{"results":[{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"O43914","secondaryAccessions":["A8K2X0","F5H389","Q6FGA5","Q9UMT3"],"uniProtkbId":"TYOBP_HUMAN","entryAudit":{"firstPublicDate":"2000-05-30","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"1998-06-01","entryVersion":196,"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":{"evidences":[{"evidenceCode":"ECO:0000312","source":"HGNC","id":"HGNC:12449"}],"value":"TYRO protein tyrosine kinase-binding protein"}},"alternativeNames":[{"fullName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"9490415"}],"value":"DNAX-activation protein 12"}},{"fullName":{"value":"Killer-activating receptor-associated protein"},"shortNames":[{"value":"KAR-associated protein"}]}],"flag":"Precursor"},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000312","source":"HGNC","id":"HGNC:12449"}],"value":"TYROBP"},"synonyms":[{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"9490415"}],"value":"DAP12"},{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"O54885"}],"value":"KARAP"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"O54885"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10449773"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10604985"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10799849"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11602640"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15557162"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16920917"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17928527"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18957693"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21727189"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23715743"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25957402"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26221034"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9490415"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9655483"}],"value":"Adapter protein which non-covalently associates with activating receptors found on the surface of a variety of immune cells to mediate signaling and cell activation following ligand binding by the receptors (PubMed:10604985, PubMed:9490415, PubMed:9655483). TYROBP is tyrosine-phosphorylated in the ITAM domain following ligand binding by the associated receptors which leads to activation of additional tyrosine kinases and subsequent cell activation (PubMed:9490415). Also has an inhibitory role in some cells (PubMed:21727189). Non-covalently associates with activating receptors of the CD300 family to mediate cell activation (PubMed:15557162, PubMed:16920917, PubMed:17928527, PubMed:26221034). Also mediates cell activation through association with activating receptors of the CD200R family (By similarity). Required for neutrophil activation mediated by integrin (By similarity). Required for the activation of myeloid cells mediated by the CLEC5A/MDL1 receptor (PubMed:10449773). Associates with natural killer (NK) cell receptors such as KIR2DS2 and the KLRD1/KLRC2 heterodimer to mediate NK cell activation (PubMed:23715743, PubMed:9490415, PubMed:9655483). Also enhances trafficking and cell surface expression of NK cell receptors KIR2DS1, KIR2DS2 and KIR2DS4 and ensures their stability at the cell surface (PubMed:23715743). Associates with SIRPB1 to mediate activation of myeloid cells such as monocytes and dendritic cells (PubMed:10604985). Associates with TREM1 to mediate activation of neutrophils and monocytes (PubMed:10799849). Associates with TREM2 on monocyte-derived dendritic cells to mediate up-regulation of chemokine receptor CCR7 and dendritic cell maturation and survival (PubMed:11602640). Association with TREM2 mediates cytokine-induced formation of multinucleated giant cells which are formed by the fusion of macrophages (PubMed:18957693). Stabilizes the TREM2 C-terminal fragment (TREM2-CTF) produced by TREM2 ectodomain shedding which suppresses the release of pro-inflammatory cytokines (PubMed:25957402). In microglia, required with TREM2 for phagocytosis of apoptotic neurons (By similarity). Required with ITGAM/CD11B in microglia to control production of microglial superoxide ions which promote the neuronal apoptosis that occurs during brain development (By similarity). Promotes pro-inflammatory responses in microglia following nerve injury which accelerates degeneration of injured neurons (By similarity). Positively regulates the expression of the IRAK3/IRAK-M kinase and IL10 production by liver dendritic cells and inhibits their T cell allostimulatory ability (By similarity). Negatively regulates B cell proliferation (PubMed:21727189). Required for CSF1-mediated osteoclast cytoskeletal organization (By similarity). Positively regulates multinucleation during osteoclast development (By similarity)"}],"commentType":"FUNCTION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"O54885"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10449773"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10604985"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10799849"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11602640"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15557162"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16920917"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17012248"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17928527"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18624290"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20890284"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23715743"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25957402"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25981043"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26221034"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26358190"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9655483"}],"value":"Homodimer; disulfide-linked (PubMed:20890284). Homotrimer; disulfide-linked (PubMed:25981043). Homotetramer; disulfide-linked (PubMed:25981043). Homotrimers and homotetramers form when low levels of partner receptors are available and are competitive with assembly with interacting receptors (PubMed:25981043). They may represent alternative oligomerization states or may be intermediates in the receptor assembly process (PubMed:25981043). Binding of a metal cation aids in homooligomerization through coordination of the metal ion by the subunits of the oligomer (PubMed:25981043). Interacts with TREM1 (PubMed:10799849). Interacts with TREM2 (PubMed:11602640, PubMed:25957402). Interacts with SIRPB1 (PubMed:10604985). Interacts with CLECSF5 (PubMed:10449773). Interacts with SIGLEC14 (PubMed:17012248). Interacts with CD300LB and CD300E (PubMed:15557162, PubMed:16920917, PubMed:17928527). Interacts with CD300C2 (By similarity). Interacts (via ITAM domain) with SYK (via SH2 domains); activates SYK mediating neutrophil and macrophage integrin-mediated activation (By similarity). Interacts with KLRC2, KIR2DS3 and KIR2DS5 (PubMed:18624290, PubMed:20890284). Interacts with CD300H (PubMed:26221034). Interacts with KIR2DS1 (PubMed:23715743). Interacts with KLRD1 (PubMed:15940674, PubMed:9655483). Interacts with SIGLEC1 (PubMed:26358190)"}],"commentType":"SUBUNIT"},{"commentType":"INTERACTION","interactions":[{"interactantOne":{"uniProtKBAccession":"O43914","intActId":"EBI-2214794"},"interactantTwo":{"uniProtKBAccession":"Q14953","geneName":"KIR2DS5","intActId":"EBI-16823921"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"O43914","intActId":"EBI-2214794"},"interactantTwo":{"uniProtKBAccession":"O95944","geneName":"NCR2","intActId":"EBI-14058375"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"O43914","intActId":"EBI-2214794"},"interactantTwo":{"uniProtKBAccession":"O00241","geneName":"SIRPB1","intActId":"EBI-2615458"},"numberOfExperiments":4,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"O43914","intActId":"EBI-2214794"},"interactantTwo":{"uniProtKBAccession":"Q969S0","geneName":"SLC35B4","intActId":"EBI-10281213"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"O43914","intActId":"EBI-2214794"},"interactantTwo":{"uniProtKBAccession":"Q8N2M4","geneName":"TMEM86A","intActId":"EBI-12015604"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"O43914","intActId":"EBI-2214794"},"interactantTwo":{"uniProtKBAccession":"Q9NZC2","geneName":"TREM2","intActId":"EBI-14036387"},"numberOfExperiments":4,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"O43914","intActId":"EBI-2214794"},"interactantTwo":{"uniProtKBAccession":"O43914","geneName":"TYROBP","intActId":"EBI-2214794"},"numberOfExperiments":2,"organismDiffer":false}]},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9655483"}],"value":"Cell membrane","id":"SL-0039"},"topology":{"evidences":[{"evidenceCode":"ECO:0000255"}],"value":"Single-pass type I membrane protein","id":"SL-9905"}}]},{"commentType":"ALTERNATIVE PRODUCTS","events":["Alternative splicing"],"isoforms":[{"name":{"value":"1"},"synonyms":[{"value":"KARAP-a"}],"isoformIds":["O43914-1"],"isoformSequenceStatus":"Displayed"},{"name":{"value":"2"},"synonyms":[{"value":"KARAP-b"}],"isoformIds":["O43914-2"],"sequenceIds":["VSP_012909"],"isoformSequenceStatus":"Described"},{"name":{"value":"3"},"isoformIds":["O43914-3"],"sequenceIds":["VSP_046066"],"isoformSequenceStatus":"Described"}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11922939"}],"value":"Expressed at low levels in the early development of the hematopoietic system and in the promonocytic stage and at high levels in mature monocytes. Expressed in hematological cells and tissues such as peripheral blood leukocytes and spleen. Also found in bone marrow, lymph nodes, placenta, lung and liver. Expressed at lower levels in different parts of the brain especially in the basal ganglia and corpus callosum"}],"commentType":"TISSUE SPECIFICITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9490415"}],"value":"Following ligand binding by associated receptors, tyrosine phosphorylated in the ITAM domain which leads to activation of additional tyrosine kinases and subsequent cell activation"}],"commentType":"PTM"},{"commentType":"DISEASE","disease":{"diseaseId":"Polycystic lipomembranous osteodysplasia with sclerosing leukoencephalopathy 1","diseaseAccession":"DI-02174","acronym":"PLOSL1","description":"A recessively inherited disease characterized by presenile dementia along with large-scale destruction of cancellous bones. Initial symptoms, starting in the twenties, are pain and swelling resulting from cysts in the wrists and ankles. Extremity bone fractures could occur with minor trauma. At around 30 years of age, patients gradually develop neuropsychiatric symptoms, including epileptic seizures, agnosia, apraxia, speech disorder, memory disturbance, euphoria, and loss of social inhibitions. The disorder usually leads to death in the fifth decade of life.","diseaseCrossReference":{"database":"MIM","id":"221770"},"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10888890"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12370476"}]},"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 TYROBP family"}],"commentType":"SIMILARITY"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":21,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000255"}]},{"type":"Chain","location":{"start":{"value":22,"modifier":"EXACT"},"end":{"value":113,"modifier":"EXACT"}},"description":"TYRO protein tyrosine kinase-binding protein","featureId":"PRO_0000022603"},{"type":"Topological domain","location":{"start":{"value":22,"modifier":"EXACT"},"end":{"value":40,"modifier":"EXACT"}},"description":"Extracellular","evidences":[{"evidenceCode":"ECO:0000305","source":"PubMed","id":"25981043"}]},{"type":"Transmembrane","location":{"start":{"value":41,"modifier":"EXACT"},"end":{"value":61,"modifier":"EXACT"}},"description":"Helical","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25981043"}]},{"type":"Topological domain","location":{"start":{"value":62,"modifier":"EXACT"},"end":{"value":113,"modifier":"EXACT"}},"description":"Cytoplasmic","evidences":[{"evidenceCode":"ECO:0000305","source":"PubMed","id":"25981043"}]},{"type":"Domain","location":{"start":{"value":80,"modifier":"EXACT"},"end":{"value":108,"modifier":"EXACT"}},"description":"ITAM"},{"type":"Region","location":{"start":{"value":75,"modifier":"EXACT"},"end":{"value":113,"modifier":"EXACT"}},"description":"Disordered","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"MobiDB-lite"}]},{"type":"Compositional bias","location":{"start":{"value":87,"modifier":"EXACT"},"end":{"value":113,"modifier":"EXACT"}},"description":"Polar residues","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"MobiDB-lite"}]},{"type":"Binding site","location":{"start":{"value":50,"modifier":"EXACT"},"end":{"value":50,"modifier":"EXACT"}},"description":"","featureCrossReferences":[{"database":"ChEBI","id":"CHEBI:29108"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25981043"}],"ligand":{"name":"Ca(2+)","id":"ChEBI:CHEBI:29108","note":"ligand shared between two neighboring subunits in homooligomer"}},{"type":"Site","location":{"start":{"value":54,"modifier":"EXACT"},"end":{"value":54,"modifier":"EXACT"}},"description":"Important for interaction with transmembrane receptors","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20890284"}]},{"type":"Modified residue","location":{"start":{"value":91,"modifier":"EXACT"},"end":{"value":91,"modifier":"EXACT"}},"description":"Phosphotyrosine","evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"O54885"}]},{"type":"Modified residue","location":{"start":{"value":102,"modifier":"EXACT"},"end":{"value":102,"modifier":"EXACT"}},"description":"Phosphotyrosine","evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"O54885"}]},{"type":"Disulfide bond","location":{"start":{"value":35,"modifier":"EXACT"},"end":{"value":35,"modifier":"EXACT"}},"description":"Interchain","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20890284"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25981043"},{"evidenceCode":"ECO:0007744","source":"PDB","id":"2L34"},{"evidenceCode":"ECO:0007744","source":"PDB","id":"2L35"},{"evidenceCode":"ECO:0007744","source":"PDB","id":"4WOL"}]},{"type":"Alternative sequence","location":{"start":{"value":20,"modifier":"EXACT"},"end":{"value":30,"modifier":"EXACT"}},"description":"in isoform 3","evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"16344560"}],"featureId":"VSP_046066","alternativeSequence":{}},{"type":"Alternative sequence","location":{"start":{"value":77,"modifier":"EXACT"},"end":{"value":77,"modifier":"EXACT"}},"description":"in isoform 2","evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"14702039"},{"evidenceCode":"ECO:0000303","source":"Reference","id":"Ref.2"},{"evidenceCode":"ECO:0000303","source":"Reference","id":"Ref.3"}],"featureId":"VSP_012909","alternativeSequence":{}},{"type":"Natural variant","location":{"start":{"value":2,"modifier":"EXACT"},"end":{"value":2,"modifier":"EXACT"}},"description":"found in patients with early-onset Alzheimer disease; uncertain significance; associated in cis with L-55 in some patients; dbSNP:rs200649978","featureCrossReferences":[{"database":"dbSNP","id":"rs200649978"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27658901"}],"featureId":"VAR_081398","alternativeSequence":{"originalSequence":"G","alternativeSequences":["E"]}},{"type":"Natural variant","location":{"start":{"value":23,"modifier":"EXACT"},"end":{"value":23,"modifier":"EXACT"}},"description":"found in patients with early-onset Alzheimer disease; uncertain significance; dbSNP:rs769635655","featureCrossReferences":[{"database":"dbSNP","id":"rs769635655"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27658901"}],"featureId":"VAR_081399","alternativeSequence":{"originalSequence":"R","alternativeSequences":["C"]}},{"type":"Natural variant","location":{"start":{"value":47,"modifier":"EXACT"},"end":{"value":47,"modifier":"EXACT"}},"description":"found in patients with early-onset Alzheimer disease; uncertain significance; dbSNP:rs372140827","featureCrossReferences":[{"database":"dbSNP","id":"rs372140827"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27658901"}],"featureId":"VAR_081400","alternativeSequence":{"originalSequence":"V","alternativeSequences":["A"]}},{"type":"Natural variant","location":{"start":{"value":50,"modifier":"EXACT"},"end":{"value":50,"modifier":"EXACT"}},"description":"found in patients with early-onset Alzheimer disease; uncertain significance; causes reduced expression; also leads to reduced expression of TREM2","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27658901"}],"featureId":"VAR_081401","alternativeSequence":{"originalSequence":"D","alternativeSequences":["DPADGRLVLGDRDGR"]}},{"type":"Natural variant","location":{"start":{"value":55,"modifier":"EXACT"},"end":{"value":55,"modifier":"EXACT"}},"description":"found in patients with early-onset Alzheimer disease; uncertain significance; associated in cis with E-2 in some patients; 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when associated with L-41 and L-49.","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25981043"}],"alternativeSequence":{"originalSequence":"G","alternativeSequences":["L"]}},{"type":"Mutagenesis","location":{"start":{"value":49,"modifier":"EXACT"},"end":{"value":49,"modifier":"EXACT"}},"description":"Does not significantly alter the formation of homotrimers or homotetramers; when associated with L-41 and L-45.","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25981043"}],"alternativeSequence":{"originalSequence":"G","alternativeSequences":["L"]}},{"type":"Mutagenesis","location":{"start":{"value":50,"modifier":"EXACT"},"end":{"value":50,"modifier":"EXACT"}},"description":"Reduced cell surface expression of KIR2DS1. Severely impairs formation of homotrimers and homotetramers. Abolishes interaction with TREM2 and stabilization of TREM2-CTF. Impairs the expression of KLRD1-KLRC2 on the cell surface.","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23715743"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25957402"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25981043"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9655483"}],"alternativeSequence":{"originalSequence":"D","alternativeSequences":["A"]}},{"type":"Mutagenesis","location":{"start":{"value":50,"modifier":"EXACT"},"end":{"value":50,"modifier":"EXACT"}},"description":"Severely impairs formation of homotrimers and homotetramers.","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25981043"}],"alternativeSequence":{"originalSequence":"D","alternativeSequences":["E","Q"]}},{"type":"Mutagenesis","location":{"start":{"value":50,"modifier":"EXACT"},"end":{"value":50,"modifier":"EXACT"}},"description":"Reduces formation of homotrimers and homotetramers.","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25981043"}],"alternativeSequence":{"originalSequence":"D","alternativeSequences":["N"]}},{"type":"Mutagenesis","location":{"start":{"value":54,"modifier":"EXACT"},"end":{"value":54,"modifier":"EXACT"}},"description":"Reduced interaction with KLRC2 and KIR2DS3. 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