{"results":[{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"Q99808","secondaryAccessions":["B3KQV7","B3KQY5","Q5T9W9","Q9UJY2"],"uniProtkbId":"S29A1_HUMAN","entryAudit":{"firstPublicDate":"2000-12-01","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2007-01-23","entryVersion":214,"sequenceVersion":3},"annotationScore":5.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"lineage":["Eukaryota","Metazoa","Chordata","Craniata","Vertebrata","Euteleostomi","Mammalia","Eutheria","Euarchontoglires","Primates","Haplorrhini","Catarrhini","Hominidae","Homo"]},"proteinExistence":"1: Evidence at protein level","proteinDescription":{"recommendedName":{"fullName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"12527552"}],"value":"Equilibrative nucleoside transporter 1"},"shortNames":[{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"12527552"}],"value":"hENT1"}]},"alternativeNames":[{"fullName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"10755314"}],"value":"Equilibrative nitrobenzylmercaptopurine riboside-sensitive nucleoside transporter"},"shortNames":[{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"10755314"}],"value":"Equilibrative NBMPR-sensitive nucleoside transporter"},{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"10755314"}],"value":"es nucleoside transporter"}]},{"fullName":{"value":"Nucleoside transporter, es-type"}},{"fullName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"35790189"}],"value":"Solute carrier family 29 member 1"}}]},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000312","source":"HGNC","id":"HGNC:11003"}],"value":"SLC29A1"},"synonyms":[{"value":"ENT1"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"O54698"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10755314"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12527552"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"14759222"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15037197"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17379602"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21795683"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23639800"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26406980"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27995448"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"35790189"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"39737929"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"39885119"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8986748"}],"value":"Uniporter involved in the facilitative transport of nucleosides and nucleobases, and contributes to maintaining their cellular homeostasis (PubMed:10722669, PubMed:10755314, PubMed:12527552, PubMed:14759222, PubMed:15037197, PubMed:17379602, PubMed:21795683, PubMed:26406980, PubMed:27995448, PubMed:35790189, PubMed:8986748). Functions as a Na(+)-independent transporter (PubMed:8986748). Involved in the transport of nucleosides such as adenosine, guanosine, inosine, uridine, thymidine and cytidine (PubMed:10722669, PubMed:10755314, PubMed:12527552, PubMed:14759222, PubMed:15037197, PubMed:17379602, PubMed:26406980, PubMed:8986748, PubMed:39737929). Also transports purine nucleobases (hypoxanthine, adenine, guanine) and pyrimidine nucleobases (thymine, uracil) (PubMed:21795683, PubMed:27995448). Also mediates the uptake of nicotinamide; contributes to maintaining nicotinamide homeostasis but also regulated nicotinamide biological roles (PubMed:39885119). Mediates basolateral nucleoside uptake into Sertoli cells, thereby regulating the transport of nucleosides in testis across the blood-testis barrier (By similarity). Regulates inosine levels in brown adipocytes tissues (BAT) and extracellular inosine levels, which controls BAT-dependent energy expenditure (PubMed:35790189)"}],"commentType":"FUNCTION"},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"adenosine(in) = adenosine(out)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:75343"},{"database":"ChEBI","id":"CHEBI:16335"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12527552"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"14759222"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15037197"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17379602"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26406980"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27995448"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"39737929"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8986748"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"guanosine(in) = guanosine(out)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:75371"},{"database":"ChEBI","id":"CHEBI:16750"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"inosine(in) = inosine(out)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:75375"},{"database":"ChEBI","id":"CHEBI:17596"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12527552"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"14759222"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"35790189"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"uridine(out) = uridine(in)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:71519"},{"database":"ChEBI","id":"CHEBI:16704"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10755314"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15037197"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26406980"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8986748"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"thymidine(in) = thymidine(out)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:75363"},{"database":"ChEBI","id":"CHEBI:17748"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12527552"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"cytidine(in) = cytidine(out)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:75367"},{"database":"ChEBI","id":"CHEBI:17562"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15037197"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"adenine(out) = adenine(in)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:71523"},{"database":"ChEBI","id":"CHEBI:16708"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21795683"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"guanine(out) = guanine(in)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:71531"},{"database":"ChEBI","id":"CHEBI:16235"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21795683"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"thymine(out) = thymine(in)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:71527"},{"database":"ChEBI","id":"CHEBI:17821"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21795683"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"uracil(in) = uracil(out)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:69404"},{"database":"ChEBI","id":"CHEBI:17568"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21795683"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"hypoxanthine(out) = hypoxanthine(in)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:71515"},{"database":"ChEBI","id":"CHEBI:17368"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21795683"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"nicotinamide(out) = nicotinamide(in)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:86679"},{"database":"ChEBI","id":"CHEBI:17154"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"39885119"}]}},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"14759222"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15037197"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17379602"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21795683"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26406980"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27995448"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"39737929"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8986748"}],"value":"Transporter activity is sensitive to low concentrations of the inhibitor nitrobenzylmercaptopurine riboside (NBMPR) (PubMed:10722669, PubMed:14759222, PubMed:15037197, PubMed:17379602, PubMed:21795683, PubMed:26406980, PubMed:27995448, PubMed:8986748). Inhibited by dilazep (PubMed:14759222, PubMed:15037197, PubMed:17379602, PubMed:39737929). Inhibited by dipyridamole (PubMed:14759222, PubMed:15037197, PubMed:17379602, PubMed:39737929). Inhibited by hypoxanthine (PubMed:15037197). Inhibited by azidothymidine (AZT) (PubMed:15037197). Inhibited by dideoxycytidine (ddC) (PubMed:15037197). Inhibited by dideoxyinosine (ddI) (PubMed:15037197). Inhibited by draflazine (PubMed:17379602). Inhibited by soluflazine (PubMed:17379602). Inhibited by cladribine (PubMed:27995448). Inhibited by capecitabine (PubMed:27995448). Inhibited by clofarabine (PubMed:27995448). Inhibited by ribavirin (PubMed:27995448). Modestly inhibited by acyclovir (PubMed:27995448). Modestly inhibited by 5-fluorouracil (PubMed:27995448)"}],"commentType":"ACTIVITY REGULATION"},{"commentType":"BIOPHYSICOCHEMICAL PROPERTIES","kineticParameters":{"maximumVelocities":[{"velocity":351.0,"unit":"pmol/min/mg","enzyme":"enzyme for adenosine uptake","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"}]},{"velocity":1280.0,"unit":"pmol/min/mg","enzyme":"enzyme for adenosine uptake","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17379602"}]},{"velocity":578.0,"unit":"nmol/min/mg","enzyme":"enzyme for adenosine uptake","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27995448"}]},{"velocity":639.0,"unit":"pmol/min/mg","enzyme":"enzyme for guanosine uptake","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"}]},{"velocity":163.0,"unit":"pmol/min/mg","enzyme":"enzyme for inosine uptake","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"}]},{"velocity":853.0,"unit":"pmol/min/mg","enzyme":"enzyme for uridine uptake","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"}]},{"velocity":698.0,"unit":"pmol/min/mg","enzyme":"enzyme for thymidine uptake","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"}]},{"velocity":1290.0,"unit":"pmol/min/mg","enzyme":"enzyme for cytidine uptake","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"}]}],"michaelisConstants":[{"constant":40.0,"unit":"uM","substrate":"adenosine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"}]},{"constant":62.0,"unit":"uM","substrate":"adenosine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12527552"}]},{"constant":215.0,"unit":"uM","substrate":"adenosine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27995448"}]},{"constant":440.0,"unit":"uM","substrate":"adenosine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26406980"}]},{"constant":11.3,"unit":"uM","substrate":"adenosine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15037197"}]},{"constant":19.2,"unit":"uM","substrate":"adenosine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17379602"}]},{"constant":44.0,"unit":"uM","substrate":"adenosine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"14759222"}]},{"constant":140.0,"unit":"uM","substrate":"guanosine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"}]},{"constant":170.0,"unit":"uM","substrate":"inosine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"}]},{"constant":65.0,"unit":"uM","substrate":"inosine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"14759222"}]},{"constant":240.0,"unit":"uM","substrate":"uridine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8986748"}]},{"constant":260.0,"unit":"uM","substrate":"uridine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"}]},{"constant":400.0,"unit":"uM","substrate":"uridine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21795683"}]},{"constant":2160.0,"unit":"uM","substrate":"uridine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26406980"}]},{"constant":43.0,"unit":"uM","substrate":"uridine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15037197"}]},{"constant":300.0,"unit":"uM","substrate":"thymidine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"}]},{"constant":580.0,"unit":"uM","substrate":"cytidine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"}]},{"constant":207.0,"unit":"uM","substrate":"cytidine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15037197"}]},{"constant":3200.0,"unit":"uM","substrate":"adenine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21795683"}]},{"constant":6300.0,"unit":"uM","substrate":"thymine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21795683"}]},{"constant":6000.0,"unit":"uM","substrate":"hypoxanthine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21795683"}]}]}},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23219802"}],"value":"Identified in a complex with STOM"}],"commentType":"SUBUNIT"},{"commentType":"SUBCELLULAR LOCATION","note":{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11584005"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12527552"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23219802"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23639800"}],"value":"Localized to the basolateral membrane of Sertoli cells (PubMed:23639800). Localized to the cell membrane of erythrocytes (PubMed:11584005, PubMed:23219802)"}]},"subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12527552"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23639800"}],"value":"Basolateral cell membrane","id":"SL-0026"},"topology":{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Multi-pass membrane protein","id":"SL-9909"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12527552"}],"value":"Apical cell membrane","id":"SL-0015"},"topology":{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Multi-pass membrane protein","id":"SL-9909"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11584005"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15501974"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23219802"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26406980"}],"value":"Cell membrane","id":"SL-0039"},"topology":{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Multi-pass membrane protein","id":"SL-9909"}}]},{"commentType":"ALTERNATIVE PRODUCTS","events":["Alternative splicing"],"isoforms":[{"name":{"value":"1"},"isoformIds":["Q99808-1"],"isoformSequenceStatus":"Displayed"},{"name":{"value":"2"},"isoformIds":["Q99808-2"],"sequenceIds":["VSP_056579"],"isoformSequenceStatus":"Described"}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10755314"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11311901"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11584005"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12527552"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15501974"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23219802"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23639800"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"35790189"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8986748"}],"value":"Expressed in testis at the blood-testis barrier (at protein level) (PubMed:23639800). Detected in erythrocytes (at protein level) (PubMed:11584005, PubMed:23219802). Expressed at relatively high levels in cerebral cortex, particularly the frontal and parietal lobes, and the thalamus and basal ganglia (at protein level) (PubMed:11311901). In the midbrain expressed at moderate levels, whereas in the other areas of the brainstem, namely medulla and pons, cerebellum and the hippocampus expressed at lower amounts when compared to the other brain regions (at protein level) (PubMed:11311901). Expressed in Langerhans cells and lymphocytes in the pancreas (at protein level) (PubMed:15501974). Expressed in kidney, in polarized renal epithelial cells (PubMed:12527552). Expressed in adipose tissues (PubMed:35790189). Expressed in placenta (PubMed:8986748). Expressed in small intestine (PubMed:10755314)"}],"commentType":"TISSUE SPECIFICITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21795683"}],"value":"Cys-414 near TM10 is a major determinant of nucleobase transport activity"}],"commentType":"DOMAIN"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10722669"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19349973"}],"value":"Glycosylated"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15501974"}],"value":"The absence of the protein in tumor cells is associated with reduced survival in patients with gemcitabine-treated pancreas adenocarcinoma"}],"commentType":"MISCELLANEOUS"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Belongs to the SLC29A/ENT transporter (TC 2.A.57) family"}],"commentType":"SIMILARITY"}],"features":[{"type":"Initiator methionine","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":1,"modifier":"EXACT"}},"description":"Removed","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8986748"}]},{"type":"Chain","location":{"start":{"value":2,"modifier":"EXACT"},"end":{"value":456,"modifier":"EXACT"}},"description":"Equilibrative nucleoside transporter 1","featureId":"PRO_0000209337"},{"type":"Topological domain","location":{"start":{"value":2,"modifier":"EXACT"},"end":{"value":14,"modifier":"EXACT"}},"description":"Cytoplasmic","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11584005"}]},{"type":"Transmembrane","location":{"start":{"value":15,"modifier":"EXACT"},"end":{"value":36,"modifier":"EXACT"}},"description":"Helical; 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