{"results":[{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"P28799","secondaryAccessions":["D3DX55","P23781","P23782","P23783","P23784","Q53HQ8","Q53Y88","Q540U8","Q9BWE7","Q9H8S1","Q9UCH0"],"uniProtkbId":"GRN_HUMAN","entryAudit":{"firstPublicDate":"1992-12-01","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2005-10-11","entryVersion":246,"sequenceVersion":2},"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":"16862116"}],"value":"Progranulin"},"shortNames":[{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"16862116"}],"value":"PGRN"}]},"alternativeNames":[{"fullName":{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P28798"}],"value":"Acrogranin"}},{"fullName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"1618805"}],"value":"Epithelin precursor"}},{"fullName":{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P28798"}],"value":"Glycoprotein of 88 Kda"},"shortNames":[{"value":"GP88"},{"value":"Glycoprotein 88"}]},{"fullName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"1542665"}],"value":"Granulin precursor"}},{"fullName":{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P28798"}],"value":"PC cell-derived growth factor"},"shortNames":[{"evidences":[{"evidenceCode":"ECO:0000303","source":"Reference","id":"Ref.4"}],"value":"PCDGF"}]},{"fullName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"12526812"},{"evidenceCode":"ECO:0000303","source":"PubMed","id":"1618805"}],"value":"Proepithelin"},"shortNames":[{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"12526812"}],"value":"PEPI"}]}],"contains":[{"recommendedName":{"fullName":{"value":"Paragranulin"}}},{"recommendedName":{"fullName":{"value":"Granulin-1"}},"alternativeNames":[{"fullName":{"value":"Granulin G"}}]},{"recommendedName":{"fullName":{"value":"Granulin-2"}},"alternativeNames":[{"fullName":{"value":"Granulin F"}}]},{"recommendedName":{"fullName":{"value":"Granulin-3"}},"alternativeNames":[{"fullName":{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P23785"}],"value":"Epithelin-2"}},{"fullName":{"value":"Granulin B"}}]},{"recommendedName":{"fullName":{"value":"Granulin-4"}},"alternativeNames":[{"fullName":{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P23785"}],"value":"Epithelin-1"}},{"fullName":{"value":"Granulin A"}}]},{"recommendedName":{"fullName":{"value":"Granulin-5"}},"alternativeNames":[{"fullName":{"value":"Granulin C"}}]},{"recommendedName":{"fullName":{"value":"Granulin-6"}},"alternativeNames":[{"fullName":{"value":"Granulin D"}}]},{"recommendedName":{"fullName":{"value":"Granulin-7"}},"alternativeNames":[{"fullName":{"value":"Granulin E"}}]}],"flag":"Precursor"},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000312","source":"HGNC","id":"HGNC:4601"}],"value":"GRN"}}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P28798"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12526812"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18378771"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"28073925"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"28453791"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"28541286"}],"value":"Secreted protein that acts as a key regulator of lysosomal function and as a growth factor involved in inflammation, wound healing and cell proliferation (PubMed:12526812, PubMed:18378771, PubMed:28073925, PubMed:28453791, PubMed:28541286). Regulates protein trafficking to lysosomes, and also the activity of lysosomal enzymes (PubMed:28453791, PubMed:28541286). Also facilitates the acidification of lysosomes, causing degradation of mature CTSD by CTSB (PubMed:28073925). In addition, functions as a wound-related growth factor that acts directly on dermal fibroblasts and endothelial cells to promote division, migration and the formation of capillary-like tubule structures (By similarity). Also promotes epithelial cell proliferation by blocking TNF-mediated neutrophil activation preventing release of oxidants and proteases (PubMed:12526812). Moreover, modulates inflammation in neurons by preserving neurons survival, axonal outgrowth and neuronal integrity (PubMed:18378771)"}],"commentType":"FUNCTION"},{"texts":[{"value":"Promotes proliferation of the epithelial cell line A431 in culture"}],"commentType":"FUNCTION","molecule":"Granulin-4"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12526812"}],"value":"Inhibits epithelial cell proliferation and induces epithelial cells to secrete IL-8"}],"commentType":"FUNCTION","molecule":"Granulin-3"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"28453791"}],"value":"Stabilizes CTSD through interaction with CTSD leading to maintain its aspartic-type peptidase activity"}],"commentType":"FUNCTION","molecule":"Granulin-7"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12526812"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21092856"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23364791"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26370502"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27789271"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"28453791"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"28541286"}],"value":"Progranulin is secreted as a homodimer (PubMed:23364791). Interacts with SLPI; interaction protects progranulin from proteolysis (PubMed:12526812). Interacts (via region corresponding to granulin-7 peptide) with CTSD; stabilizes CTSD and increases its proteolytic activity (PubMed:28453791). Interacts (via region corresponding to granulin-7 peptide) with SORT1; this interaction mediates endocytosis and lysosome delivery of progranulin; interaction occurs at the neuronal cell surface in a stressed nervous system (PubMed:21092856). Interacts with PSAP; facilitates lysosomal delivery of progranulin from the extracellular space and the biosynthetic pathway (PubMed:26370502). Forms a complex with PSAP and M6PR; PSAP bridges the binding between progranulin and M6PR (PubMed:26370502). Forms a complex with PSAP and SORT1; progranulin bridges the interaction between PSAP and SORT1; facilitates lysosomal targeting of PSAP via SORT1; interaction enhances PSAP uptake in primary cortical neurons (PubMed:28541286). Interacts (via regions corresponding to granulin-2 and granulin-7 peptides) with GBA1; this interaction prevents aggregation of GBA1-SCARB2 complex via interaction with HSPA1A upon stress (PubMed:27789271). Interacts (via region corresponding to granulin-7 peptide) with HSPA1A; mediates recruitment of HSPA1A to GBA1 and prevents GBA1 aggregation in response to stress (PubMed:27789271)"}],"commentType":"SUBUNIT"},{"commentType":"INTERACTION","interactions":[{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q6UY14-3","geneName":"ADAMTSL4","intActId":"EBI-10173507"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q9UIJ7","geneName":"AK3","intActId":"EBI-3916527"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q9NYG5","geneName":"ANAPC11","intActId":"EBI-2130187"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q8N6T3","geneName":"ARFGAP1","intActId":"EBI-716933"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q8N6T3-3","geneName":"ARFGAP1","intActId":"EBI-10694449"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q9Y575-3","geneName":"ASB3","intActId":"EBI-14199987"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q96DX5-3","geneName":"ASB9","intActId":"EBI-25843552"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q6XD76","geneName":"ASCL4","intActId":"EBI-10254793"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q96FT7-4","geneName":"ASIC4","intActId":"EBI-9089489"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q8IXM2","geneName":"BACC1","intActId":"EBI-4280811"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"P46379-2","geneName":"BAG6","intActId":"EBI-10988864"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q16611","geneName":"BAK1","intActId":"EBI-519866"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q14457","geneName":"BECN1","intActId":"EBI-949378"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q96LC9","geneName":"BMF","intActId":"EBI-3919268"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q9GZL8","geneName":"BPESC1","intActId":"EBI-25861458"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q8WUW1","geneName":"BRK1","intActId":"EBI-2837444"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q9Y297","geneName":"BTRC","intActId":"EBI-307461"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q8TAB5","geneName":"C1orf216","intActId":"EBI-747505"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q6P5X5","geneName":"C22orf39","intActId":"EBI-7317823"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q6P5X5-2","geneName":"C22orf39","intActId":"EBI-10692329"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q53FE4","geneName":"C4orf17","intActId":"EBI-715110"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"O00555","geneName":"CACNA1A","intActId":"EBI-766279"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P28799","intActId":"EBI-747754"},"interactantTwo":{"uniProtKBAccession":"Q96NX5","geneName":"CAMK1G","intActId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