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Solute carrier family 22 member 4 (Ergothioneine transporter) (ET transporter) (Organic cation/carnitine transporter 1)

 S22A4_HUMAN             Reviewed;         551 AA.
Q9H015; O14546;
19-JUL-2004, integrated into UniProtKB/Swiss-Prot.
01-MAY-2007, sequence version 3.
12-SEP-2018, entry version 150.
RecName: Full=Solute carrier family 22 member 4;
AltName: Full=Ergothioneine transporter;
Short=ET transporter;
AltName: Full=Organic cation/carnitine transporter 1;
Name=SLC22A4; Synonyms=ETT, OCTN1, UT2H;
Homo sapiens (Human).
Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi;
Mammalia; Eutheria; Euarchontoglires; Primates; Haplorrhini;
Catarrhini; Hominidae; Homo.
NCBI_TaxID=9606;
[1]
NUCLEOTIDE SEQUENCE [MRNA], TISSUE SPECIFICITY, BIOPHYSICOCHEMICAL
PROPERTIES, AND VARIANTS THR-306 AND PHE-503.
TISSUE=Fetal liver;
PubMed=9426230; DOI=10.1016/S0014-5793(97)01441-5;
Tamai I., Yabuuchi H., Nezu J., Sai Y., Oku A., Shimane M., Tsuji A.;
"Cloning and characterization of a novel human pH-dependent organic
cation transporter, OCTN1.";
FEBS Lett. 419:107-111(1997).
[2]
NUCLEOTIDE SEQUENCE [MRNA], AND FUNCTION.
PubMed=15795384; DOI=10.1073/pnas.0408624102;
Gruendemann D., Harlfinger S., Golz S., Geerts A., Lazar A.,
Berkels R., Jung N., Rubbert A., Schoemig E.;
"Discovery of the ergothioneine transporter.";
Proc. Natl. Acad. Sci. U.S.A. 102:5256-5261(2005).
[3]
NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
PubMed=15372022; DOI=10.1038/nature02919;
Schmutz J., Martin J., Terry A., Couronne O., Grimwood J., Lowry S.,
Gordon L.A., Scott D., Xie G., Huang W., Hellsten U., Tran-Gyamfi M.,
She X., Prabhakar S., Aerts A., Altherr M., Bajorek E., Black S.,
Branscomb E., Caoile C., Challacombe J.F., Chan Y.M., Denys M.,
Detter J.C., Escobar J., Flowers D., Fotopulos D., Glavina T.,
Gomez M., Gonzales E., Goodstein D., Grigoriev I., Groza M.,
Hammon N., Hawkins T., Haydu L., Israni S., Jett J., Kadner K.,
Kimball H., Kobayashi A., Lopez F., Lou Y., Martinez D., Medina C.,
Morgan J., Nandkeshwar R., Noonan J.P., Pitluck S., Pollard M.,
Predki P., Priest J., Ramirez L., Retterer J., Rodriguez A.,
Rogers S., Salamov A., Salazar A., Thayer N., Tice H., Tsai M.,
Ustaszewska A., Vo N., Wheeler J., Wu K., Yang J., Dickson M.,
Cheng J.-F., Eichler E.E., Olsen A., Pennacchio L.A., Rokhsar D.S.,
Richardson P., Lucas S.M., Myers R.M., Rubin E.M.;
"The DNA sequence and comparative analysis of human chromosome 5.";
Nature 431:268-274(2004).
[4]
NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
TISSUE=Colon;
PubMed=15489334; DOI=10.1101/gr.2596504;
The MGC Project Team;
"The status, quality, and expansion of the NIH full-length cDNA
project: the Mammalian Gene Collection (MGC).";
Genome Res. 14:2121-2127(2004).
[5]
FUNCTION.
PubMed=10215651;
Yabuuchi H., Tamai I., Nezu J., Sakamoto K., Oku A., Shimane M.,
Sai Y., Tsuji A.;
"Novel membrane transporter OCTN1 mediates multispecific,
bidirectional, and pH-dependent transport of organic cations.";
J. Pharmacol. Exp. Ther. 289:768-773(1999).
[6]
INDUCTION, TISSUE SPECIFICITY, AND INVOLVEMENT IN RA.
PubMed=14608356; DOI=10.1038/ng1267;
Tokuhiro S., Yamada R., Chang X., Suzuki A., Kochi Y., Sawada T.,
Suzuki M., Nagasaki M., Ohtsuki M., Ono M., Furukawa H., Nagashima M.,
Yoshino S., Mabuchi A., Sekine A., Saito S., Takahashi A., Tsunoda T.,
Nakamura Y., Yamamoto K.;
"An intronic SNP in a RUNX1 binding site of SLC22A4, encoding an
organic cation transporter, is associated with rheumatoid arthritis.";
Nat. Genet. 35:341-348(2003).
[7]
VARIANTS THR-306 AND GLU-462.
PubMed=12436193; DOI=10.1007/s100380200088;
Saito S., Iida A., Sekine A., Ogawa C., Kawauchi S., Higuchi S.,
Nakamura Y.;
"Catalog of 238 variations among six human genes encoding solute
carriers (hSLCs) in the Japanese population.";
J. Hum. Genet. 47:576-584(2002).
[8]
CHARACTERIZATION OF VARIANT GLU-462, AND BIOPHYSICOCHEMICAL
PROPERTIES.
PubMed=15459889; DOI=10.1002/jps.20190;
Kawasaki Y., Kato Y., Sai Y., Tsuji A.;
"Functional characterization of human organic cation transporter OCTN1
single nucleotide polymorphisms in the Japanese population.";
J. Pharm. Sci. 93:2920-2926(2004).
[9]
VARIANT PHE-503, AND TISSUE SPECIFICITY.
PubMed=15107849; DOI=10.1038/ng1339;
Peltekova V.D., Wintle R.F., Rubin L.A., Amos C.I., Huang Q., Gu X.,
Newman B., Van Oene M., Cescon D., Greenberg G., Griffiths A.M.,
St George-Hyslop P.H., Siminovitch K.A.;
"Functional variants of OCTN cation transporter genes are associated
with Crohn disease.";
Nat. Genet. 36:471-475(2004).
-!- FUNCTION: Sodium-ion dependent, low affinity carnitine
transporter. Probably transports one sodium ion with one molecule
of carnitine. Also transports organic cations such as
tetraethylammonium (TEA) without the involvement of sodium.
Relative uptake activity ratio of carnitine to TEA is 1.78. A key
substrate of this transporter seems to be ergothioneine (ET).
{ECO:0000269|PubMed:10215651, ECO:0000269|PubMed:15795384}.
-!- BIOPHYSICOCHEMICAL PROPERTIES:
Kinetic parameters:
KM=0.470 mM for TEA (at 37 degrees Celsius and pH 7.4)
{ECO:0000269|PubMed:15459889, ECO:0000269|PubMed:9426230};
Vmax=0.974 nmol/min/mg enzyme toward TEA (at 37 degrees Celsius
and pH 7.4) {ECO:0000269|PubMed:15459889,
ECO:0000269|PubMed:9426230};
pH dependence:
More active at neutral and alkaline pHs than at acidic pHs.
{ECO:0000269|PubMed:15459889, ECO:0000269|PubMed:9426230};
-!- SUBUNIT: Interacts with PDZK1. {ECO:0000250}.
-!- SUBCELLULAR LOCATION: Membrane; Multi-pass membrane protein.
-!- TISSUE SPECIFICITY: Widely expressed. Highly expressed in whole
blood, bone marrow, trachea and fetal liver. Weakly expressed in
kidney, skeletal muscle, prostate, lung, pancreas, placenta,
heart, uterus, spleen and spinal cord. Highly expressed in
intestinal cell types affected by Crohn disease, including
epithelial cells. Expressed in CD68 macrophage and CD43 T-cells
but not in CD20 B-cells. Predominantly expressed in CD14 cells in
peripheral blood mononuclear cells. {ECO:0000269|PubMed:14608356,
ECO:0000269|PubMed:15107849, ECO:0000269|PubMed:9426230}.
-!- INDUCTION: Overexpressed upon TNF treatment.
{ECO:0000269|PubMed:14608356}.
-!- DISEASE: Rheumatoid arthritis (RA) [MIM:180300]: An inflammatory
disease with autoimmune features and a complex genetic component.
It primarily affects the joints and is characterized by
inflammatory changes in the synovial membranes and articular
structures, widespread fibrinoid degeneration of the collagen
fibers in mesenchymal tissues, and by atrophy and rarefaction of
bony structures. {ECO:0000269|PubMed:14608356}. Note=Disease
susceptibility is associated with variations affecting the gene
represented in this entry.
-!- SIMILARITY: Belongs to the major facilitator (TC 2.A.1)
superfamily. Organic cation transporter (TC 2.A.1.19) family.
{ECO:0000305}.
-!- CAUTION: PubMed:9426230 reported that this protein does not
transport carnitine, however, experiments were done with the Phe-
503 variant, which affects the ability to transport carnitine.
PubMed:15459889 showed that, although weakly, it can also
transport carnitine at some level. Its function in carnitine
transport is therefore unclear. {ECO:0000305}.
-----------------------------------------------------------------------
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EMBL; AB007448; BAA23356.1; -; mRNA.
EMBL; Y09881; CAA71007.1; -; mRNA.
EMBL; AC008599; -; NOT_ANNOTATED_CDS; Genomic_DNA.
EMBL; AC034220; -; NOT_ANNOTATED_CDS; Genomic_DNA.
EMBL; BC028313; AAH28313.1; -; mRNA.
CCDS; CCDS4153.1; -.
RefSeq; NP_003050.2; NM_003059.2.
UniGene; Hs.310591; -.
ProteinModelPortal; Q9H015; -.
BioGrid; 112470; 1.
STRING; 9606.ENSP00000200652; -.
ChEMBL; CHEMBL2073668; -.
DrugBank; DB00594; Amiloride.
DrugBank; DB00345; Aminohippuric acid.
DrugBank; DB01053; Benzylpenicillin.
DrugBank; DB00122; Choline.
DrugBank; DB00501; Cimetidine.
DrugBank; DB00575; Clonidine.
DrugBank; DB01151; Desipramine.
DrugBank; DB00536; Guanidine.
DrugBank; DB00458; Imipramine.
DrugBank; DB00332; Ipratropium bromide.
DrugBank; DB00125; L-Arginine.
DrugBank; DB00583; L-Carnitine.
DrugBank; DB00123; L-Lysine.
DrugBank; DB01137; Levofloxacin.
DrugBank; DB06691; Mepyramine.
DrugBank; DB00184; Nicotine.
DrugBank; DB01165; Ofloxacin.
DrugBank; DB01035; Procainamide.
DrugBank; DB01917; Putrescine.
DrugBank; DB00908; Quinidine.
DrugBank; DB00468; Quinine.
DrugBank; DB03566; Spermidine.
DrugBank; DB00127; Spermine.
DrugBank; DB00624; Testosterone.
DrugBank; DB08837; Tetraethylammonium.
DrugBank; DB01409; Tiotropium.
DrugBank; DB00661; Verapamil.
TCDB; 2.A.1.19.2; the major facilitator superfamily (mfs).
iPTMnet; Q9H015; -.
PhosphoSitePlus; Q9H015; -.
BioMuta; SLC22A4; -.
DMDM; 146345508; -.
EPD; Q9H015; -.
PaxDb; Q9H015; -.
PeptideAtlas; Q9H015; -.
PRIDE; Q9H015; -.
ProteomicsDB; 80197; -.
DNASU; 6583; -.
Ensembl; ENST00000200652; ENSP00000200652; ENSG00000197208.
GeneID; 6583; -.
KEGG; hsa:6583; -.
UCSC; uc003kwq.4; human.
CTD; 6583; -.
DisGeNET; 6583; -.
EuPathDB; HostDB:ENSG00000197208.5; -.
GeneCards; SLC22A4; -.
H-InvDB; HIX0024844; -.
HGNC; HGNC:10968; SLC22A4.
HPA; CAB015468; -.
HPA; HPA036974; -.
MalaCards; SLC22A4; -.
MIM; 180300; phenotype.
MIM; 604190; gene.
neXtProt; NX_Q9H015; -.
OpenTargets; ENSG00000197208; -.
PharmGKB; PA332; -.
eggNOG; ENOG410IRIE; Eukaryota.
eggNOG; ENOG410XSRI; LUCA.
GeneTree; ENSGT00760000118852; -.
HOGENOM; HOG000234570; -.
HOVERGEN; HBG061545; -.
InParanoid; Q9H015; -.
KO; K08202; -.
OMA; SWEMFTV; -.
OrthoDB; EOG091G05AC; -.
PhylomeDB; Q9H015; -.
TreeFam; TF315847; -.
Reactome; R-HSA-549127; Organic cation transport.
ChiTaRS; SLC22A4; human.
GeneWiki; SLC22A4; -.
GenomeRNAi; 6583; -.
PRO; PR:Q9H015; -.
Proteomes; UP000005640; Chromosome 5.
Bgee; ENSG00000197208; Expressed in 140 organ(s), highest expression level in bronchial epithelial cell.
CleanEx; HS_SLC22A4; -.
Genevisible; Q9H015; HS.
GO; GO:0016324; C:apical plasma membrane; IDA:BHF-UCL.
GO; GO:0005887; C:integral component of plasma membrane; TAS:ProtInc.
GO; GO:0005739; C:mitochondrion; IDA:MGI.
GO; GO:0005886; C:plasma membrane; TAS:Reactome.
GO; GO:0005524; F:ATP binding; IEA:UniProtKB-KW.
GO; GO:0015226; F:carnitine transmembrane transporter activity; IDA:MGI.
GO; GO:0015491; F:cation:cation antiporter activity; IDA:BHF-UCL.
GO; GO:0000166; F:nucleotide binding; TAS:ProtInc.
GO; GO:0008514; F:organic anion transmembrane transporter activity; IBA:GO_Central.
GO; GO:0030165; F:PDZ domain binding; IPI:BHF-UCL.
GO; GO:0015651; F:quaternary ammonium group transmembrane transporter activity; IDA:BHF-UCL.
GO; GO:0008513; F:secondary active organic cation transmembrane transporter activity; TAS:Reactome.
GO; GO:0015293; F:symporter activity; IEA:UniProtKB-KW.
GO; GO:0007589; P:body fluid secretion; TAS:ProtInc.
GO; GO:0009437; P:carnitine metabolic process; IBA:GO_Central.
GO; GO:0015879; P:carnitine transport; IDA:MGI.
GO; GO:0015695; P:organic cation transport; TAS:ProtInc.
GO; GO:0015697; P:quaternary ammonium group transport; IDA:BHF-UCL.
GO; GO:0006814; P:sodium ion transport; IEA:UniProtKB-KW.
GO; GO:0006641; P:triglyceride metabolic process; IEA:Ensembl.
CDD; cd06174; MFS; 1.
InterPro; IPR020846; MFS_dom.
InterPro; IPR005828; MFS_sugar_transport-like.
InterPro; IPR036259; MFS_trans_sf.
InterPro; IPR004749; Orgcat_transp/SVOP.
InterPro; IPR005829; Sugar_transporter_CS.
Pfam; PF00083; Sugar_tr; 1.
SUPFAM; SSF103473; SSF103473; 1.
TIGRFAMs; TIGR00898; 2A0119; 1.
PROSITE; PS50850; MFS; 1.
PROSITE; PS00216; SUGAR_TRANSPORT_1; 1.
1: Evidence at protein level;
ATP-binding; Complete proteome; Disease mutation; Glycoprotein;
Ion transport; Membrane; Nucleotide-binding; Polymorphism;
Reference proteome; Sodium; Sodium transport; Symport; Transmembrane;
Transmembrane helix; Transport.
CHAIN 1 551 Solute carrier family 22 member 4.
/FTId=PRO_0000220497.
TOPO_DOM 1 20 Cytoplasmic. {ECO:0000255}.
TRANSMEM 21 41 Helical; Name=1. {ECO:0000255}.
TOPO_DOM 42 141 Extracellular. {ECO:0000255}.
TRANSMEM 142 162 Helical; Name=2. {ECO:0000255}.
TOPO_DOM 163 171 Cytoplasmic. {ECO:0000255}.
TRANSMEM 172 192 Helical; Name=3. {ECO:0000255}.
TOPO_DOM 193 197 Extracellular. {ECO:0000255}.
TRANSMEM 198 218 Helical; Name=4. {ECO:0000255}.
TOPO_DOM 219 232 Cytoplasmic. {ECO:0000255}.
TRANSMEM 233 253 Helical; Name=5. {ECO:0000255}.
TOPO_DOM 254 257 Extracellular. {ECO:0000255}.
TRANSMEM 258 278 Helical; Name=6. {ECO:0000255}.
TOPO_DOM 279 337 Cytoplasmic. {ECO:0000255}.
TRANSMEM 338 358 Helical; Name=7. {ECO:0000255}.
TOPO_DOM 359 371 Extracellular. {ECO:0000255}.
TRANSMEM 372 392 Helical; Name=8. {ECO:0000255}.
TOPO_DOM 393 399 Cytoplasmic. {ECO:0000255}.
TRANSMEM 400 420 Helical; Name=9. {ECO:0000255}.
TOPO_DOM 421 426 Extracellular. {ECO:0000255}.
TRANSMEM 427 447 Helical; Name=10. {ECO:0000255}.
TOPO_DOM 448 460 Cytoplasmic. {ECO:0000255}.
TRANSMEM 461 481 Helical; Name=11. {ECO:0000255}.
TOPO_DOM 482 486 Extracellular. {ECO:0000255}.
TRANSMEM 487 507 Helical; Name=12. {ECO:0000255}.
TOPO_DOM 508 551 Cytoplasmic. {ECO:0000255}.
NP_BIND 218 225 ATP. {ECO:0000255}.
CARBOHYD 57 57 N-linked (GlcNAc...) asparagine.
{ECO:0000255}.
CARBOHYD 64 64 N-linked (GlcNAc...) asparagine.
{ECO:0000255}.
CARBOHYD 91 91 N-linked (GlcNAc...) asparagine.
{ECO:0000255}.
VARIANT 306 306 I -> T (in dbSNP:rs272893).
{ECO:0000269|PubMed:12436193,
ECO:0000269|PubMed:9426230}.
/FTId=VAR_019528.
VARIANT 462 462 G -> E (abrogates TEA transport activity;
dbSNP:rs4646201).
{ECO:0000269|PubMed:12436193,
ECO:0000269|PubMed:15459889}.
/FTId=VAR_019529.
VARIANT 503 503 L -> F (reduces the ability to transport
carnitine; dbSNP:rs1050152).
{ECO:0000269|PubMed:15107849,
ECO:0000269|PubMed:9426230}.
/FTId=VAR_019530.
SEQUENCE 551 AA; 62155 MW; C827A99AA78C9443 CRC64;
MRDYDEVIAF LGEWGPFQRL IFFLLSASII PNGFNGMSVV FLAGTPEHRC RVPDAANLSS
AWRNNSVPLR LRDGREVPHS CSRYRLATIA NFSALGLEPG RDVDLGQLEQ ESCLDGWEFS
QDVYLSTVVT EWNLVCEDNW KVPLTTSLFF VGVLLGSFVS GQLSDRFGRK NVLFATMAVQ
TGFSFLQIFS ISWEMFTVLF VIVGMGQISN YVVAFILGTE ILGKSVRIIF STLGVCTFFA
VGYMLLPLFA YFIRDWRMLL LALTVPGVLC VPLWWFIPES PRWLISQRRF REAEDIIQKA
AKMNNIAVPA VIFDSVEELN PLKQQKAFIL DLFRTRNIAI MTIMSLLLWM LTSVGYFALS
LDAPNLHGDA YLNCFLSALI EIPAYITAWL LLRTLPRRYI IAAVLFWGGG VLLFIQLVPV
DYYFLSIGLV MLGKFGITSA FSMLYVFTAE LYPTLVRNMA VGVTSTASRV GSIIAPYFVY
LGAYNRMLPY IVMGSLTVLI GILTLFFPES LGMTLPETLE QMQKVKWFRS GKKTRDSMET
EENPKVLITA F


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