: FlyBase
Molecular_function	RNA polymerase II core promoter proximal region sequence-specific DNA binding transcription factor activity involved in positive regulation of transcription Inferred from direct assay PubMed 1648449. Source: FlyBase RNA polymerase II regulatory region sequence-specific DNA binding Inferred from direct assay PubMed 1988156PubMed 9774673PubMed 9806924. Source: FlyBase
Complete GO annotation...

Binary interactions

With	Entry	#Exp.	IntAct	Notes
cact	Q03017	4	EBI-198375,EBI-200600
tamo	Q9W1A4	4	EBI-198375,EBI-91385
tub	P22812	3	EBI-198375,EBI-93181

Alternative products

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

Isoform C (identifier: P15330-1)

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.

Isoform A (identifier: P15330-2)

Also known as: B;

The sequence of this isoform differs from the canonical sequence as follows:
     330-677: GKHTFWNLHR...NSQARKPETP → DPAHLRRKRQ...LQISNLSIST
     678-999: Missing.

Note: Nuclear localization signal at positions 335-340.

Sequence annotation (Features)

	Feature key	Position(s)	Length	Description	Graphical view	Feature identifier
Molecule processing
	Chain	1 – 999	999	Embryonic polarity protein dorsal		PRO_0000205164
Regions
	Domain	47 – 342	296	RHD
	Motif	756 – 773	18	Nuclear localization signal Potential
	Compositional bias	395 – 464	70	Gln-rich
	Compositional bias	635 – 705	71	Pro-rich
Amino acid modifications
	Modified residue	312	1	Phosphoserine; by PKA Potential
Natural variations
	Alternative sequence	330 – 677	348	GKHTF…KPETP → DPAHLRRKRQKTGGDPMHLL LQQQQKQQLQNDHQDGRQTN MNCWNTQNIPPIKTEPRDTS PQPFGLSYRAPPELTPSPQP LSPSSNYNHNSTPSPYNMAS AVTPTNGQQQLMSPNHPQQQ QQQQQYGATDLGSNYNPFAQ QVLAQQQQHQQQQQQHQHQH QQQHQQQQQQQQQQQQQSLQ FHANPFGNPGGNSWESKFSA AAVAAAAATATGAAPANGNS NNLSNLNNPFTMHNLLTSGG GPGNANNLQWNLTTNHLHNQ HTLHQQQQLQQQQQQQYDNT APTNNNANLNNNNNNNNTAG NQADNNGPTLSNLLSFDSGQ LVHINSEDQQILRLNSEDLQ ISNLSIST in isoform A.		VSP_005581
	Alternative sequence	678 – 999	322	Missing in isoform A.		VSP_005582
Experimental info
	Sequence conflict	236	1	T → S in AAT94434. Ref.6
	Sequence conflict	391	1	H → Q in AAC35296. Ref.3
	Sequence conflict	401	1	L → F in AAC35296. Ref.3
	Sequence conflict	407	1	V → SQQEQYTQEQSL in AAC35296. Ref.3
	Sequence conflict	698 – 699	2	DK → EQ in AAC35296. Ref.3
	Sequence conflict	965 – 999	35	ASEFD…AANGI → PVNLTRPPPTMLPWMLARF in AAC35296. Ref.3
Isoform A:
	Sequence conflict	506	1	Q → QQ in AAA28479. Ref.1

Sequences

	Sequence		Length	Mass (Da)
	Isoform C [UniParc]. Last modified June 1, 2001. Version 2. Checksum: E29C6594AC07D662	FASTA	999	111,551
	10 20 30 40 50 60 MFPNQNNGAA PGQGPAVDGQ QSLNYNGLPA QQQQQLAQST KNVRKKPYVK ITEQPAGKAL 70 80 90 100 110 120 RFRYECEGRS AGSIPGVNST PENKTYPTIE IVGYKGRAVV VVSCVTKDTP YRPHPHNLVG 130 140 150 160 170 180 KEGCKKGVCT LEINSETMRA VFSNLGIQCV KKKDIEAALK AREEIRVDPF KTGFSHRFQP 190 200 210 220 230 240 SSIDLNSVRL CFQVFMESEQ KGRFTSPLPP VVSEPIFDKK AMSDLVICRL CSCSATVFGN 250 260 270 280 290 300 TQIILLCEKV AKEDISVRFF EEKNGQSVWE AFGDFQHTDV HKQTAITFKT PRYHTLDITE 310 320 330 340 350 360 PAKVFIQLRR PSDGVTSEAL PFEYVPMDSG KHTFWNLHRH LKRKPDEDLF QQILRLDAKR 370 380 390 400 410 420 EVQPPTIEVI DLDTPKIDVQ REIPSEMEFN HEESQQSEPA LEQEQSVQQE QYTQEQSLQQ 430 440 450 460 470 480 EQYTQEQSLQ QEQYLQQLEQ QQSFQLEEPM QQDQELPAQQ SFDQAIDHLP DHTSDHIPED 490 500 510 520 530 540 MEAADAHAEA EAHRLRSEQE KEIDTIIDEK VRELEQLDLG QQLEPRPLTA NDKITEWMKS 550 560 570 580 590 600 SEIEQQVHEP SPTAEADVLD SALEISKADK TLDELLETVA ELDEIYTDFK VQRDTYKNTI 610 620 630 640 650 660 QNELAGLQGR APLQVEDSFD DAATYTSLQI AFKNPVLIPM DDIMPPTPPM SQCAPEDAHQ 670 680 690 700 710 720 HYDPVEVNSQ ARKPETPMRP VPPVPPAILT IQYPPEEDKL PPLPPKRIRK QDSNAENRSI 730 740 750 760 770 780 EANTVQTKPS TGESPLNKRL PPAPKNPNFN TLPRQKKPGF FSKLFSRRKS KPDLAQGQEN 790 800 810 820 830 840 SSILDSKANS REPSIGHFNM QDPMRASLRS SKSAAPFISN PAPAKSSPVK AKKPGSKLTK 850 860 870 880 890 900 PVGRSVSSVS GKRPAYLNAD VVHIPLKGDS VNSLPQQQRT EGYSQSSTIS VGAGLDRRTA 910 920 930 940 950 960 SALQLADIPI SEGGMELVAI ADRQSLHNLV SSIEGHFNVQ LDPNLDLTEA EHFALYTSIP 970 980 990 PLAAASEFDE TSAYYAPVDA GEILTPDEVA KRLAAANGI
	Isoform A (B) [UniParc]. Checksum: 10FEFCEB1D3A5789	FASTA	677	75,347
	10 20 30 40 50 60 MFPNQNNGAA PGQGPAVDGQ QSLNYNGLPA QQQQQLAQST KNVRKKPYVK ITEQPAGKAL 70 80 90 100 110 120 RFRYECEGRS AGSIPGVNST PENKTYPTIE IVGYKGRAVV VVSCVTKDTP YRPHPHNLVG 130 140 150 160 170 180 KEGCKKGVCT LEINSETMRA VFSNLGIQCV KKKDIEAALK AREEIRVDPF KTGFSHRFQP 190 200 210 220 230 240 SSIDLNSVRL CFQVFMESEQ KGRFTSPLPP VVSEPIFDKK AMSDLVICRL CSCSATVFGN 250 260 270 280 290 300 TQIILLCEKV AKEDISVRFF EEKNGQSVWE AFGDFQHTDV HKQTAITFKT PRYHTLDITE 310 320 330 340 350 360 PAKVFIQLRR PSDGVTSEAL PFEYVPMDSD PAHLRRKRQK TGGDPMHLLL QQQQKQQLQN 370 380 390 400 410 420 DHQDGRQTNM NCWNTQNIPP IKTEPRDTSP QPFGLSYRAP PELTPSPQPL SPSSNYNHNS 430 440 450 460 470 480 TPSPYNMASA VTPTNGQQQL MSPNHPQQQQ QQQQYGATDL GSNYNPFAQQ VLAQQQQHQQ 490 500 510 520 530 540 QQQQHQHQHQ QQHQQQQQQQ QQQQQQSLQF HANPFGNPGG NSWESKFSAA AVAAAAATAT 550 560 570 580 590 600 GAAPANGNSN NLSNLNNPFT MHNLLTSGGG PGNANNLQWN LTTNHLHNQH TLHQQQQLQQ 610 620 630 640 650 660 QQQQQYDNTA PTNNNANLNN NNNNNNTAGN QADNNGPTLS NLLSFDSGQL VHINSEDQQI 670 LRLNSEDLQI SNLSIST

References

	« Hide 'large scale' referencesShow 'large scale' references »
[1]	"Dorsal, an embryonic polarity gene in Drosophila, is homologous to the vertebrate proto-oncogene, c-rel." Steward R. Science 238:692-694(1987) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM A).
[2]	"Relocalization of the dorsal protein from the cytoplasm to the nucleus correlates with its function." Steward R. Cell 59:1179-1188(1989) [PubMed] [Europe PMC] [Abstract] Cited for: SEQUENCE REVISION, SUBCELLULAR LOCATION.
[3]	"Dorsal-B, a splice variant of the Drosophila factor Dorsal, is a novel Rel/NF-kappaB transcriptional activator." Gross I., Georgel P., Oertel-Buchheit P., Schnarr M., Reichhart J.-M. Gene 228:233-242(1999) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM A), FUNCTION, TISSUE SPECIFICITY, DEVELOPMENTAL STAGE, ALTERNATIVE SPLICING. Strain: Oregon-R.
[4]	"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.
[5]	"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.
[6]	Stapleton M., Carlson J.W., Chavez C., Frise E., George R.A., Pacleb J.M., Park S., Wan K.H., Yu C., Rubin G.M., Celniker S.E. Submitted (AUG-2004) to the EMBL/GenBank/DDBJ databases Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM A). Strain: Berkeley. Tissue: Embryo.
[7]	"Tamo selectively modulates nuclear import in Drosophila." Minakhina S., Yang J., Steward R. Genes Cells 8:299-310(2003) [PubMed] [Europe PMC] [Abstract] Cited for: INTERACTION WITH TAMO, SUBCELLULAR LOCATION.
[8]	"The Drosophila nucleoporin DNup88 localizes DNup214 and CRM1 on the nuclear envelope and attenuates NES-mediated nuclear export." Roth P., Xylourgidis N., Sabri N., Uv A.E., Fornerod M., Samakovlis C. J. Cell Biol. 163:701-706(2003) [PubMed] [Europe PMC] [Abstract] Cited for: INTERACTION WITH EMB, SUBCELLULAR LOCATION.
+	Additional computationally mapped references.

Cross-references

Sequence databases
EMBL GenBank DDBJ	M23702 mRNA. Translation: AAA28479.1. AF053614 mRNA. Translation: AAC35296.1. AE014134 Genomic DNA. Translation: AAF53611.1. AE014134 Genomic DNA. Translation: AAF53612.1. BT015205 mRNA. Translation: AAT94434.1.
PIR	A30350.
RefSeq	NP_001163000.1. NM_001169529.1. NP_001163001.1. NM_001169530.1. NP_724052.1. NM_165217.2. NP_724053.1. NM_165218.2. NP_724054.1. NM_165219.1.
UniGene	Dm.3233.
3D structure databases
ProteinModelPortal	P15330.
SMR	P15330. Positions 47-354.
ModBase	Search...
Protein-protein interaction databases
DIP	DIP-17423N.
IntAct	P15330. 5 interactions.
MINT	MINT-277507.
Proteomic databases
PaxDb	P15330.
Protocols and materials databases
StructuralBiologyKnowledgebase	Search...
Genome annotation databases
EnsemblMetazoa	FBtr0081007; FBpp0080560; FBgn0260632.
GeneID	35047.
KEGG	dme:Dmel_CG6667.
UCSC	CG6667-RA. d. melanogaster.
Organism-specific databases
CTD	35047.
FlyBase	FBgn0260632. dl.
Phylogenomic databases
eggNOG	NOG119056.
GeneTree	ENSGT00500000044765.
InParanoid	P15330.
KO	K09255.
OMA	QLMSPNH.
OrthoDB	EOG4HDR8J.
PhylomeDB	P15330.
Gene expression databases
Bgee	P15330.
GermOnline	CG6667. Drosophila melanogaster.
Family and domain databases
Gene3D	2.60.40.10. 1 hit. 2.60.40.340. 1 hit.
InterPro	IPR013783. Ig-like_fold. IPR014756. Ig_E-set. IPR002909. IPT_TIG_rcpt. IPR000451. NF_Rel_Dor. IPR008967. p53-like_TF_DNA-bd. IPR011539. RHD. [Graphical view]
Pfam	PF00554. RHD. 1 hit. [Graphical view]
PRINTS	PR00057. NFKBTNSCPFCT.
SMART	SM00429. IPT. 1 hit. [Graphical view]
SUPFAM	SSF81296. Ig_E-set. 1 hit. SSF49417. P53_like_DNA_bnd. 1 hit.
PROSITE	PS01204. REL_1. 1 hit. PS50254. REL_2. 1 hit. [Graphical view]
ProtoNet	Search...
Other
ChiTaRS	Dl. drosophila.
GenomeRNAi	35047.
NextBio	791548.

Entry information

Entry name

DORS_DROME

Accession

Primary (citable) accession number: P15330
Secondary accession number(s): O77088 , Q0E8P9, Q6AWP3, Q9VJD9, Q9VJE0

Entry history

Integrated into UniProtKB/Swiss-Prot:	April 1, 1990
Last sequence update:	June 1, 2001
Last modified:	May 1, 2013
This is version 153 of the entry and version 2 of the sequence. [Complete history]

Entry status

Reviewed (UniProtKB/Swiss-Prot)

Annotation program

Drosophila annotation project