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Q9UBP9 (GULP1_HUMAN) Reviewed, UniProtKB/Swiss-Prot

Last modified April 16, 2014. Version 97. Feed History...

Clusters with 100%, 90%, 50% identity | Documents (3) | Third-party data text xml rdf/xml gff fasta
to top of pageNames·Attributes·General annotation·Ontologies·Alt products·Sequence annotation·Sequences·References·Cross-refs·Entry info·DocumentsCustomize order

Names and origin

Protein namesRecommended name:
PTB domain-containing engulfment adapter protein 1
Alternative name(s):
Cell death protein 6 homolog
PTB domain adapter protein CED-6
Protein GULP
Gene names
Name:GULP1
Synonyms:CED6, GULP
OrganismHomo sapiens (Human) [Reference proteome]
Taxonomic identifier9606 [NCBI]
Taxonomic lineageEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresPrimatesHaplorrhiniCatarrhiniHominidaeHomo

Protein attributes

Sequence length304 AA.
Sequence statusComplete.
Protein existenceEvidence at protein level

General annotation (Comments)

Function

May function as an adapter protein. Required for efficient phagocytosis of apoptotic cells. Modulates cellular glycosphingolipid and cholesterol transport. May play a role in the internalization and endosomal trafficking of various LRP1 ligands, such as PSAP. Increases cellular levels of GTP-bound ARF6. Ref.1 Ref.2 Ref.8 Ref.10

Subunit structure

Homodimer. Interacts with clathrin. Interacts with GDP-bound ARF6, but not with GTP-bound ARF6. Part of a complex composed of GULP1, ACAP1 and ARF6. Interacts with ACAP1, LRP1, MEGF10 and STAB2. Ref.7 Ref.9 Ref.10 Ref.11 Ref.12

Subcellular location

Cytoplasm. Note: May associate with the cytoplasmic side of the plasma membrane and early endosomes. Ref.8

Tissue specificity

Widely expressed. Detected in macrophages, pancreas, kidney, skeletal muscle, heart, colon, intestine, lung, placenta and ovary. Ref.2

Sequence similarities

Belongs to the ced-6 family.

Contains 1 PID domain.

Ontologies

Keywords
   Biological processApoptosis
Lipid transport
Phagocytosis
Transport
   Cellular componentCytoplasm
   Coding sequence diversityAlternative splicing
   DomainCoiled coil
   PTMPhosphoprotein
   Technical termComplete proteome
Reference proteome
Gene Ontology (GO)
   Biological_processapoptotic process

Traceable author statement Ref.2. Source: ProtInc

lipid transport

Inferred from electronic annotation. Source: UniProtKB-KW

phagocytosis, engulfment

Inferred from direct assay PubMed 11729193. Source: MGI

signal transduction

Traceable author statement Ref.2Ref.1. Source: GOC

   Cellular_componentcytoplasm

Inferred from electronic annotation. Source: UniProtKB-SubCell

   Molecular_functionsignal transducer activity

Traceable author statement Ref.2Ref.1. Source: ProtInc

Complete GO annotation...

Alternative products

This entry describes 4 isoforms produced by alternative splicing. [Align] [Select]
Isoform 1 (identifier: Q9UBP9-1)

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 2 (identifier: Q9UBP9-2)

The sequence of this isoform differs from the canonical sequence as follows:
     134-167: AEEITLTIGQAFDLAYRKFLESGGKDVETRKQIA → VSIPDVVGWFVLFYKPGIVLLLALAKYLKMNNFL
     168-304: Missing.
Isoform 3 (identifier: Q9UBP9-3)

The sequence of this isoform differs from the canonical sequence as follows:
     31-133: Missing.
Note: No experimental confirmation available.
Isoform 4 (identifier: Q9UBP9-4)

The sequence of this isoform differs from the canonical sequence as follows:
     282-304: EGFKMGLTLEGTVFCLDPLDSRC → RQRWRGSKWD
Note: No experimental confirmation available.

Sequence annotation (Features)

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifier

Molecule processing

Chain1 – 304304PTB domain-containing engulfment adapter protein 1
PRO_0000296679

Regions

Domain21 – 176156PID
Coiled coil158 – 20245 Potential

Amino acid modifications

Modified residue2231Phosphoserine Ref.13

Natural variations

Alternative sequence31 – 133103Missing in isoform 3.
VSP_044703
Alternative sequence134 – 16734AEEIT…RKQIA → VSIPDVVGWFVLFYKPGIVL LLALAKYLKMNNFL in isoform 2.
VSP_027250
Alternative sequence168 – 304137Missing in isoform 2.
VSP_027251
Alternative sequence282 – 30423EGFKM…LDSRC → RQRWRGSKWD in isoform 4.
VSP_045540

Experimental info

Mutagenesis1761L → P: Loss of dimerization; when associated with P-183. Ref.7
Mutagenesis1831L → P: Loss of dimerization; when associated with P-176. Ref.7
Sequence conflict2921G → S in BAG60802. Ref.3

Sequences

Sequence LengthMass (Da)Tools
Isoform 1 [UniParc].

Last modified May 1, 2000. Version 1.
Checksum: 6FCDE040073DBE8B

FASTA30434,490
        10         20         30         40         50         60 
MNRAFSRKKD KTWMHTPEAL SKHFIPYNAK FLGSTEVEQP KGTEVVRDAV RKLKFARHIK 

        70         80         90        100        110        120 
KSEGQKIPKV ELQISIYGVK ILEPKTKEVQ HNCQLHRISF CADDKTDKRI FTFICKDSES 

       130        140        150        160        170        180 
NKHLCYVFDS EKCAEEITLT IGQAFDLAYR KFLESGGKDV ETRKQIAGLQ KRIQDLETEN 

       190        200        210        220        230        240 
MELKNKVQDL ENQLRITQVS APPAGSMTPK SPSTDIFDMI PFSPISHQSS MPTRNGTQPP 

       250        260        270        280        290        300 
PVPSRSTEIK RDLFGAEPFD PFNCGAADFP PDIQSKLDEM QEGFKMGLTL EGTVFCLDPL 


DSRC 

« Hide

Isoform 2 [UniParc].

Checksum: 6A33DC720CB9A2DE
Show »

FASTA16719,423
Isoform 3 [UniParc].

Checksum: 139143B94E5121C4
Show »

FASTA20122,567
Isoform 4 [UniParc].

Checksum: 2736C095A6140AB5
Show »

FASTA29133,333

References

« Hide 'large scale' references
[1]"Human CED-6 encodes a functional homologue of the Caenorhabditis elegans engulfment protein CED-6."
Liu Q.A., Hengartner M.O.
Curr. Biol. 9:1347-1350(1999) [PubMed] [Europe PMC] [Abstract]
Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 1), FUNCTION.
[2]"The human homologue of Caenorhabditis elegans CED-6 specifically promotes phagocytosis of apoptotic cells."
Smits E., Van Criekinge W., Plaetinck G., Bogaert T.
Curr. Biol. 9:1351-1354(1999) [PubMed] [Europe PMC] [Abstract]
Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 1), FUNCTION, TISSUE SPECIFICITY.
[3]"Complete sequencing and characterization of 21,243 full-length human cDNAs."
Ota T., Suzuki Y., Nishikawa T., Otsuki T., Sugiyama T., Irie R., Wakamatsu A., Hayashi K., Sato H., Nagai K., Kimura K., Makita H., Sekine M., Obayashi M., Nishi T., Shibahara T., Tanaka T., Ishii S. expand/collapse author list , Yamamoto J., Saito K., Kawai Y., Isono Y., Nakamura Y., Nagahari K., Murakami K., Yasuda T., Iwayanagi T., Wagatsuma M., Shiratori A., Sudo H., Hosoiri T., Kaku Y., Kodaira H., Kondo H., Sugawara M., Takahashi M., Kanda K., Yokoi T., Furuya T., Kikkawa E., Omura Y., Abe K., Kamihara K., Katsuta N., Sato K., Tanikawa M., Yamazaki M., Ninomiya K., Ishibashi T., Yamashita H., Murakawa K., Fujimori K., Tanai H., Kimata M., Watanabe M., Hiraoka S., Chiba Y., Ishida S., Ono Y., Takiguchi S., Watanabe S., Yosida M., Hotuta T., Kusano J., Kanehori K., Takahashi-Fujii A., Hara H., Tanase T.-O., Nomura Y., Togiya S., Komai F., Hara R., Takeuchi K., Arita M., Imose N., Musashino K., Yuuki H., Oshima A., Sasaki N., Aotsuka S., Yoshikawa Y., Matsunawa H., Ichihara T., Shiohata N., Sano S., Moriya S., Momiyama H., Satoh N., Takami S., Terashima Y., Suzuki O., Nakagawa S., Senoh A., Mizoguchi H., Goto Y., Shimizu F., Wakebe H., Hishigaki H., Watanabe T., Sugiyama A., Takemoto M., Kawakami B., Yamazaki M., Watanabe K., Kumagai A., Itakura S., Fukuzumi Y., Fujimori Y., Komiyama M., Tashiro H., Tanigami A., Fujiwara T., Ono T., Yamada K., Fujii Y., Ozaki K., Hirao M., Ohmori Y., Kawabata A., Hikiji T., Kobatake N., Inagaki H., Ikema Y., Okamoto S., Okitani R., Kawakami T., Noguchi S., Itoh T., Shigeta K., Senba T., Matsumura K., Nakajima Y., Mizuno T., Morinaga M., Sasaki M., Togashi T., Oyama M., Hata H., Watanabe M., Komatsu T., Mizushima-Sugano J., Satoh T., Shirai Y., Takahashi Y., Nakagawa K., Okumura K., Nagase T., Nomura N., Kikuchi H., Masuho Y., Yamashita R., Nakai K., Yada T., Nakamura Y., Ohara O., Isogai T., Sugano S.
Nat. Genet. 36:40-45(2004) [PubMed] [Europe PMC] [Abstract]
Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORMS 1; 3 AND 4).
Tissue: Neuroblastoma and Trachea.
[4]"Generation and annotation of the DNA sequences of human chromosomes 2 and 4."
Hillier L.W., Graves T.A., Fulton R.S., Fulton L.A., Pepin K.H., Minx P., Wagner-McPherson C., Layman D., Wylie K., Sekhon M., Becker M.C., Fewell G.A., Delehaunty K.D., Miner T.L., Nash W.E., Kremitzki C., Oddy L., Du H. expand/collapse author list , Sun H., Bradshaw-Cordum H., Ali J., Carter J., Cordes M., Harris A., Isak A., van Brunt A., Nguyen C., Du F., Courtney L., Kalicki J., Ozersky P., Abbott S., Armstrong J., Belter E.A., Caruso L., Cedroni M., Cotton M., Davidson T., Desai A., Elliott G., Erb T., Fronick C., Gaige T., Haakenson W., Haglund K., Holmes A., Harkins R., Kim K., Kruchowski S.S., Strong C.M., Grewal N., Goyea E., Hou S., Levy A., Martinka S., Mead K., McLellan M.D., Meyer R., Randall-Maher J., Tomlinson C., Dauphin-Kohlberg S., Kozlowicz-Reilly A., Shah N., Swearengen-Shahid S., Snider J., Strong J.T., Thompson J., Yoakum M., Leonard S., Pearman C., Trani L., Radionenko M., Waligorski J.E., Wang C., Rock S.M., Tin-Wollam A.-M., Maupin R., Latreille P., Wendl M.C., Yang S.-P., Pohl C., Wallis J.W., Spieth J., Bieri T.A., Berkowicz N., Nelson J.O., Osborne J., Ding L., Meyer R., Sabo A., Shotland Y., Sinha P., Wohldmann P.E., Cook L.L., Hickenbotham M.T., Eldred J., Williams D., Jones T.A., She X., Ciccarelli F.D., Izaurralde E., Taylor J., Schmutz J., Myers R.M., Cox D.R., Huang X., McPherson J.D., Mardis E.R., Clifton S.W., Warren W.C., Chinwalla A.T., Eddy S.R., Marra M.A., Ovcharenko I., Furey T.S., Miller W., Eichler E.E., Bork P., Suyama M., Torrents D., Waterston R.H., Wilson R.K.
Nature 434:724-731(2005) [PubMed] [Europe PMC] [Abstract]
Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
[5]Mural R.J., Istrail S., Sutton G.G., Florea L., Halpern A.L., Mobarry C.M., Lippert R., Walenz B., Shatkay H., Dew I., Miller J.R., Flanigan M.J., Edwards N.J., Bolanos R., Fasulo D., Halldorsson B.V., Hannenhalli S., Turner R. expand/collapse author list , Yooseph S., Lu F., Nusskern D.R., Shue B.C., Zheng X.H., Zhong F., Delcher A.L., Huson D.H., Kravitz S.A., Mouchard L., Reinert K., Remington K.A., Clark A.G., Waterman M.S., Eichler E.E., Adams M.D., Hunkapiller M.W., Myers E.W., Venter J.C.
Submitted (SEP-2005) to the EMBL/GenBank/DDBJ databases
Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
[6]"The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC)."
The MGC Project Team
Genome Res. 14:2121-2127(2004) [PubMed] [Europe PMC] [Abstract]
Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM 2).
Tissue: Placenta.
[7]"Identification and characterization of a dimerization domain in CED-6, an adapter protein involved in engulfment of apoptotic cells."
Su H.P., Brugnera E., Van Criekinge W., Smits E., Hengartner M., Bogaert T., Ravichandran K.S.
J. Biol. Chem. 275:9542-9549(2000) [PubMed] [Europe PMC] [Abstract]
Cited for: SUBUNIT, MUTAGENESIS OF LEU-176 AND LEU-183.
[8]"The lipoprotein receptor-related protein-1 (LRP) adapter protein GULP mediates trafficking of the LRP ligand prosaposin, leading to sphingolipid and free cholesterol accumulation in late endosomes and impaired efflux."
Kiss R.S., Ma Z., Nakada-Tsukui K., Brugnera E., Vassiliou G., McBride H.M., Ravichandran K.S., Marcel Y.L.
J. Biol. Chem. 281:12081-12092(2006) [PubMed] [Europe PMC] [Abstract]
Cited for: FUNCTION, SUBCELLULAR LOCATION.
[9]"Cooperation between engulfment receptors: the case of ABCA1 and MEGF10."
Hamon Y., Trompier D., Ma Z., Venegas V., Pophillat M., Mignotte V., Zhou Z., Chimini G.
PLoS ONE 1:E120-E120(2006) [PubMed] [Europe PMC] [Abstract]
Cited for: INTERACTION WITH MEGF10.
[10]"Regulation of Arf6 and ACAP1 signaling by the PTB-domain-containing adaptor protein GULP."
Ma Z., Nie Z., Luo R., Casanova J.E., Ravichandran K.S.
Curr. Biol. 17:722-727(2007) [PubMed] [Europe PMC] [Abstract]
Cited for: FUNCTION, INTERACTION WITH ARF6 AND ACAP1, IDENTIFICATION IN A COMPLEX WITH ACAP1 AND ARF6.
[11]"MEGF10 is a mammalian ortholog of CED-1 that interacts with clathrin assembly protein complex 2 medium chain and induces large vacuole formation."
Suzuki E., Nakayama M.
Exp. Cell Res. 313:3729-3742(2007) [PubMed] [Europe PMC] [Abstract]
Cited for: INTERACTION WITH MEGF10.
[12]"Requirement of adaptor protein GULP during stabilin-2-mediated cell corpse engulfment."
Park S.Y., Kang K.B., Thapa N., Kim S.Y., Lee S.J., Kim I.S.
J. Biol. Chem. 283:10593-10600(2008) [PubMed] [Europe PMC] [Abstract]
Cited for: INTERACTION WITH STAB2.
[13]"Combining protein-based IMAC, peptide-based IMAC, and MudPIT for efficient phosphoproteomic analysis."
Cantin G.T., Yi W., Lu B., Park S.K., Xu T., Lee J.-D., Yates J.R. III
J. Proteome Res. 7:1346-1351(2008) [PubMed] [Europe PMC] [Abstract]
Cited for: PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT SER-223, IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS].
Tissue: Cervix carcinoma.
[14]"Initial characterization of the human central proteome."
Burkard T.R., Planyavsky M., Kaupe I., Breitwieser F.P., Buerckstuemmer T., Bennett K.L., Superti-Furga G., Colinge J.
BMC Syst. Biol. 5:17-17(2011) [PubMed] [Europe PMC] [Abstract]
Cited for: IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS].
[15]"System-wide temporal characterization of the proteome and phosphoproteome of human embryonic stem cell differentiation."
Rigbolt K.T., Prokhorova T.A., Akimov V., Henningsen J., Johansen P.T., Kratchmarova I., Kassem M., Mann M., Olsen J.V., Blagoev B.
Sci. Signal. 4:RS3-RS3(2011) [PubMed] [Europe PMC] [Abstract]
Cited for: IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS].
+Additional computationally mapped references.

Cross-references

Sequence databases

EMBL
GenBank
DDBJ
AF200715 mRNA. Translation: AAF08006.1.
AF191771 mRNA. Translation: AAF18975.1.
AK055718 mRNA. No translation available.
AK298626 mRNA. Translation: BAG60802.1.
AK314498 mRNA. Translation: BAG37098.1.
AC092598 Genomic DNA. Translation: AAX93242.1.
AC125490 Genomic DNA. Translation: AAY24122.1.
AC104131 Genomic DNA. No translation available.
AC108493 Genomic DNA. No translation available.
AC133606 Genomic DNA. No translation available.
CH471058 Genomic DNA. Translation: EAX10913.1.
CH471058 Genomic DNA. Translation: EAX10914.1.
CH471058 Genomic DNA. Translation: EAX10915.1.
CH471058 Genomic DNA. Translation: EAX10917.1.
BC001103 mRNA. Translation: AAH01103.1.
RefSeqNP_001239597.1. NM_001252668.1.
NP_001239598.1. NM_001252669.1.
NP_057399.1. NM_016315.3.
UniGeneHs.470887.
Hs.712910.

3D structure databases

ProteinModelPortalQ9UBP9.
SMRQ9UBP9. Positions 18-150.
ModBaseSearch...
MobiDBSearch...

Protein-protein interaction databases

BioGrid119550. 5 interactions.
IntActQ9UBP9. 5 interactions.
MINTMINT-259085.
STRING9606.ENSP00000352047.

PTM databases

PhosphoSiteQ9UBP9.

Polymorphism databases

DMDM74720076.

Proteomic databases

PaxDbQ9UBP9.
PRIDEQ9UBP9.

Protocols and materials databases

DNASU51454.
StructuralBiologyKnowledgebaseSearch...

Genome annotation databases

EnsemblENST00000359135; ENSP00000352047; ENSG00000144366. [Q9UBP9-1]
ENST00000409580; ENSP00000386289; ENSG00000144366. [Q9UBP9-1]
ENST00000409609; ENSP00000386867; ENSG00000144366. [Q9UBP9-1]
ENST00000409637; ENSP00000386402; ENSG00000144366. [Q9UBP9-2]
ENST00000409805; ENSP00000387171; ENSG00000144366. [Q9UBP9-3]
ENST00000409830; ENSP00000386732; ENSG00000144366. [Q9UBP9-1]
ENST00000409843; ENSP00000387144; ENSG00000144366. [Q9UBP9-4]
ENST00000410051; ENSP00000387013; ENSG00000144366. [Q9UBP9-2]
GeneID51454.
KEGGhsa:51454.
UCSCuc002uqc.4. human. [Q9UBP9-2]
uc002uqf.3. human. [Q9UBP9-1]

Organism-specific databases

CTD51454.
GeneCardsGC02P189156.
H-InvDBHIX0022389.
HGNCHGNC:18649. GULP1.
HPAHPA020587.
MIM608165. gene.
neXtProtNX_Q9UBP9.
PharmGKBPA134993283.
GenAtlasSearch...

Phylogenomic databases

eggNOGNOG314249.
HOGENOMHOG000039986.
HOVERGENHBG056801.
InParanoidQ9UBP9.
OMAPFDPFSC.
OrthoDBEOG7RBZ90.
PhylomeDBQ9UBP9.
TreeFamTF314159.

Gene expression databases

ArrayExpressQ9UBP9.
BgeeQ9UBP9.
CleanExHS_GULP1.
GenevestigatorQ9UBP9.

Family and domain databases

Gene3D2.30.29.30. 1 hit.
InterProIPR011993. PH_like_dom.
IPR006020. PTB/PI_dom.
[Graphical view]
PfamPF00640. PID. 1 hit.
[Graphical view]
SMARTSM00462. PTB. 1 hit.
[Graphical view]
PROSITEPS01179. PID. 1 hit.
[Graphical view]
ProtoNetSearch...

Other

ChiTaRSGULP1. human.
GenomeRNAi51454.
NextBio55065.
PROQ9UBP9.
SOURCESearch...

Entry information

Entry nameGULP1_HUMAN
AccessionPrimary (citable) accession number: Q9UBP9
Secondary accession number(s): B2RB51 expand/collapse secondary AC list , B4DQ40, B8ZZ72, D3DPH1, E9PB86, Q53PC1, Q53RF3, Q9BVL3
Entry history
Integrated into UniProtKB/Swiss-Prot: July 24, 2007
Last sequence update: May 1, 2000
Last modified: April 16, 2014
This is version 97 of the entry and version 1 of the sequence. [Complete history]
Entry statusReviewed (UniProtKB/Swiss-Prot)
Annotation programChordata Protein Annotation Program
DisclaimerAny medical or genetic information present in this entry is provided for research, educational and informational purposes only. It is not in any way intended to be used as a substitute for professional medical advice, diagnosis, treatment or care.

Relevant documents

SIMILARITY comments

Index of protein domains and families

MIM cross-references

Online Mendelian Inheritance in Man (MIM) cross-references in UniProtKB/Swiss-Prot

Human chromosome 2

Human chromosome 2: entries, gene names and cross-references to MIM