{"results":[{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"P31645","secondaryAccessions":["Q5EE02"],"uniProtkbId":"SC6A4_HUMAN","entryAudit":{"firstPublicDate":"1993-07-01","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"1993-07-01","entryVersion":227,"sequenceVersion":1},"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":{"value":"Sodium-dependent serotonin transporter"},"shortNames":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P31652"}],"value":"SERT"}]},"alternativeNames":[{"fullName":{"value":"5HT transporter"},"shortNames":[{"value":"5HTT"}]},{"fullName":{"value":"Solute carrier family 6 member 4"}}]},"genes":[{"geneName":{"value":"SLC6A4"},"synonyms":[{"value":"HTT"},{"value":"SERT"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P31652"},{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q60857"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10407194"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12869649"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17506858"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18317590"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21730057"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27049939"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27756841"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"34851672"}],"value":"Serotonin transporter that cotransports serotonin with one Na(+) ion in exchange for one K(+) ion and possibly one proton in an overall electroneutral transport cycle. Transports serotonin across the plasma membrane from the extracellular compartment to the cytosol thus limiting serotonin intercellular signaling (PubMed:10407194, PubMed:12869649, PubMed:21730057, PubMed:27049939, PubMed:27756841, PubMed:34851672). Essential for serotonin homeostasis in the central nervous system. In the developing somatosensory cortex, acts in glutamatergic neurons to control serotonin uptake and its trophic functions accounting for proper spatial organization of cortical neurons and elaboration of sensory circuits. In the mature cortex, acts primarily in brainstem raphe neurons to mediate serotonin uptake from the synaptic cleft back into the pre-synaptic terminal thus terminating serotonin signaling at the synapse (By similarity). Modulates mucosal serotonin levels in the gastrointestinal tract through uptake and clearance of serotonin in enterocytes. Required for enteric neurogenesis and gastrointestinal reflexes (By similarity). Regulates blood serotonin levels by ensuring rapid high affinity uptake of serotonin from plasma to platelets, where it is further stored in dense granules via vesicular monoamine transporters and then released upon stimulation (PubMed:17506858, PubMed:18317590). Mechanistically, the transport cycle starts with an outward-open conformation having Na1(+) and Cl(-) sites occupied. The binding of a second extracellular Na2(+) ion and serotonin substrate leads to structural changes to outward-occluded to inward-occluded to inward-open, where the Na2(+) ion and serotonin are released into the cytosol. Binding of intracellular K(+) ion induces conformational transitions to inward-occluded to outward-open and completes the cycle by releasing K(+) possibly together with a proton bound to Asp-98 into the extracellular compartment. Na1(+) and Cl(-) ions remain bound throughout the transport cycle (PubMed:10407194, PubMed:12869649, PubMed:21730057, PubMed:27049939, PubMed:27756841, PubMed:34851672). Additionally, displays serotonin-induced channel-like conductance for monovalent cations, mainly Na(+) ions. The channel activity is uncoupled from the transport cycle and may contribute to the membrane resting potential or excitability (By similarity)"}],"commentType":"FUNCTION"},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"serotonin(out) + K(+)(in) + Na(+)(out) + H(+)(in) = serotonin(in) + K(+)(out) + Na(+)(in) + H(+)(out)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:75839"},{"database":"ChEBI","id":"CHEBI:15378"},{"database":"ChEBI","id":"CHEBI:29101"},{"database":"ChEBI","id":"CHEBI:29103"},{"database":"ChEBI","id":"CHEBI:350546"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10407194"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12869649"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21730057"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27049939"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27756841"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"34851672"}]},"physiologicalReactions":[{"directionType":"left-to-right","reactionCrossReference":{"database":"Rhea","id":"RHEA:75840"},"evidences":[{"evidenceCode":"ECO:0000305","source":"PubMed","id":"10407194"},{"evidenceCode":"ECO:0000305","source":"PubMed","id":"12869649"},{"evidenceCode":"ECO:0000305","source":"PubMed","id":"21730057"},{"evidenceCode":"ECO:0000305","source":"PubMed","id":"27049939"},{"evidenceCode":"ECO:0000305","source":"PubMed","id":"27756841"},{"evidenceCode":"ECO:0000305","source":"PubMed","id":"34851672"}]}]},{"commentType":"BIOPHYSICOCHEMICAL PROPERTIES","kineticParameters":{"maximumVelocities":[{"velocity":1.73,"unit":"pmol/min/mg","enzyme":"enzyme for serotonin","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12869649"}]},{"velocity":1.58,"unit":"pmol/min/mg","enzyme":"enzyme for serotonin","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17506858"}]},{"velocity":15.0,"unit":"pmol/min/mg","enzyme":"enzyme for serotonin","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10407194"}]}],"michaelisConstants":[{"constant":1.28,"unit":"uM","substrate":"serotonin","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12869649"}]},{"constant":0.98,"unit":"uM","substrate":"serotonin","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17506858"}]},{"constant":0.28,"unit":"uM","substrate":"serotonin","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10407194"}]},{"constant":1.9,"unit":"uM","substrate":"serotonin","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27049939"}]},{"constant":0.9,"unit":"uM","substrate":"serotonin","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21730057"}]}]}},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P31652"},{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q60857"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16803896"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16870614"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18317590"}],"value":"Monomer or homooligomer (By similarity). Interacts with TGFB1I1 (PubMed:16803896). Interacts with SEC23A, SEC24C and PATJ. Interacts with NOS1; the interaction may diminish the cell surface localization of SERT in the brain and, correspondingly, reduce serotonin reuptake. Interacts with filamentous actin and STX1A (By similarity). Interacts (via the N-terminus) with STX1A (via the H3 domain); this interaction regulates SLC4A6 channel conductance (By similarity). Interacts (via C-terminus) with SCAMP2; the interaction is direct and retains transporter molecules intracellularly (PubMed:16870614). Interacts with ITGAV:ITGB3 (PubMed:18317590). Interacts (via C-terminus) with ITGB3; this interaction regulates SLC6A4 trafficking (PubMed:18317590)"}],"commentType":"SUBUNIT"},{"commentType":"SUBCELLULAR LOCATION","note":{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P31652"},{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q60857"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16870614"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17506858"}],"value":"Could be part of recycling endosomes (PubMed:16870614). Density of transporter molecules on the plasma membrane is itself regulated by STX1A (By similarity). Density of transporter molecules on the plasma membrane is also regulated by serotonin (PubMed:17506858). Density of transporter molecules seems to be modulated by ITGAV:ITGB3 (By similarity)"}]},"subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12869649"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16870614"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17506858"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27049939"}],"value":"Cell membrane","id":"SL-0039"},"topology":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27049939"}],"value":"Multi-pass membrane protein","id":"SL-9909"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16870614"}],"value":"Endomembrane system","id":"SL-0147"},"topology":{"evidences":[{"evidenceCode":"ECO:0000255"}],"value":"Multi-pass membrane protein","id":"SL-9909"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16870614"}],"value":"Endosome membrane","id":"SL-0100"},"topology":{"evidences":[{"evidenceCode":"ECO:0000255"}],"value":"Multi-pass membrane protein","id":"SL-9909"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q60857"}],"value":"Synapse","id":"SL-0258"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q60857"}],"value":"Cell junction, focal adhesion","id":"SL-0118"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q60857"}],"value":"Cell projection, neuron projection","id":"SL-0518"}}]},{"commentType":"ALTERNATIVE PRODUCTS","events":["Alternative splicing"],"isoforms":[{"name":{"value":"1"},"isoformIds":["P31645-1"],"isoformSequenceStatus":"Displayed"},{"name":{"value":"2"},"isoformIds":["P31645-2"],"sequenceIds":["VSP_046553"],"isoformSequenceStatus":"Described"}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17506858"}],"value":"Expressed in platelets (at protein level)"}],"commentType":"TISSUE SPECIFICITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17913921"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21992875"}],"value":"Phosphorylation at Thr-276 increases 5-HT uptake and is required for cGMP-mediated SERT regulation"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12869766"}],"value":"A polymorphism in the promoter region (5-HTT gene-linked polymorphic region, 5-HTTLPR) is located approximately 1 kb upstream of the transcription initiation site and is composed of 16 repeat elements. The polymorphism consists of a 44-bp insertion or deletion involving repeat elements 6 to 8. The short allele is associated with lower transcriptional efficiency of the promoter compared with the long allele. Over half of the Caucasian population has a short allele. Individuals with one or two copies of the short allele exhibit more depressive symptoms, diagnosable depression and suicidality in relation to stressful life events than individuals homozygous for the long allele"}],"commentType":"POLYMORPHISM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15108187"}],"value":"The 5-HTTLPR polymorphism may influence susceptibility to anxiety [MIM:607834]"}],"commentType":"POLYMORPHISM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12869649"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"14593431"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17913921"}],"value":"The polymorphism Val-425 seems to be linked to a susceptibility to obsessive-compulsive disorder (OCD) [MIM:164230]"}],"commentType":"POLYMORPHISM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15635638"}],"value":"Genetic variations in SLC6A4 determine the genetic susceptibility to alcoholism [MIM:103780]"}],"commentType":"POLYMORPHISM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15995945"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18957375"}],"value":"Polymorphisms that alter SLC6A4 expression or function may increase the susceptibility to autism"}],"commentType":"POLYMORPHISM"},{"texts":[{"value":"This protein is the target of psychomotor stimulants such as amphetamines or cocaine"}],"commentType":"MISCELLANEOUS"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Belongs to the sodium:neurotransmitter symporter (SNF) (TC 2.A.22) family. SLC6A4 subfamily"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19270731"},{"evidenceCode":"ECO:0000305","source":"PubMed","id":"30707744"}],"value":"Was reported to interact with VIM, however the paper was retracted as some results and conclusions are not reliable"}],"commentType":"CAUTION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18227069"},{"evidenceCode":"ECO:0000305","source":"PubMed","id":"31201246"}],"value":"Was reported to interact with RAB4 and to be induced by serotonin, however the paper was retracted as some results and conclusions are not reliable"}],"commentType":"CAUTION"},{"commentType":"WEB RESOURCE","resourceName":"Wikipedia","resourceUrl":"https://en.wikipedia.org/wiki/Serotonin_transporter","ftp":false,"note":"Serotonin transporter entry"},{"commentType":"WEB RESOURCE","resourceName":"Protein 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Recruits onto early endosomes in a Rab5- and HAP40-dependent fashion (PubMed:16476778)"}]},"subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15654337"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16476778"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"7647777"}],"value":"Cytoplasm","id":"SL-0086"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15654337"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16391387"}],"value":"Nucleus","id":"SL-0191"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16476778"}],"value":"Early endosome","id":"SL-0094"}}]},{"commentType":"SUBCELLULAR LOCATION","molecule":"Huntingtin, myristoylated N-terminal fragment","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"24459296"}],"value":"Cytoplasmic vesicle, autophagosome","id":"SL-0023"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16391387"}],"value":"Expressed in the brain cortex (at protein level). Widely expressed with the highest level of expression in the brain (nerve fibers, varicosities, and nerve endings). In the brain, the regions where it can be mainly found are the cerebellar cortex, the neocortex, the striatum, and the hippocampal formation"}],"commentType":"TISSUE SPECIFICITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19748341"}],"value":"The N-terminal Gln-rich and Pro-rich domain has great conformational flexibility and is likely to exist in a fluctuating equilibrium of alpha-helical, random coil, and extended conformations"}],"commentType":"DOMAIN"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10770929"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"29802276"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8696339"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9535906"}],"value":"Cleaved by caspases downstream of the polyglutamine stretch (PubMed:10770929, PubMed:29802276, PubMed:8696339, PubMed:9535906). The resulting N-terminal fragments are cytotoxic and provokes apoptosis (PubMed:10770929)"}],"commentType":"PTM","molecule":"Huntingtin"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17141218"}],"value":"Forms with expanded polyglutamine expansion are specifically ubiquitinated by SYVN1, which promotes their proteasomal degradation"}],"commentType":"PTM","molecule":"Huntingtin"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17611284"}],"value":"Phosphorylation at Ser-1179 and Ser-1199 by CDK5 in response to DNA damage in nuclei of neurons protects neurons against polyglutamine expansion as well as DNA damage mediated toxicity"}],"commentType":"PTM","molecule":"Huntingtin"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"24459296"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"29802276"}],"value":"Myristoylated at Gly-551, following proteolytic cleavage at Asp-550"}],"commentType":"PTM","molecule":"Huntingtin, myristoylated N-terminal fragment"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8696339"}],"value":"The poly-Gln region of HTT is highly polymorphic (10 to 35 repeats) in the normal population and is expanded to about 36-120 repeats in Huntington disease patients. The repeat length usually increases in successive generations, but also contracts on occasion. The adjacent poly-Pro region is also polymorphic and varies between 7-12 residues. Polyglutamine expansion leads to elevated susceptibility to apopain cleavage and likely result in accelerated neuronal apoptosis (PubMed:8696339)"}],"commentType":"POLYMORPHISM"},{"commentType":"DISEASE","disease":{"diseaseId":"Huntington disease","diseaseAccession":"DI-01754","acronym":"HD","description":"A neurodegenerative disorder characterized by involuntary movements (chorea), general motor impairment, psychiatric disorders and dementia. Onset of the disease occurs usually in the third or fourth decade of life. Onset and clinical course depend on the degree of poly-Gln repeat expansion, longer expansions resulting in earlier onset and more severe clinical manifestations. Neuropathology of Huntington disease displays a distinctive pattern with loss of neurons, especially in the caudate and putamen.","diseaseCrossReference":{"database":"MIM","id":"143100"},"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8458085"}]},"note":{"texts":[{"value":"The disease is caused by variants affecting the gene represented in this entry"}]}},{"commentType":"DISEASE","disease":{"diseaseId":"Lopes-Maciel-Rodan syndrome","diseaseAccession":"DI-04988","acronym":"LOMARS","description":"An autosomal recessive neurodevelopmental disorder characterized by developmental regression in infancy, delayed psychomotor development, severe intellectual disability, and cerebral and cerebellar atrophy. Additional features include swallowing problems, dystonia, bradykinesia, and continuous manual stereotypies without chorea. Some patients manifest seizures.","diseaseCrossReference":{"database":"MIM","id":"617435"},"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26740508"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27329733"}]},"note":{"texts":[{"value":"The disease is caused by variants affecting the gene represented in this entry"}]}},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Belongs to the huntingtin family"}],"commentType":"SIMILARITY"},{"commentType":"WEB RESOURCE","resourceName":"Wikipedia","resourceUrl":"https://en.wikipedia.org/wiki/Huntingtin","ftp":false,"note":"Huntingtin entry"}],"features":[{"type":"Chain","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":3142,"modifier":"EXACT"}},"description":"Huntingtin","featureId":"PRO_0000083942"},{"type":"Chain","location":{"start":{"value":551,"modifier":"EXACT"},"end":{"value":584,"modifier":"EXACT"}},"description":"Huntingtin, myristoylated N-terminal 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